N.P.O. Green, G.W. Stout, D.J. Taylor - Editor - Roland Soper - Biological Science 2. 2-Cambridge University Press (1985)

BIOLOGICAL “=
Me
WILBERFORCE
SCIENCE 2
systems, Maintenance and Change
N.P.O. GREEN B.Sc., M.I.Biol

Headmaster
Sutton Manor High School for Boys,
Sutton, Surrey
G.W. STOUT B.Sc., M.Ed., M.LBiol

Secretary for Biological Sciences
University of Cambridge Local Examinations Syndicate
D.j. TAYLOR B.Sc., Ph.D., M.I.Biol

Head of Biology
Strode's Sixth Form College, Egham
Editor
R. SOPER B.Sc., F.I.Biol
Formerly Vice-principal and Head of Science
Collyers Sixth Form College, Horsham
The right of the

Un iversity of Cambridge
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©Cambridge University Press 1984
First published 1984

Sixth printing 1987
Printed in Great Britain by Ebenezer Baylis & Son Ltd

The Trinity Press, Worcester and London
British Library cataloguing in publication data

Green, Nigel
Biological science 2: systems, maintenance
and change.
1. Biology
I. Title II. Stout, Wilf III. Taylor, Dennis
IV. Soper, R.
574 QH308.7
ISBN 0 521 26951 2
CAMBISE ISBN 0 521 33586 8
Cover design by Andrew Weall and Associates
WV
Contents
Preface Vv Chapter Seventeen Movement and support 625
Acknowledgements Vv 17.1 Skeletal systems 625
17.2 Skeletal tissue 627
Chapter Fourteen Transport 473 17.3. Anatomy of the skeleton of a mammal
Transport in plants 473 (the rabbit) 629
14.1 Plant water relations 474 17.4 The muscle system 635
14.2 Movement of water through the flowering 17.5 Innervation of skeletal muscle 641
plant 481 17.6 Locomotion in a variety of invertebrates 645
14.3 Transpiration and movement of water 17.7 Locomotion in vertebrates 652
through the leaf 481 17.8 Support in plants 663
14.4 Ascent of water in the xylem
14.5 Uptake of water by roots Chapter Eighteen Homeostasis 665
493
14.6 Uptake of mineral salts and their transport 18.1. Control systems in biology 665
across roots 495 18.2 Temperature regulation 671
Translocation of mineral salts through 18.3 Ectothermic animals 674
plants 496 18.4 Endothermic animals 675
Translocation of organic solutes in phloem 497 18.5 The mammalian liver 684
Transport in animals 507 Chapter Nineteen Excretion and
General characteristics of a circulatory osmoregulation 693
system 507 19.1 The significance of excretion and osmo-
14.10 The development of transport systems in regulation 693
animals 507 19.2 Nitrogenous excretory products 695
14.11 Composition of mammalian blood 512 19.3 Nitrogenous excretion and osmoregulation 696
14.12 The mammalian circulatory system S17 19.4 A phylogenetic review of organs and
14.13 Functions of mammalian blood i processes of nitrogenous excretion and
14.14 The immune system 534 osmoregulation 699
19.5 The mammalian kidney 710
Chapter Fifteen Coordination and control in
19.6 Anti-diuretic hormone (ADH) and the
plants 541 formation of a hypertonic or hypotonic
i5sl Plant movements 541 urine 719
1522 Plant growth substances 542 19.7. Control of blood sodium level 720
[Seo Synergism and antagonism 556 19.8 Control of blood pH qn
15.4 Phytochrome and effects of light on
plant development 559 Chapter Twenty Reproduction 723
155 Vernalisation and flowering 562 20.1 Asexual and sexual reproduction 723
20.2 Sexual reproduction in plants 732
Chapter Sixteen Coordination and control in
20.3. Sexual reproduction in Man 749
animals 565 20.4 Phylogenetic review of sexual reproduction
16.1 The nervous system 565 in vertebrates 766
16.2 The vertebrate nervous system 574
16.3 The phylogenetic development of the Chapter Twenty-one Growth and development 769
nervous system 585 Zlet Measurement of growth 769
16.4 Sensory receptors 586 Zl Patterns of growth TI
16.5 Structure and function of receptors 590 Zits Control of growth and development 774
16.6 Effectors — the endocrine system 601 21.4 Development 774
16.7, The study of behaviour (ethology) 611 ZAC Morphogenesis TS
16.8 Innate behaviour 611 ZAG Growth and development in the flowering
16.9 Learned behaviour 622 plant 776
ill
21.7 . Metamorphosis 786 24.5 Natural selection 864
21.8 Development in vertebrates 790 24.6 Modern views on evolution 866
21.9 Repair and regeneration 12 24.7 Evidence for the theory of evolution 866
21.10 Dormancy 192 Mechanisms of
Chapter Twenty-five
Chapter Twenty-two Continuity of life 796 speciation 885
22.1 The cell cycle 796 Zoe Population genetics 885
22.2 Mitosis 197 Factors producing changes in populations 888
22.3 Meiosis 801 Selection 890
22.4 The structure of chromosomes 811 Artificial selection 893
22.5 The nature of genes 815 Natural selection 896
22.6 Protein synthesis 820 The concept of species 900
22.7 Genetic control 825 Speciation 900
22.8 The genetic control of development 827 Intraspecific speciation 900
2500 Interspecific hybridisation 903
Chapter Twenty-three Variation and genetics 831
23.1 Mendel’s work 831 905
Answers and discussion
23.2 The chromosomal basis of inheritance 836
23.3 Linkage 838
Chapter
23.4 Gene mapping 841
Chapter
23.5 Linkage groups and chromosomes 842
Chapter
23.6 Sex discrimination 843
Chapter
23.7. Gene interactions 846
Chapter
23.8 Variation 851
Chapter
23.9 Mutation 852
Chapter
Chapter Twenty-four Evolution — history of life 857 Chapter
24.1 Theories of the origin of life 857 Chapter
24.2 The nature of the earliest organisms 861 Chapter
24.3 Summary of the ‘theories’ of the origin of Chapter
life 862
24.4 The theory of evolution 862 Index
Preface
The fundamental aim underlying the writing of Biological fields and by practising teachers and examiners. The text
Science was the desire to emphasise the unifying scientific includes:
nature of biological systems despite the amazing diversity — clearly written factual material,
in structure and function seen at all levels of biological —a carefully selected series of thoroughly pretested
organisation. practical investigations relevant to the A-level course,
— a variety of types of question designed to stimulate an
Books 1 and 2 comprise a complete text for the A-level enquiring approach and answers to them.
student, following all syllabuses in Biological Sciences and
incorporating all the topic areas recommended by the GCE Whilst it is recognised that the study of Biological Science
Interboard Working Party on the A-level common core in follows no set pattern, the content of books 1 and 2 has
Biology (published 1983). The text will also be relevant to been arranged so that each book contains material
all first-year University and Further Education College approximating to each year of a two-year course.
students studying the Biological Sciences. The appendices, which provide information and techni-
ques vital to the study of Biological Science at this level,
Each chapter is designed to provide comprehensive, recognise that many students do not study Chemistry and
up-to-date information on all topics in Biological Sciences, Physics to the same level. Mathematical, physical and
and the accuracy and relevance of this information has chemical concepts related to Biological Sciences are
been checked by leading authorities in the appropriate emphasised throughout the text, as appropriate.
Acknowledgements
The authors and publisher wish to acknowledge the many friends, However, the authors accept full responsibility for the final
colleagues, students and advisers who have helped in the content of these books.
production of Biological Science. Finally, the authors wish to express their thanks to their wives and
In particular, we wish to thank: families for the constant support and encouragement shown
Dr R. Batt, Dr Claudia Berek, Professor R.J. Berry, Dr A.C. throughout the preparation and publication of these books.
Blake, Dr John C. Bowman, Mr R. Brown, Dr Fred Burke, Mr We also wish to thank the following for permission to use their
Richard Carter, Dr Norman R. Cohen, Dr K.J.R. Edwards, Mr illustrations, tables and questions.
Malcolm Emery, Mr Nick Fagents, Dr James T. Fitzsimons, Dr
Figures: 14.10, 14.13, 14.15c, 14.15d, 20.1, 20.26, 21.17, Centre
John Gay, Dr Brij L. Gupta, Dr David E. Hanke, the late Dr
for Cell and Tissue Research, York; 14.11a, 20.2, 24.8b, Heather
R.N. Hardy, Reverend J.R. Hargreaves, Dr S.A. Henderson,
Angel; 14.116, A-Z Collection; 14.12, data from L.J. Briggs &
Mr Michael J. Hook, Mr Colin S. Hutchinson, Illustra Design
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Ltd, Dr Alick Jones, Mrs Susan Kearsey, Dr Simon P. Maddrell
18.41.19:4, 20.39, 20.414, 21. 26a,.21.26b, 22.185, 23.15, 23.38,
FRS, Professor Aubrey Manning, Dr Chris L. Mason, Mrs Ruth
Biophoto Associates; 14.19, data from R.N. Robertson & J.S.
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Dr R.E. Riley, Dr David Secher, Dr John M. Squire, the late 14.21b, 14.21c, Dr Chris Marshall; 14.22, Anderson & Cronshaw
Professor James F. Sutcliffe, Miss Anne C. Tallantire, Dr R.M. (1970) Planta, 91, 173-80, Springer-Verlag; 14.25a, 14.25b, Dr
Taylor, Dr Eric R. Turner, Dr Paul Wheater, Dr Brian E.J. Martin Zimmerman, Harvard University; 14.28, Professor
Wheeler, Dr Michael Wheeler. B.E.S. Gunning (1977) Science Progress, 64, 539-68, Blackwell
The authors are particularly indebted to Mrs Adrienne Oxley, Scientific Publications Ltd.; 14.29, 14.47, E.G. Springthorpe
who patiently and skilfully organised the pretesting of all the (1973) An introduction to functional systems in animals,
practical exercises. Her perseverance has produced exercises that Longman by permission of the Longman Group Limited;
teachers, pupils and laboratory technicians can depend upon. 14.30, A.E. Vines & N. Rees (1972) Plant and animal biology
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approach to lower animals (2nd ed.) Cambridge University Press; Lockwood (1963) Animal body fluids and their regulation,
14.38a, 14.38b, 14.39a, 14.39b, 14.48, 14.49, 16.8, 19.21a, 19.215, Heinemann Educational; 19.19, Professor A. Clifford Barger,
20.32, Dr Paul Wheater; 14.40, 14.59, A.G. Clegg & P.C. Clegg Harvard Medical School; 20.11a, 20.11b, Gurdon (1977) Proc.
(1963) Biology of the mammal (2nd ed.) Heinemann Medical Roy. Soc. Lond. B, 198, 211-47; 20.23; Hermann Eisenbeiss;
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Animal physiology (2nd ed.) Cambridge University Press; 14.54, no.4, by permission of the International Planned Parenthood
14.55, J.H. Green (1968) An introduction to human physiology, Federation; 20.416, Dr Everett Anderson/Science Photo Library;
Oxford University Press; 14.63, E. Florey (1967) An introduction 20.48, Research in reproduction (1972) 4, no. 5, by permission of
to general and comparative animal physiology, W.B. Saunders & the International Planned Parenthood Federation; 20.49a,
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functions, Studies in Biology no. 8, Edward Arnold; 14.70, Emil Wallace (1948) J. Agric. Sci., 38, 93, and H. Palsson & B. Vergés
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173, cover 27 August 1971, copyright © 1971 by the American animal growth and development, Studies in Biology no. 116,
Association for the Advancement of Science; 14.72, from The Edward Arnold; 21.8, J. Hammond (ed.) (1955) Progress in the
development of the immune system, Max Cooper & Alex Lawton, physiology of farm animals, 2,341, Butterworths; 21.12, R. Soper
copyright © 1974 by Scientific American Inc., all rights reserved; & §.T. Smith (1979) Modern biology, by permission of Macmil-
14.78, Macfarlane Burnet (1971) Genes, dreams and realities, lan, London and Basingstoke; 21.13, J.L. Durrer & J.P. Hannon
MUP Press Ltd.; 15.2¢,,15:2b, 17.484, 17.485, 17.48c,. Roy (1962) Am. J. Physiol. , 202, 375; 21.15, data from R. Desveaux &
Edwards; 15.3, W.O. James (1963) An introduction to plant M. Kogane-Charles (1952) Annls. Inst. Nat. Rech. Agron. Paris,
physiology (6th ed.) Oxford University Press; 15.16, Dr B.E. 3, 385-416, © INRA-PARIS 1952; 21.29, Gene Cox; 21.34, The
Juniper; 15.17, P.E. Pilet (1975) Planta, 122, 299-302; 15.18, T Natural History Photographic Agency; 22.2, 22.7a, 22.7b, 22.8,,
Swarbrick Harnessing the hormone, Grower Publications Ltd.; 22.9, Dr S.A. Henderson, Department of Genetics, University of
15.20, 20.28, Long Ashton Research Station; 15.24, Centre Cambridge; 22.3a, 23.26, ARC Poultry Research Centre; 22.11b,
Nationale de la Recherche Scientifique Regulateurs naturels de la 22,l1¢, H. G.. Callan (1963) Unt. Reva Cytol... 15, 120224
croissance vegetale (1964); 15.27, Dr Peter Evans, Southampton Professor H.G. Callan, University of St Andrews; 22.14, 22.24
University; 15.33, Professor Anton Lang (1957) Proc. Natl. Acad. E.J. Ambrose & D.M. Easty (1977) Cell biology (2nd ed.),
Sci. USA, 43, 709-17; 16.25a, E.D. Adrian & Y. Zotterman Nelson, by permission of Van Nostrand Reinhold (UK); 22.20,
(1926) J. Physiol., 61, 151-71; 16.255, B. Katz (1950) J. Physiol., from The genetic code I, F.H.C. Crick, copyright © 1962 by
111, 261-82; 16.31, R. Schmidt (ed.) (1978) Fundamentals of Scientific American Inc., all rights reserved; 22.25a, O.L. Miller
sensory physiology, Springer-Verlag; 16.43a, 16.43b, Dr I. Jr & B.A. Hamkalo, Visualization of bacterial genes in action,
Hunter-Duvar, The Hospital for Sick Children, Toronto; 16.46, Science, 169, 392-5, 24 July 1970, copyright © 1970 by the
P.J. Bentley (1976) Comparative vertebrate endocrinology, American Association fer the Advancement of Science; 24.3,
Cambridge University Press; 16.65, 16.66, Caroline E.G. Tutin; RIDA Photo Library; 24.6, G. Matthews (1939) Climate of
16.57a, N. Tinbergen (1953) The herring gull’s world, Collins; evolution (2nd ed.) vol. 1, New York Academy of Sciences; 24.7,
16.57b, N. Tinbergen & A. Purdeck (1950) Behaviour, 3, 1-38; Eric Hosking; 24.8, D. Lack (1947) Darwin’s finches, Cambridge
16.58, J. Brady (1979) Biological clocks, Studies in Biology no. University Press; 24.9a, Bruce Coleman Ltd.; 24.9b, Pig farming
104, Edward Arnold; 16.59, Niall Rankin/Eric and David magazine, Ipswich; 24.14, T.H. Hamilton (1967) Process and
Hosking; 16.60, A. Watson (1970) J. Reprod. Fert., Suppl. 11, pattern in evolution, Macmillan, London and Basingstoke; 24.18,
3-14; 16.61, J.S. Huxley (1914) Proc. Zool. Soc. Lond., 1914(2), The Zoological Society of London; 24.19, V.M. Ingram (1963)
491-562; 16.63, R.A. Hinde (1970) Animal behaviour (2nd ed.) Haemoglobins in genetics and evolution, © 1963, Columbia
McGraw-Hill Book Company, reproduced with permission; University Press, reprinted by permission; 25.3; M.N. Karn &
16.64, 16.65, N. Tinbergen (1951) A study of instinct, Oxford L.S. Penrose (1951) Ann. Eugenics, London, 16, 147-64; 25.4,
University Press; 16.68, J.B. Messenger (1977) Symp. Zool. D.S. Falconer (1953) J. Genetics, 51, 470-501; 25.5a, 25.5b,
Soc. Lond., 38, 347-76 by permission of the Zoological Photo A.G.P.M.; 25.6, Semences Nickerson, France; 25.8, D.F.
Society of London; 17.15, Lee D. Peachey (1965) Journal of Jones, Connecticut Agricultural Experiment Station 25.9, John
Cell Biology, 25, 209-31; 17.17, P.M.G. Munro, Biopoly- Haywood; 25.10a, 25.10b, Dr H.B.D. Kettlewell; PSA tile ELIS By.
mer Group, Imperial College; 17.18, A. Freundlich, Biopolymer Kettlewell (1958) Heredity, 12, 51-72, by permission of Longman
Group, Imperial College; 17.19, Dr J. Squire, Biopolymer 25.12, M.A. Tribe, I. Tallan and M.R. Eraut (1978) Case Studies
Group, Imperial College; 17.25b, David Harrison (1971) Ad- in Genetics, Cambridge University Press; 25.13, John Haywood.
vanced biology notes, by permission of Macmillan, London and
Basingstoke; 17.31, 17.35, from Cells and organelles, 2nd edition
by Alex B. Novikoff & Eric Holtzman, copyright © 1970, 1976 by Tables: 14.7, T.E. Weier, C.R. Stocking & M.G. Barbour (1970)
Holt, Rinehart and Winston, reprinted by permission of Holt, Botany: an introduction to plant biology (4th ed.), John Wiley &
Rinehart and Winston, CBS College Publishing; 17.32, M.A. Sons Inc.;16.1, A.L. Hodgkin (1958) Proc. Roy. Soc. Lond., 148,
Sleigh (1974) Cilia and flagella, Academic Press Inc.; 17.49, E.J. 1-37; 16.18, W.H. Thorpe (1963) Learning and instinct in
Sains, The Racing Pigeon, London; 17.57, Topham; 18.9, C.J.
animals, Methuen; 19.2, K. Schmidt-Nielsen (1979) Animal
Martin (1930) Lancet, (2) 108, 561; 18.14, R.N. Hardy (1979) physiology (2nd ed.), Cambridge University Press; 24.6, M.O.
Temperature and animal life, Studies in Biology no. 35 (2nd ed.) Dayhoff & R.V. Eck (1967-8) Atlas of protein sequence and
Edward Arnold; 18.21, Royal Veterinary College, Histology structure, National Biomedical Research Foundation, Silver
Department; 19.2, J. Zanefeld (1937) J. Ecology, 25, 431-68; Spring, Md.
19.3, E.H. Mercer (1959) Proc. Roy. Soc. Lond. B, 150, 216-36,
plate 16; 19.8, J.W.L. Beament (1958) J. Exp. Biol., 35, 494-519; Cover: Cockchafer in flight. Stephen Dalton/NHPA.
Vi
Chapter Fourteen
Transport
The exchange of substances between individual cells and In larger and more complex organisms, cells may be too
their environments takes place by the physical process of widely separated from each other and from their external
diffusion (which includes osmosis) and the active processes environments for these processes to be adequate. Special-
of active transport and endocytosis or exocytosis. Within ised long-distance transport systems which can move
cells, substances generally move by diffusion, but active substances more rapidly become necessary. Materials are
processes, such as cytoplasmic streaming, can also occur. generally moved by a mass flow system, mass flow being the
Over short distances these means of transport are rapid and bulk transport of materials from one point to another as a
efficient and in unicellular organisms, and multicellular result of a pressure difference between the two points. It is
organisms which possess a large surface area to volume a characteristic of mass flow that all the materials, whether
ratio, they operate efficiently enough for specific trans- they be in solution or suspension, are swept along at similar
port systems to be unnecessary. For example, respira- speeds as in a river, unlike diffusion, where molecules
tory gases are exchanged by diffusion between the body move independently of each other according to their
surface and the environment in small organisms such as the diffusion gradients. Some of the mass flow systems of plants
earthworm. and animals are summarised in table 14.1.
Transport in plants
Table 14.2 page 474 summarises the major groups of The study of translocation has important economic
substances that move through plants and gives details of the applications. For example, it is useful to know how
major routes and mechanisms for uptake, transport and herbicides, fungicides, growth regulators and nutrients
elimination. enter plants, and the routes that they take through plants,
The movement of substances through the conducting, or in order to know how best to apply them, and to judge
vascular, tissues of plants is termed translocation. In possible effects that they might have. Also, plant
vascular plants (Pteridophytes and Spermatophytes) the pathogens such as fungi, bacteria and viruses are some-
vascular tissues are highly specialised and are called xylem times translocated and such knowledge could influence
and phloem. Xylem translocates mainly water, mineral treatment or preventive measures. One relatively recent
salts, some organic nitrogen and hormones from the roots innovation has been the introduction of fungicides de-
to the aerial parts of the plant. Phloem translocates a scribed as systemic which are translocated through the
variety of organic and inorganic solutes, mainly from the plants and provide longer-term, and more thorough,
expanded leaves to other parts of the plant. protection from such important diseases as mildews.
Table 14.1 Some mass flow systems of animals and plants.
Mass flow system Material(s) moved Driving force Location
Plants
vascular system:
xylem (chapter 14) mainly water and transpiration and vascular plants:
mineral salts root pressure Pteridophyta and
mainly organic food, mechanism not Spermatophyta
phloem (chapter 14)
e.g. sucrose fully understood
Animals
alimentary system food and water muscles of Annelida, Mollusca,
(chapter 10) alimentary canal Arthropoda, Chordata
respiratory system air or water respiratory muscles Mollusca, Chordata

(chapter 11)
blood heart or contractile Annelida, Mollusca,
blood vascular system
blood vessels Arthropoda, Chordata
(chapter 14)
lymphatic system lymph general muscular Mammalia
(chapter 14) activity in the body
eee SSS S790—_.0—. 00660—500—0o00—oOamnaa—as—,
473
Table 14.2 Movement of substances through plants.
hmhah—
eA
Uptake Transport Elimination

oa
Water
route root hairs xylem stomata (cuticle and lenticels)
mechanism osmosis mass flow (root pressure) diffusion of water vapour
Solutes ae
route root hairs xylem (mainly inorganic shedding of leaves, bark, fruits and seeds; otherwise
solutes) retained until death or passed to next generation in
phloem (mainly organic embryo of seed
solutes)
mechanism __ diffusion or active transport mass flow controlled by plant growth substances
Gases*
route stomata, lenticels, root intercellular spaces and stomata, lenticels, root epidermis
epidermis through cells
mechanism __ diffusion diffusion diffusion
* Movement of gases is considered in further detail in chapter 11.
14.1 Plant water relations 14.1.3 Water potential (symbol w, the Greek
letter psi)
14.1.1 Osmosis Plant physiologists now use the term water
An understanding of plant water relations potential,* w, when describing the tendency of water
depends upon an understanding of the physical processes molecules to move from one place to another. More
of osmosis and diffusion, which are explained in section traditional terms that may be encountered are described in
A1.5. There it is pointed out that osmosis can be regarded the addendum, section 14.1.10.
as a special kind of diffusion in which water molecules are Water moves from a region of higher water potential to
the only molecules diffusing owing to the presence of a one of lower water potential.
differentially permeable membrane which does not allow By convention, water potential for pure water at
the passage of solute particles. Water molecules move from atmospheric pressure is zero. All solutions at atmospheric
a region of their high concentration (a dilute solution) to a pressure have lower water potentials and therefore
region of their low concentration (a more concentrated negative values of w. Osmosis can be redefined as the
solution) through a differentially permeable membrane. It movement of water molecules from a region of higher
is known that this process occurs more rapidly than can be water potential to a region of lower water potential through
accounted for by straightforward diffusion, and that mass a differentially permeable membrane.
flow is involved. It can, however, be explained in terms of If a pressure greater than atmospheric pressure is applied
diffusion. to pure water or a solution, its water potential increases.
Thus, water potential can be raised to a positive value, as
14.1.2 Osmotic pressure (OP) and osmotic for plasma in the glomerulus of the kidney. At atmospheric
potential (7) pressure, the water potential of a solution is determined by
its osmotic pressure, and is usually expressed in pressure
Osmotic pressure is the pressure* a solution units:
would generate if enclosed within an osmometer and
wo we OPlution
allowed to come to equilibrium with pure water. The more
concentrated a solution, the greater is its OP.
The term osmotic potential is now more commonly used, Advantage of using water potential
since under normal circumstances the ‘pressure’ referred to Water potential can be regarded as the tendency of water to
is only a potential (theoretical) pressure, and only becomes leave a system. A higher (that is less negative) water
a real pressure under the special circumstances of the
“ The origin of the term water potential may be of interest. It is a
osmometer experiment (see section A1.5.2). By conven- fundamental term derived from thermodynamics and describes the free
tion, osmotic potential is given a negative sign, and osmotic energy of water in a given system, such as in soil or in a cell. Water
pressure a positive sign. Throughout this chapter the molecules, like other molecules, always move down gradients of free
energy, from higher to lower free energy, according to the laws of entropy.
abbreviation OP is used to mean osmotic pressure, with a The process is called diffusion. The water potential of a system is the
positive sign, because negative signs can be confusing. difference between the free energy of water in that system and of pure
water at atmospheric pressure. It is usually expressed in pressure units by
* Pressure was formerly measured in atmospheres (atm), but now pascals biologists, though it can also be expressed in energy units.
are used (Pa) (see section A4.3). For a more detailed account, written with biology teachers and A-level
1 Pa = 1 Nm“ (N = newton) students in mind, see Bradbeer, Thomason & Bradbeer
1 bar = 0.987 atm = 10° Pa = 100 kPa (Dec. 1980)
School Science Review, 62, 272-3, no. 219 and Hutchinson. C.S. &
1 atm = 1.0132 bar = 1.0132 x 10° Pa = 101.32 kPa Sutcliffe, J.F. (Summer 1983) J. Biol.Ed., 17, 123-30.
474
potential implies a greater tendency to leave. If two 14.1.5 Osmosis and plant cells
systems are in contact, water will move from the system
with the higher water potential to the one with the lower The differentially permeable membranes of
potential. particular relevance in the water relations of plant cells are
Using the term water potential, the tendency for water to shown in fig 14.2. The cell wall is usually freely permeable
move between any two systems can therefore be measured, to substances in solution, so is not an osmotic barrier. The
not just from cell to cell in a plant, but also for example cell contains a large central vacuole whose contents, the
from soil to root, from leaf to air or from soil to air. Water cell sap or vacuolar sap, contribute to the osmotic pressure
can be said to move through a plant down a continuous of the cell. The two important membranes are the plasma
gradient of water potential from soil to air. The steeper the membrane and the tonoplast. In plant water relations, the
potential gradient, the faster the flow of water along it. plasma membrane, cytoplasm and tonoplast can be
regarded as acting together as one differentially permeable
membrane.
14.1.4 Movement of water between solutions
by osmosis Experiment 14.1: To investigate osmosis in
living plant cells
The terms mentioned above can only be used
with confidence if they are properly understood. The Materials
following question, together with questions 14.5-14.8, can onion bulb or young rhubarb distilled water
be a useful test of this understanding. epidermis 1 M sucrose solution
differentially permeable
microscope 2 teat pipettes
membrane 2 slides and cover-slips filter paper
solution A solution B scalpel and forceps
O solute molecules or ions Method

e.g. sugar, salt
Remove a strip of epidermis from the inner surface of one
© solvent molecules (water)
of the fleshy storage leaves of the onion bulb, or from the
solute + solvent =
solution young rhubarb petiole. Rhubarb has the advantage of
having coloured cell sap, but onion epidermis is easier to
Fig. 14.1 Two solutions separated by a differentially permeable peel off. The epidermis can be removed by first slitting it
membrane with a scalpel, then lifting and tearing back the single layer
of cells with fingers or forceps. Quickly transfer the
epidermal strip to a slide and add two or three drops of
are. separ ed | "a ferential permeable mem- distilled water. Carefully add a cover-slip and examine the
brane. cells with a microscope. Identify and draw a few epidermal
(a)Which solution has the 0 concentration of cells. Repeat using another strip of epidermis and 1M
water molecules? . sucrose solution instead of distilled water. Observe the
_ (b) Which solution is moreconcentrated, that is has strip over a period of 15min and draw any changes
the higher concentration of solute? observed in one or more representative cells at high
tc) In which direction will osmosis power. The possibility of reversing the process observed
__(d) Which solution has the highero can be investigated by irrigating with distilled water under
. PEWhich of the following two values the cover-slip to wash away the sucrose solution. Use filter
“ly 2 000 kPa, (ii) —1 000 kPa’ paper to absorb any excess liquid.
a ioWhich solutionhas the higher water potential(ys2 Results

AGS What is the relationship between OP and y of a Fig 14.3 shows the appearance of onion epidermal cells
_ solution at rele pressure? — left in 1 M sucrose solution for varying lengths of time.
cell wall - freely permeable to small molecules

plasma membrane
cytoplasm
tonoplast
plasma membrane
; : rotoplast — surrounds cytoplasm
Fig 14.2 Differentially :living Siaiars vacuolar sap, differentially
permeable membranes of part of a solution of sugars, tonoplast permeable
cell — membrane
a typical vacuolated plant salts and other solutes
surrounding vacuole
membranes
cell. Note that the plasma
membrane would normally nucleus
be pressed tightly up
against the cell wall
475
protoplast shrinking tendency for water to enter a cell is exactly balanced by
away from cell wall
turgor pressure, the amount of water leaving the cell equals
1.5 min that entering the cell. There is no further net uptake of
water and the cell is now in equilibrium with the
1.75 min different degrees of surrounding solution. The contents of the cell are still likely
plasmolysis of a
single cell (rate is
to be of higher osmotic pressure than the external solution
KK =D 2 min rapid initially, but because only a small amount of water is needed to raise the
slows)
turgor pressure to the equilibrium point, and this is not
sufficient to dilute the cell contents significantly. Turgor
So aa sm pressure therefore accounts for the fact that at equilibrium
the osmotic pressure of a plant cell can be greater than that
shrunken plasma membrane of the external solution.
protoplast
cell wall Turgor pressure is a real pressure rather than a potential
After 10 min
one, and can only develop to any extent if a cell wall is
plasmolysed cells present. Animal cells have no cell wall and the plasma
Fig 14.3 Appearance of onion epidermal cells during
membrane is too delicate to prevent the cell expanding and
plasmolysis. Strips of epidermal cells were left in 1 M sucrose bursting in a hypotonic solution. Animal cells must
solution for varying lengths of time therefore be protected by osmoregulation (chapter 19).
14.1.6 Plasmolysis and turgor pressure (TP)

14.4 Whi
s?
ch organisms, apartfrom plants,ce
If a cell is in contact with a hypertonic possess cell wall
solution, that is a solution of lower water potential than its
own contents, then water leaves the cell by osmosis through
the plasma membrane. Water is lost first from the 14.1.7 Movement of water between
cytoplasm and then from the vacuole through the tono- solutions and cells by osmosis
plast. The protoplast, that is the living contents of the cell When considering the tendency for water to
within the cell wall, shrinks and eventually pulls away from move by osmosis between a plant cell and an external
the cell wall. This process is called plasmolysis, and the cell solution, the water potential of the cell must be compared
is said to be plasmolysed. The point at which plasmolysis is with the water potential of the solution, since water always
just about to happen is called incipient plasmolysis. At moves from higher to lower water potential. The water
incipient plasmolysis the protoplast has just ceased to exert potential of the cell must equal the water potential of the
any pressure against the cell wall, so the cell is flaccid. external solution at equilibrium.
Water will continue to leave the protoplast until its As previously stated, at atmospheric pressure the water
contents are of the same water potential as the external potential of a solution is determined by its osmotic
solution. No further shrinkage then occurs.
pressure:
: 14.2 What occupies the space between tea 3 OP- a
__ the cell wall and the shrunken protoplast in plasmo--

lysed cells? i... The water potential of a cell is determined by two
factors, its osmotic pressure and its turgor pressure. As
The process of plasmolysis is usually reversible without with the external solution, the higher the osmotic pressure
permanent damage to the cell. If a plasmolysed cell is of the solution inside the cell, the lower its water potential.
placed in pure water or a hypotonic solution, that is a However, an increase in turgor pressure inside a cell
solution of higher water potential than the contents of the increases the tendency for water to leave the cell (or resists
cell, water enters the cell by osmosis. As the volume of the water entry), that is an increase in turgor pressure increases
protoplast increases it begins to exert pressure against the the water potential of the cell. This can be summarised in
cell wall and stretches it. The wall is relatively rigid, so the the equation below:
pressure inside the cell rises rapidly. The pressure exerted yor = Tp Op@!!*
by the protoplast against the cell wall is called turgor
pressure (TP). Sometimes the term wall pressure (WP) is At Piers
equilibrium, we ll =qpoeeee
i or i
yn =p
used instead, referring to the pressure of the wall against

the protoplast. The two pressures are equal and opposite in where ‘int’ means internal solution, and ‘ext’ means
direction. As the turgor pressure of the cell gradually external solution.
increases, due to water entering by osmosis, the cell
becomes turgid. Full turgidity, that is maximum TP, is * cell .
ao ' 0) better expressed as Wy = y, + Y, (see Addendum, section
achieved when a cell is placed in pure water. When the
476
The problems below are useful in testing understanding Experiment 14.2: To determine the mean osmotic
of the terms used so far. It will be necessary to remember pressure of the cell sap in a
the following points: sample of plant cells using the
(1) water always moves from higher water potential to method of incipient plas-
lower water potential; molysis
(2) for two systems to be in equilibrium, they must have
equal water potential; There are several methods available for determining
(3) in most cases the quantities of water entering and osmotic pressure of plant cells, but the most convenient is
leaving plant cells are not sufficient to significantly that of incipient plasmolysis. It makes use of the following
change their osmotic pressures (osmotic pressure is relationships:
therefore assumed to remain constant in the following (1) w of a cell = TP— OP; » of a solution =—OP.
calculations) though turgor pressures do change signifi- (2) p*" = wp" when the two are in equilibrium.
cantly.
Samples of the tissue being investigated are allowed to
come to equilibrium in a range of solutions of different
14.5. Deeaibe what happens if a cell at concentrations (water potentials) and the aim is to find
incipient plasmolysis, with an OP of 2 000 kPa, is which solution causes incipient plasmolysis, that is
placed in a solution ofOP 1 200 kPa. What are the » shrinkage of the protoplasts to the point where they
and ia oft llat equilibrium? just begin to pull away from the cell walls. At this point
If the OP of the vacuolar sap of acell c turgor pressure is zero since no pressure is exerted
hpure water at atmospheric pres- _ by the protoplasts against the cell walls, so
a, what is its TP? po = —oOp!! = wes me =—OPsauton (from (1) and (2)
Ifa cell is equilibrated with pure — above). In other words, the solution causing incipient
water ee thentransferred to a sucrose solution with | plasmolysis has the same osmotic pressure as the cell
a water potential of —800 kPa, sap.
(a) what is the difference in w between the cell In practice, osmotic pressure varies between cells in the
contents and the external solution at the Lue of same tissue and so some plasmolyse in more dilute
transfer? solutions than others. Incipient plasmolysis is said to have
. (b) would water enter or leave the been reached when 50% of the cells have plasmolysed. At
_ (c) would the TP of the cell increas crease? — this point 50% of the cells are unplasmolysed and the
average cell can be said to be at incipient plasmolysis.
The osmotic pressure obtained is a mean value for the
14.1.8 Movement of water between cells by tissue.
osmosis Materials
Consider the situation in fig 14.4, in which two onion bulb or rhubarb petiole
vacuolated cells possessing different water potentials are in 6 petri dishes
contact. 6 test-tubes
test-tube rack
freely permeable cell walls
labels or wax pencil
2 x10 cm? or 25 cm? graduated pipettes
2 x 100 cm? beakers
differentially permeable
plasma membranes brush (fine paintbrush)
distilled water
1 M sucrose solution
fine forceps
Fig 14.4 Two Pasteur pipettes
adjacent vacuolated slides and cover-slips
cells microscope
graph paper
14.8 isthe gherwater
(a) Which cell has razor blade or sharp scalpel
~ potential? :
Method
ceIn which direction will water moveSy osmosis? (An alternative method using beetroot is described after
(c) What will be the water potential of the oe at this first method.)
equilibrium? —
(1) Label six petri dishes and six test-tubes appropriately
(d)What will be ihe’ osmotic pressures and turgor for each of the following sucrose solutions: 0.3 M,
"pressures ofthe cells at equilibrium? 0.35 M, 0.4 M, 0.45 M, 0.5 M and 0.6 m.
477
Table 14.3 Sucrose dilution table for experiment 14.2. Table 14.4 Osmotic pressures of given sucrose solutions at
oe a 20 °C.
Concentration of | Volume of distilled Volume of IM oo;
sucrose solution water/cm sucrose solution/cm* Concentration of

sucrose solution Osmotic Osmotic
0.30 M 14 6 (molarity) pressure/k Pa pressure/atm
0.35 M 13 i
0.40 M 12 8 0.05 130 i3
0.45 M 11 9 0.10 260 2.6
0.50 M 10 10 0.15 410 4.0
0.60 M 8 12 0.20 540 5:3
0.25 680 6.7
0.30 820 8.1
(2) Using a 10 cm? or 25 cm* graduated pipette, a beaker 0.35 970 9.6
of distilled water and a beaker of 1M sucrose solution, 0.40 1120 11.1
make up 20 cm® of sucrose solution of the required 0.45 1280 12.6
concentration in each test-tube. Table 14.3 shows the 0.50 1 450 14.3
0.55 1620 16.0
amounts used. 0.60 1 800 17.8
(3) Make sure that the solutions are mixed thoroughly by 0.65 1980 19.5
shaking. This is very important. Add the solutions to 0.70 2 180 ZnS
the appropriate petri dishes. 0.75 2370 2323
(4) Onion. Remove one of the fleshy storage leaves of an 0.80 2 580 2505
0.85 2790 ZIeS
onion. While it is still attached to the leaf, cut the inner 0.90 3010 29.7
epidermis into six squares of approximately 5mm 0.95 3 250 32.1
side using a razor blade or scalpel. Remove each of 1.00 3510 34.6
the six squares using fine forceps and immediately 1.50 6670 65.8
place one square of tissue into each petri dish. Agitate 2.00 11 810 116.6
each dish gently to ensure that the tissue is
completely immersed and washed with the sucrose
solution. Leave for about 20 min. Results
Rhubarb. Score the outer epidermis into six squares A typical graph for onion epidermis is shown in fig 14.5.
of approximately 5mm side and remove the Similar results are obtained using rhubarb epidermis.
epidermis as described for the onion.
(5) Remove the tissue from the 0.60 M solution and, using 100
a brush, mount it on a slide in sucrose solution of the
same concentration. Add a cover-slip and examine
with a microscope. 50
(6) Select a suitable area of cells using low power. Switch
to a medium or high power objective and move the
plasmolysed/%
Cells
slide through the selected area, recording the state 0 Ol 02 03, 04905 sOemoul
(plasmolysed or unplasmolysed) of the first 100 cells Molarity of sucrose solution
viewed. Cells in which there is any sign of the Fig 14.5 Percentage of onion epidermal cells plasmolysed in
protoplast pulling away from the cell wall should be different concentrations of sucrose solution
counted as plasmolysed.
(7) Repeat for all other squares of tissue, mounting them
in their respective solutions. — 1 osmotic pressure
(8) From the total number of cells counted and number onion epidermal cells if50% of the cells eo a _
plasmolysed determine the percentage of plasmo- in 0.38 mM sucrose solution? oo.
lysed cells for each solution. Plot a graph of
percentage of plasmolysed cells (vertical axis)
against molarity of sucrose solution (horizontal axis). Use of beetroot tissue
(9) Read off from the graph the molarity of the sucrose Beetroot is a less convenient material to use, but a
solution which causes 50% of the cells to plasmolyse. combination of this experiment with experiment 14.3 would
(10) Plot a graph of osmotic pressure (vertical axis) enable an estimation of the turgor pressure of beetroot
against molarity of sucrose solution (horizontal axis) cells to be made, although it should be pointed out that
using the data provided in table 14.4. different beetroots may have different water potentials and
(11) From this graph determine the osmotic pressure of osmotic pressures. Beetroots normally have a higher
the solution which caused 50% plasmolysis. This is osmotic pressure than onion or rhubarb because they
equal to the mean osmotic pressure of the cell sap. have more sugar and inorganic salts in their vacuoles.
478
Modifications to the method are as follows. Materials
(1)-(3) As for onion and rhubarb experiment above, except
fresh potato tuber or fresh beetroot
use solutions of the following concentrations: 0.4 M,
0.45 M, 0.5 M, 0.55 M, 0.6 M and 0.7 M. 6 petri dishes
(4) Cut a rectangular ‘chip’ of beetroot with square ends of 5 test-tubes
approximately 5 x5mm. test-tube rack
Thin sections (maximum
0.5 mm thick) should be cut from the end of this chip using labels or wax pencil
2 x 10 cm® or 25 cm? graduated pipettes
a razor blade. The thinner the sections, the easier it is to
tile
count plasmolysed cells. The coloured sap enables easy
distilled water
detection of plasmolysis. The sections could be cut
1 M sucrose solution
immediately before the practical class and kept in distilled
scalpel or knife
water. Add several sections of beetroot tissue to each
sucrose solution, and leave for about 30 min. Meanwhile
2 x 100 cm? beakers
graph paper
examine similar sections, mounted in distilled water, with a
microscope to become familiar with the appearance of the
unplasmolysed cells. The margins of the sections are likely
to be thinner and easier to examine. Some damaged cells Method
may be colourless, and some small cells near vascular (1) Label six petri dishes appropriately, one for each of
tissue may be seen. These can be ignored in subsequent the following: distilled water, 0.1 M, 0.25, 0.5 M,
counts. 0.75M and 1.0M sucrose solutions. Label five
(5)—-(11) As before, starting with tissue from 0.7 m solution. test-tubes appropriately, one for each of the sucrose
solutions.
Results
(2) Using a graduated pipette, a beaker of distilled water
A set of results obtained for beetroot is given in table 14.5. and a beaker of 1 Msucrose solution, make up 20 cm?
of sucrose solution of the required concentration in
Table 14.5 Percentage of beetroot cells plasmolysed in each test-tube. A dilution table is useful as described
different concentrations of sucrose solution. in experiment 14.2 (table 14.3).
(3) Shake the tubes to mix the solutions thoroughly.
Molarity of sucrose solution Percentage of plasmolysed cells* (4) Pour the solutions into the appropriate petri dishes.
Add 20 cm of distilled water to the sixth petri
0.30 23 dish.
0.40 ae
0.45 13:5 (5) Place the petri dishes on graph paper, making sure
0.50 74.0 their lower surfaces are dry.
0.55 100.0 (6) Using the knife or scalpel, cut 12 rectangular strips of
0.60 100.0 tissue approximately 2 mm thick, 5 mm wide and as
long as possible (about 5 cm) from aslice of tissue
* Sample size 200 cells. (2 mm thick) taken from the middle of a large beetroot
or potato. It is important to work quickly to avoid loss of
. isthe mean osmotic pres- water through evaporation as this would lower the
water potential of the tissue.
sure be lls usedin this experiment?
—=
(7) Completely immerse two strips in each petri dish and
| v a graph to determine this.)
immediately measure their lengths against the graph
paper seen through the bottoms of the dishes. Agitate
the contents of each dish to wash the strips.
(8) Leave in covered petri dishes for at least 1 h,
Experiment 14.3: To determine the water poten- preferably 24 h.
tial of a plant tissue (9) Measure the lengths again, and calculate the mean
Water potential is a measure of the tendency of water percentage change in length. Plot a graph of the mean
molecules to pass from one place to another. The principle percentage change in length (vertical axis) against the
in this experiment is to discover a solution, of known water molarity of the sucrose solution (horizontal axis).
potential, in which the tissue being examined neither gains Changes in length are proportional to changes in
nor loses water. Samples of the tissue are allowed to come volume.
to equilibrium in a range of solutions of different concentra- (10) Read off from the graph the molarity of the sucrose
tions and the solution which induces neither an increase solution which causes no change in length.
nor a decrease in mass or volume of the tissue has the (11) Plot a graph of osmotic pressure (vertical axis)
same water potential as the tissue. The method described against molarity of sucrose solution (horizontal axis)
below relies on volume rather than mass changes. using the data provided in table 14.4.
479
(12) From this graph, determine the osmotic pressure of
the solution which caused no change in length. The
What |is ‘the.mean. water potel :
water potential of the tissue is determined according -tialofbe ells from the dataintable 14.6?(You |
will needtodetermine mean percentage Ghapoee in>
to the following:
length and draw graphs. J
y cell saponins solution ____ Opeenal solution
14.12 ed are at caw tipsoftissue
(13) If beetroot has been used and its osmotic pressure _ added to each dish?—
determined from experiment 14.2, calculate the 14.13 Whyaiareth
turgor pressure from: _ when left?”
wy = TP —OP 14, 14 If thes ‘osm
> cells is 1400 kPa and tl
Results - —950 kPa, what is: their mea
More accurate results are likely to be obtained by pooling 14. 15, Consider ‘the
class results. See table 14.6 for specimen results.
Table 14.6 Lengths of beetroot strips left in distilled water stalk (scape).of a dandelion (Taakeo 1 officinale) is
or different concentrations of sucrose solution for 24 h. firstcut
Looe into four ae3a eSoe
Molarity of sucrose Length of beetroot Length of beetroot

solution strip at startlcm strip after 24 h/cm
i 2 3 1 2 5
0.00 (distilled water) 4.8 5.0 5.3 eooe =kG

n concentrated /
~ (c)Why did eget bend inwards|in
0.10 5.1 48 4.9 Sea! a ee 501
0.20 5.17) 4.99 4:9 5264,90 5.0
0.25 5.246 4-8 320 doce) Wd20 sucrose solution?
0.30 49 49 5.0 AD id; emo (d) Why did scape retain
A 'thesam :curvature 1p
0.40 49 50 4.8 49 50 4.8 _ dilute sucrose solution?—
0.50 3.0% 4:8" 3.8 48 47 5.0
0.60 #85001 3.0 46 49 4.9
_ (e) Which of the following coud be
0.75 49 49 5.0 46 47 48 _ scape cells using this metho
0.90 4:9. 5.0. =4.9 45 4.7 4.7 water potential or turgor
1.00 48 49 4.9 47 46 4.4 o open to determine
LU 49 49 4.9 45 41 4.5 giving full experimental dete Is.
cut portions of scape

immediately curl
thick-walled backwards
cells of
epidermis
thin-walled
cells of cortex
—— ee
15 s after cutting
plane of cut epidermis

TS scape During cutting
epidermis
A \ B Cc
ee A, B, C Strips after they have been allowed to come to

equilibrium in different concentrations of sucrose
solution or distilled water (about 30 min)
dilute sucrose solution
Fig 14.6 Experiment on dandelion scapes.

distilled water concentrated sucrose Investigation of the effects of distilled water
solution (1 M) or drying and sucrose solutions on the curvature of strips
in air of dandelion scape
480
14.1.9 Effect of heat and alcohols on (2) Instead of the equation
membranes
The differential permeability of cell mem- yor = TP! =OP™
branes can be destroyed by certain chemicals and treat-
ments, such as ethanol and high temperatures. The
membranes are still present but behave as if holes had been plant physiologists prefer to use the equation
punched through them and they no longer provide a barrier
to the passage of large molecules such as sucrose. High Vw = Pt,
temperature and alcohols denature membrane proteins
and increase fluidity of membrane lipids; alcohols at high
concentrations can also dissolve lipids. where w, is the water potential of the cell, w, is the
contribution made by the solute concentration to the water
14,16 The red colour of beetroot is con- potential (1, is negative and equals osmotic pressure) and
tained in the cell vacuoles. Using this information, p, is the contribution made by the turgor pressure to the
riments to investigate the effects of heat water potential (this is zero or positive).
the differential permeability of —
branes.
14.2 Movement of water through the
flowering plant
14.1.10 Addendum
(1) Use of the term water potential has Water in the plant is in direct contact with
replaced use of the older terms of diffusion pressure deficit water in the soil and with water vapour in the air around the
and suction pressure. The origins and meanings of the older plant. It has already been stated that water moves from
terms are explained below. higher to lower water potentials. Plant physiologists
Diffusion pressure deficit (DPD), and suction pressure therefore think of water as moving through plants from a
(SP). It has been stated that osmosis can be regarded as a region of higher water potential in the soil to a region of
special kind of diffusion in which only water molecules lower water potential in the atmosphere, down a gradient
move. The tendency for water molecules to move used to of water potentials. The water potential of moderately dry
be expressed by the term diffusion pressure (DP). The air is several tens of thousands of kilopascals below that of
greater the concentration of water molecules in a solution, the plant; hence there is a great tendency for water to leave
the greater is the DP of the solution. At atmospheric the plant.
Most of the water entering the plant does so via the root
pressure (that is in the absence of hydrostatic pressure),
hairs. It travels across the root cortex to the xylem, ascends
pure water therefore has the maximum DP, and when
in the xylem to the leaves and is lost by evaporation from
substances dissolve in water they lower the DP of the
the surface of the mesophyll cells before diffusing out
solution. Solutions always have a lower DP than pure water
through the stomata. This latter process is called transpira-
at atmospheric pressure; the more concentrated a solution,
tion, and the flow of water from the roots to the transpiring
the less concentrated the water and the lower its DP.
surfaces forms the transpiration stream. It is estimated that
A solution was therefore said to have a diffusion pressure
less than 1% of the water absorbed is used by the average
deficit (DPD) with respect to pure water, whose DPD is set
plant. Uses of water are given in section 5.1.2.
arbitrarily at zero. The more concentrated the solution, the
greater is its DPD. For a given solution DPD = OP at
atmospheric pressure. The term DPD replaced the term
suction pressure (SP). The origin of the latter came from 14.3 Transpiration and movement of
the idea that in osmosis a hypertonic solution sucked water water through the leaf
from a hypotonic solution. The hypertonic solution was
therefore said to have a higher suction pressure. It is now Water normally leaves the plant as a vapour.
realised that the driving force for water movement comes The change fromaliquid state to a vapour state requires
from the hypotonic solution with its higher water potential the addition of energy, called the latent heat of evaporation.
(or diffusion pressure). This energy is provided by the Sun (solar energy) and
according to the most widely accepted theory of transpira-
The relationship between the terms discussed is summation, the cohesion-tension theory, it is this energy that
rised thus: maintains the flow of water through the entire plant.
—y = DPD = SP Transpiration is the loss of water vapour from the surface of
For a solution at atmospheric pressure: a plant, and may occur from the following three sites
(relative losses when stomata are open are indicated in
—y = DPD = OP = SP brackets).
481
s
(1) Stomata: by evaporation of water from cells and

diffusion of the water vapour through stomata, the Why does transpiration occ
pores found in the epidermis of leaves and green stems
mainly( leaves?
(about 90%).
(2) Cuticle: by evaporation of water from the outer walls of
epidermal cells through the waxy cuticle covering the Water is brought to the leaf in the xylem of vascular
epidermis of leaves and stems (about 10%, varying with bundles which spread to form a fine branching network
thickness of cuticle). throughout the leaf. The branches end in one or a few
(3) Lenticels: by evaporation of water through lenticels xylem vessels or tracheids possessing little lignification.
(minute proportions, although this is the main source of Water can therefore escape easily through their cellulose
water loss from deciduous trees after leaf fall). walls to the mesophyll cells they supply. Fig 14.7 shows the
The quantities of water lost by transpiration can be very three pathways which water can subsequently follow,
large. A herbaceous plant, such as cotton or sunflower, can namely the apoplast pathway (cell walls), the symplast
lose between 1-2 dm’ of water per day, and a large oak tree pathway (cytoplasm and plasmodesmata) and the vacuolar
more than 600 dm’ per day. pathway (from vacuole to vacuole).
(a) lignin cellulose xylem vessel

or tracheid
plasma deere ee
tonoplast permeable
cell wall - freely permeable mesophyll
cytoplasm cell
SYMPLAST cell sap inside vacuole

PATHWAY
VACUOLAR APOPLAST
PATHWAY PATHWAY
evaporation of water from cell

surface followed by diffusion
of water vapour through air
spaces
substomatal
air space
epidermal
cell
cuticular
stomatal transpiration
guard cell transpiration (evaporation)
(diffusion +
stoma mass flow of
molecules caused
by air movements)
482
Fig 14.7 (a) (facing page) Diagrammatic (b) cell plasmodesma cytoplasm plasma tonoplast vacuole
representation of water movement through a wall membrane
leaf. There are three possible pathways: the
symplast and vacuolar pathways are shown
to the left, the apoplast pathway to the right.
Cells A, B and C are referred to in the text.
Thickness of cell walls has been
exaggerated. (b) (right) Diagrammatic
representation of a group of cells tC, (1 it its,
aI SS IS
summarising possible pathways of water
(and solute) movement. More than one
pathway may be used simultaneously. Such
pathways may be used across the leaf and
across the root cortex. Movement of ions by —— apoplast pathway - through cell walls
the vacuolar pathway would involve active ——— symplast pathway - through cytoplasm and plasmodesmata
transport. The apoplast pathway is the most = >vacuolar pathway - through plasma membranes, cytoplasm
important, and the vacuolar pathway the and tonoplasts of vacuoles (relatively little water moves
least important (negligible) through this pathway)
14.3.1 The apoplast (apoplasm) pathway B would then have a higher water potential than cell A (at
equilibrium they would be equal). Water will therefore
The apoplasm is the system of adjacent cell
move from cell B to cell A, thus lowering the water
walls which is continuous throughout the plant (except for
potential of cell B relative to cell C. In this way a gradient of
the Casparian strip in roots, section 14.5.2). Up to 50% of
water potential is set up across the leaf from a higher
a cellulose cell wall may be ‘free space’ which can be
potential in the xylem to a lower potential in the mesophyll
occupied by water. As water evaporates from the
cells. Water enters the mesophyll cells from the xylem by
mesophyll cell walls into the intercellular air spaces, osmosis. Although it is convenient to describe the
tension develops in the continuous stream of water in the movement of water in a step-by-step fashion, it should be
apoplasm, and water is drawn through the walls in a mass stressed that the water potential gradient that develops
flow by the cohesion of water molecules (section 14.4). across the leaf is a continuous one, and water moves
Water in the apoplasm is supplied from the xylem. smoothly down the gradient as a liquid would in moving
along a wick.
14.3.2 The symplast (symplasm) pathway It is sometimes imagined that water moves across the leaf
The symplasm is the system of interconnected in response to a gradient of osmotic pressures. However,
protoplasts in the plant. The cytoplasm of neighbouring although a water potential gradient exists, there is no
protoplasts is linked by the plasmodesmata, the cytoplas- evidence to suggest that osmotic pressures of the relevant
mic strands which extend through pores in adjacent cell cells differ significantly from one another. Differences in
walls (fig 14.76). The exact structure of plasmodesmata is water potential are due mainly to differences in turgor
not yet fully explained, so to what extent they form pressure (remember that loss of a small amount of water
channels for the movement of materials is not known. from a cell has a much greater effect on turgor pressure
However, it seems likely that once water, and any solutes it than on osmotic pressure). The same applies to the root
contains, is taken into the cytoplasm of one cell it can move (section 14.5) where gradients of turgor pressure and water
through the symplasm without having to cross further potential, but not necessarily of osmotic pressure, exist.
membranes. Water would move down a water potential
gradient, as in the vacuolar pathway (section 14.3.3). 14.3.4 Exit of water through stomata
Movement might be aided by cytoplasmic streaming. The The three pathways described above end with
symplasm is a more important pathway of water movement water evaporating into air spaces. From here water vapour
than the vacuolar pathway. diffuses through the stomata, following the path of least
resistance in moving from a high water potential inside the
14.3.3 The vacuolar pathway leaf to a much lower one outside the leaf. In dicotyledons,
In the vacuolar pathway water moves from stomata are usually confined to, or are more numerous in,
vacuole to vacuole through neighbouring cells, crossing the the lower epidermis. Control of stomatal opening is
symplasm and apoplasm in the process and moving through discussed in section 14.3.9. Immediately next to the leafis a
plasma membranes and tonoplasts by osmosis (fig 14.7).
layer of stationary air whose thickness depends on the
dimensions and surface features of the leaf, such as
It moves down a water potential gradient, set up as follows.
Water evaporates from the wet walls of the mesophyll hairiness, and also on wind speed. Water vapour must
diffuse through this layer before being swept away by
cells into the intercellular air spaces, particularly into the
larger substomatal air spaces. Taking cell A in fig 14.7a as moving air (mass flow). The thinner the stationary layer,
an example, loss of water from the cell would result in a the faster is the rate of transpiration. There is a diffusion
gradient from the stationary layer back to the mesophyll
decrease in its turgor pressure and its water potential. Cell
483
lower epidermis Fig 14.8 Diffusion of water molecules from
a stoma. Note that the diffusion gradient is
stoma steeper at the edges of the pore
diffusion shell
in still air
Lines represent contours of equal concentration
of water molecules (equal water potential); the
steeper the water potential gradient, the closer
together the contours and the faster the rate of
moving air diffusion. The fastest rates are therefore from
the edges of the pores. This ‘edge effect’ means
that water loss and gaseous exchange are more
rapid through a large number of small holes than
Arrows represent curved paths of diffusion through a smaller number of large holes with the
of water molecules same total area
cells. Theoretically each stoma has a diffusion gradient, or Method

‘diffusion shell’ around it, as shown in fig 14.8. In practice (1) Select a suitable leafy plant, cut off the shoot and
the diffusion shells of neighbouring stomata overlap in still immerse the cut end immediately in a bucket of water
air to form one overall diffusion shell. to minimise the risk of air being drawn into the xylem.
Immediately cut the shoot again under water, with a
14.3.5 Measuring the rate of transpiration slanting cut, a few centimetres above the original cut.
The stem must be thick enough to fit tightly into the
Transpiration can easily be demonstrated by
bung of the potometer.
placing a bell jar over a potted plant with the pot enclosed
(2) Submerge a conical filter flask in a sink of water to fill it
in a plastic bag to prevent water loss from the soil. As
with water. Transfer the leafy shoot from bucket to sink
transpiration occurs, a fluid collects on the inside of the bell
and again immediately make a slanting cut a few
jar which is shown to contain water when tested with
centimetres above the last cut. Fit the shoot into the
cobalt(II) chloride paper (blue to pink in water) or
bung of the flask under water and push the bung in to
anhydrous copper(II) sulphate crystals (white to blue in
make atight fit.
water).
(3) Submerge the graduated capillary tube, with rubber
Measuring absolute rates of transpiration can be dif-
tubing attached, in the sink, fill it with water and attach it
ficult, but satisfactory results, at least for the purposes of
to the side arm of the filter flask.
comparison, can be obtained by means of the two simple
(4) Remove the apparatus from the sink and set up the
experiments described below.
syringe with the needle pushed into the rubber tubing
as shown in fig 14.9. The syringe can be clamped ina
Experiment 14.4: To investigate and measure vertical position. The joint between shoot and bung
factors affecting rate of tran- should be smeared with vaseline to make certain it is
Spiration using a potometer airtight.
A potometer is a piece of apparatus designed to measure (5) As the shoot takes up water the end of the water
the rate of water uptake by a cut shoot or young seedling. It column in the capillary tube can be seen to move. It
does not measure transpiration directly, but since most of may be returned to the open end of the tube by pushing
the water taken up is lost by transpiration, the two in water from the syringe. Allow the shoot to equilibrate
processes are closely related. Potometers are available for 5 min whilst regularly replacing the water taken up.
commercially, but a simple version may be set up as (6) Measure the time taken for the water column to move a
shown in fig 14.9. given distance along the capillary tube and express the
rate of water uptake in convenient units, such as
Materials cm min~'. A number of readings should be taken to
potometer (fig 14.9: conical filter flask, short rubber ensure that the rate is fairly constant, and the mean
tubing, rubber bung with a single hole, hypodermic result calculated. The temperature of the air around the
syringe and needle, graduated capillary tube) plant should be noted.
large black polythene bag _ stop clock (7) Each time the air bubble reaches the end of the
large transparent thermometer graduated section of the tube return it to its Original
polythene bag vaseline (petroleum jelly) position with the syringe.
small electric fan leafy shoot, such as lilac (8) The effects of some of the following factors on rate of
retort stand and clamp bucket uptake of water could be investigated:
484
Fig 14.9 A simple potometer
WY, O
Be NN“SX \
leafy shoot
fT
a7 retort stand
syringe containing water for pushing air

back to right-hand end of capillary tube
rubber bung
YYMy graduated capillary tube
water
Yl
L
conical filter
rubber tubing
seas fee [hans
(a) wind — use a small electric fan (do not strongly 14.3.6 Effects of environmental factors on
buffet the leaves or the stomata will close); transpiration
(b) humidity — enclose the shoot in a transparent
Plants exhibit many morphological and anato-
plastic bag;
mical features which enable them to reduce transpiration
(c) darkness — enclose the shoot in a black polythene
losses if dry conditions are encountered. Such features are
bag; described as xeromorphic and are considered below. Plants
(d) removal of half the leaves —is the transpiration rate
growing in dry habitats and thus subjected to drought are
halved?
called xerophytes and possess many xeromorphic features
(e) vaselining upper and/or lower epidermises of the
which are described in more detail in section 19.3.2. Plants
leaves to prevent water loss.
growing under conditions in which there is normally an
In each case sufficient time should be allowed to ensure
adequate water supply are called mesophytes, but never-
that the new rate has been attained. It is not always
theless can show xeromorphic features.
possible to change only one condition at a time; for
example, enclosing the plant in a transparent bag will also Light
lead to some reduction in light intensity. Light affects transpiration because stomata usually open in
Absolute rate of water uptake the light and close in darkness. At night, therefore, only
small amounts of water are lost by cuticular, and possibly
Results can be converted to actual volume of water taken lenticular transpiration. As stomata open in the morning,
up per unit time, such as cm’ h"', if the volume of the transpiration rates increase.
graduated scale corresponding to each division is deter- One group of succulent plants, the Crassulaceae, open
mined. their stomata at night and close them during the day as a
Most of the water taken up is lost through the leaves. An means of reducing water losses. Carbon dioxide enters the
estimate of rate of water loss per unit leaf area can be stomata at night, is fixed into an organic acid, and then
obtained by measuring the volume of water lost as released again inside the leaf for photosynthesis during the
described above and then removing all the leaves and day.
determining their surface area. The latter can be obtained
by drawing the outlines of the leaves on graph paper and Temperature
counting the enclosed squares. Using these data results Given the presence of light, the external factor which has
can be expressed as cm® h 'm”” leaf area. the greatest effect on transpiration is temperature. The
higher the temperature, the greater the rate of evaporation
Results of water from mesophyll cells and the greater the saturation
The effects of wind, temperature, humidity and darkness of the leaf atmosphere with water vapour. At the same
are discussed in section 14.3.6. time, a rise in temperature lowers the relative humidity of
the air outside the leaf. Both events result in a steeper .
concentration gradient of water molecules from leaf
atmosphere to external atmosphere. The steeper this
485
gradient, the faster is the rate of diffusion. Alternatively, it the moist leaf atmosphere to the external atmosphere
can be said that water potential increases inside the leaf steeper. As the concentration of water vapour in the
while decreasing outside the leaf. external atmosphere, that is the humidity, rises, the
The temperature of the leaf is raised by solar radiation. diffusion gradient becomes less steep. Water vapour
Pale-coloured leaves reflect more of this radiation than pressure of the atmosphere also decreases with altitude as
normal leaves and therefore do not heat up as rapidly. The atmospheric pressure decreases. High altitude plants
pale colour is usually due to a thick coat of epidermal hairs, therefore often show xeromorphic adaptations to reduce
waxy deposits or scales, and is a xeromorphic feature. transpiration rates.
A xeromorphic feature of some leaves is the presence of
sunken stomata, that is stomata in grooves or infoldings of
Humidity and vapour pressure the epidermis, around which a high humidity can build up
Low humidity (low water vapour pressure) outside the leaf and reduce transpiration losses. In some cases the whole
favours transpiration because it makes the diffusion leaf may roll up enclosing a humid atmosphere, such as in
gradient of water vapour (or water potential gradient) from Ammophila (marram grass) (fig 14.10). A coat of
xylem vessel
thick cuticle of
lower epidermis.
Stomata absent
from epidermis
a photosynthetic
=< cells of mesophyll
bite
hinge cells
Soa
T-dotdilied behest’
atmosphere
s
2.
ites?
ae
ir
rh
«get
unicellular
hairs trap humid
air
stoma
upper epidermis
with thin cuticle and
stomata
Fig 14.10 A transverse section of the xeromorphic leaf of Ammophila (marram grass) to show distribution of tissues. The leaf
is shown in the rolled condition
486
epidermal hairs or scales will tend to trap a layer of still transpiration is minimal and water is scarce.
moist air next to the leaf, thus reducing transpiration. The upper surfaces of dicotyledonous leaves, which are
Wind exposed to direct solar radiation and are less protected
from air currents than the lower surfaces, often possess
In still air a shell of highly saturated air builds up around
thicker cuticles than their lower surfaces. Increased wax
the leaf, thus reducing the steepness of the diffusion
deposits on leaves can virtually eliminate cuticular tran-
gradient between leaf atmosphere and external atmos-
spiration. Also, waxy leaves are usually shiny and so reflect
phere. Any disturbance, that is mass flow of the air, will
more solar radiation.
generally sweep away this shell. Thus windy conditions
result in increased transpiration rates, the increase being Stomata
most pronounced at low \wind speeds. High winds may In general, the greater the number of stomata per unit area,
result in stomatal closure and cessation of stomatal the greater is the rate of stomatal transpiration; however,
transpiration. their distribution is also important. For example, the lower
Hairs and scales trap still air as described above, tending surfaces of dicotyledonous leaves usually have more
to reduce transpiration rates. stomata than their upper surfaces (table 14.7), whereas
Availability of soil water monocotyledonous leaves, which are generally held verti-
cally rather than horizontally, have similar upper and lower
As soil dries out, water usually binds more tightly to soil surfaces with similar stomatal distributions (see maize and
particles. Also, though of less importance, the soil solution oat, table 14.7). On average, fewer stomata occur in plants
becomes more concentrated, that is its water potential adapted to dry conditions. The number may vary within the
decreases. There is therefore less tendency for water to same species as a result.
enter by osmosis. Reduced water uptake is followed shortly
by a reduction in transpiration rate as there is greater Table 14.7 Stomatal densities in the leaves of some common
resistance to movement of water through the plant, and a plants
less steep water potential gradient from the soil through the
plant to the atmosphere. Number of stomatalcm™
upper lower
14.3.7 Effect of plant or internal factors on Plant epidermis epidermis
the rate of transpiration
The effects of some xeromorphic adaptations Monocotyledons
maize (Zea mais) 5 200 6 800
on transpiration rates have been considered above. Further oat (Avena sativa) 2500 2 300
examples of the ways in which such ‘internal’ as opposed to
Dicotyledons
‘external’ (environmental) factors can operate are given apple (Malus spp.) 0 29 400
below. bean (Phaseolus vulgaris) 4000 28 100
Leaf surface area and surface area to cabbage (Brassica spp.) 14 100 22 600
lucerne (Medicago sativa) 16 900 13 800
volume ratio Nasturtium 0 13 000
Transpiration of a plant increases with its total leaf surface oak (Quercus spp.) 0 45 000
potato (Solanum tuberosum) 5 100 16 100
area, and with leaf surface area to volume ratio. Reduction 13 000
tomato (Lycopersicon esculentum) 1200
of leaf surface is achieved when leaves are reduced to
needles, such as in Pinus and other conifers, or to spines, as
Based on Weier, T. E., Stocking, C. R. and Barbour, M. G. (1970)
in cacti. There may also simply be a reduction in size in dry Botany, an Introduction to Plant Biology, 4th ed., John Wiley & Sons, p.
conditions. The shedding of leaves in dry or cold seasons by 192.
deciduous plants is a xeromorphic adaptation. In cold
seasons water may be unavailable through being frozen and Experiment 14.5: To investigate stomatal dis-
the rate of water movement through the plant is also lower tribution
at low temperatures.
Materials
Surface area to volume ratio can be reduced by using
the stem as the main photosynthetic organ, as in cacti. clear nail varnish
Fig 14.11 shows the characteristic reduction in leaf surface slides and cover-slips
area of succulents like cacti. fine forceps
fresh fully expanded leaves
Cuticle microscope
In general, the thinner the cuticle the greater the rate of .
Method
cuticular transpiration, although the composition of the
cuticle is also important. Where it is thin, as in ferns, A convenient means of examining stomatal distribution is
30-45% of the transpiration losses can be through the to make a replica of the leaf surface using clear nail
cuticle. It may assume particular importance if stomatal varnish. Spread athin layer of the nail varnish over the leaf
487
Fig 14.11 Succulent plants. (left) Silver Cholla (Opuntia
echinocarpa) (right) Kalarchoe verticilitta
14.3.8 Functions of transpiration

Transpiration has been described as a ‘neces-
sary evil’ because it is an inevitable, but potentially
harmful, consequence of the existence of wet cell walls
from which evaporation occurs. Water vapour escapes
along the route used for gaseous exchange between the
plant and its environment, which is essential for the
processes of photosynthesis and respiration. The route is
mainly through stomata, as already discussed. If there was
using the brush in the bottle. Allow it to dry, then peel off the no cuticle, stomata would be unnecessary and gaseous
thin replica with fine forceps, lay it on a slide and add a exchange would be even more efficient. However, loss of
cover-slip. It may be mounted in water for convenience. water could not then be controlled. The cuticle reduces
Examine with a microscope. Count the number of stomata water loss and further control is exercised by the stomata,
in a given field of view and repeat several times in different which in most plants are highly sensitive to water stress and
areas. Obtain a mean value. Determine the area of the field close, for example, under conditions of drought. They also
of view of the microscope by measuring the diameter with a usually close during the night when photosynthesis ceases.
calibrated slide or transparent ruler and using the formula Loss of water can lead to wilting, serious desiccation, and
mr2 for the area, where ris the radius and 7=3.142. The often death of a plant if conditions of drought are
number of stomata per square centimetre can then be experienced. There is good evidence that even mild water
calculated. stress results in reduced growth rate and, in crops, to
Compare the densities of stomata’in upper and lower economic losses through reductions in yield.
epidermises of the same leaf, and in different species. Is Despite its apparent inevitability, it is worth questioning
there any correlation between stomatal densities and the whether there might be some advantages associated with
habitats of plants? transpiration. Two possibilities are as follows.
(1) It has been suggested that the transpiration stream is
_ 14.18 Examine fig 14.12. Describe and necessary to distribute mineral salts throughout the
expl
the relation
ain veen the three variables
plant, since these move with the water. Whilst this may
be true, it seems probable that very low transpiration
rates would suffice. For example, mineral salt supply to
leaves is just as great at night, when transpiration is
low, as during the day because the xylem sap is more
_ 100 concentrated at night. Uptake of mineral salts is largely
a air temperature independent of the transpiration stream, but high rates
& 15 at crop level
s of water uptake may serve to draw water and dissolved
3B
3 transpiration
substances from more remote regions of the soil.
‘g 50 rate (2) The evaporation of water from mesophyll cells that
2 accompanies transpiration requires energy and there-
a, light fore results in cooling of the leaves in the same way that
mA intensity
sweating cools the skin of mammals. This is sometimes
0
3 6 9 noon 3 6 9 pm important under conditions of direct sunlight when
Time leaves absorb large amounts of radiant energy and
experience rises in temperature which, under extreme
Fig 14.12 Relationship between light intensity, air
temperature and transpiration rate from lucerne leaves. (From conditions, can inhibit photosynthesis. However, it is
data by L. J. Briggs & H. L. Shantz (1916) J. Agr. Res., 5, unlikely that the cooling effect is of significance under
583-649; cited by A. C. Leopold (1964) Plant growth and normal conditions. Plants that live in hot climates
development, p. 396. McGraw-Hill) usually have other means of counteracting heat stress.
488
form hoops around the sausage-shaped guard cells as
shown in fig 14.14b. As the cells inflate with water, that is
become turgid, the hoops tend to restrict the cells to an
increase in length only. Because the ends of the guard cells
are joined, and also because the thin dorsal walls stretch
more easily than the thick ventral walls, each cell assumes a
semicircular shape (fig 14.14). Thus a hole, the stoma,
appears between the guard cells. The same effect can be
sspo 8 obtained by inflating a sausage-shaped balloon which has
had a piece of adhesive tape stuck along one side to mimic
the non-elastic ventral wall of the guard cell.
Conversely, when the guard cells lose water and
turgidity, the pore closes. The question remains as to how
the turgidity changes are brought about.
A traditional hypothesis, the ‘starch-sugar hypothesis’,
yi ‘guard ‘cells
suggested that an increase in sugar concentration in guard
cells during the day led to an increase in their osmotic
pressure and entry of water by osmosis. However, sugar
has never been shown to accumulate in guard cells to the
extent necessary to cause the observed changes in osmotic
pressure. It has now been shown that an accumulation of
Fig 14.13 Scanning electron micrograph of stomata on the
lower surface of a leaf potassium ions and associated anions occurs in guard cells
during the day in response to light and is sufficient to
account for the observed changes. In darkness, potassium
14.3.9 Stomata — structure and mechanism
ions (K*) move out of the guard cells into surrounding
of opening and closing
epidermal cells. There is still doubt about which anion
Stomata are pores in the epidermis through balances the potassium. In some, but not all, species
which gaseous exchange occurs. They are found mainly in studied large quantities of organic acid anions, such as
leaves, but also in stems. Each stoma is bounded by two malate, accumulate. At the same time the starch grains that
guard cells which, unlike the other epidermal cells, possess appear in guard cell chloroplasts in darkness decrease in
chloroplasts. The guard cells control the size of the stoma size, Suggesting that starch is converted to malate in light.
by changes in their turgidity. The appearance of guard cells A possible route is:
and stomata is well revealed by the scanning electron PEP carboxylase malate dehydrogenase
microscope, as shown in fig 14.13. PEP + CO,—~» oxaloacetate malate
The appearance of epidermal cells, guard cells and phosphoenolpyruvate
NADPH, NADP
stomata in surface view, as seen with the light microscope, series of reactions
is dealt with in section 8.1. Fig 14.14 is a diagram of a sec- starch
tion through a typical stoma, and shows that the guard cell
walls are unevenly thickened, the wall furthest from the (Compare C, photosynthesis, section 9.8.2.)
pore (termed the dorsal wall) being thinner than that next Some species, such as Allium cepa (onion), have no
to the pore (the ventral wall). Also, the cellulose starch in their guard cells. Here malate does not accumu-
microfibrils that make up the walls are arranged so that the late during stomatal opening and inorganic anions, such as
ventral wall is less elastic than the dorsal wall, and some chloride (Cl-), may be taken up with the cations.
Certain questions remain to be answered. For example,
(a) stoma thick ventral wall of why is light necessary for stomatal opening, and what
guard cell function is served by the chloroplasts apart from starch
ROROpIAS! thin dorsal wall of
guard cell storage? Is malate converted back to starch in darkness? In
‘ ) (b) 1979 it was shown that the enzymes of the Calvin cycle are
small vacuole
cellulose
absent from the chloroplasts of guard cells of Vicia faba
cytoplasm microfibrils (broad bean), and the thylakoid system is poorly de-
pattern of cellulose veloped, although chlorophyll is present. Normal C,
microfibrils around photosynthesis therefore cannot occur and starch cannot
circumference of
guard cells be made by this route. This might help to explain why
two guard cells in a
surface view
epidermal cell starch is made at night rather than during the day as in
normal photosynthetic cells. Another interesting fact is
Fig 14.14 (a) Vertical section through a stoma, showing also that guard cells lack plasmodesmata and are therefore
part of the lower surface of the leat. (b) Pattern of cellulose
relatively isolated from other epidermal cells.
microfibrils in guard cell walls
489
Vascular bundle
(a) HP detail seebelow _(b)
intrafascicular
cambium
interfascicular
cambium
epidermal hair. cuticle
(multicellular in as dicot
Helianthus)
epidermis
cuticle as dicot
made of cutin
hypodermis
epidermis collenchyma, outer cells
one cell thick may possess chloroplasts
cortex collenchyma— ground tissue
outer cells parenchyma (no separate
may possess parenchyma cortex and pith)
chloroplasts vascular bundles
endodermis 9 HP detail see below
one cell thick, may form scattered and numerous,
a ‘starch sheath’ i.e. larger towards centre
cells may store starch,
rarely obvious unless
specially stained
parenchyma |
P
pericycle pericycle or bundle sheath
sclerenchyma fibres sclerenchyma, surrounding bundle Ii
phloem phloem
sieve tubes, companion cells, phloem sieve tubes and companion
parenchyma, phloem fibres cells only
intrafascicular cambium metaxylem
metaxylem two large vessels
mature xylem vessels, fibres and protoxylem
parenchyma endarch (see dicot)
\ TS vascular
TS vascular bundle
bundle
protoxylem endarch, i.e. nearer centre lysigenous canal formed from

than metaxylem; annular and spirally breakdown of protoxylem
thickened vessels
(d)
epidermis
collenchyma
parenchyma
ground tissue
~22— of cortex
t»-—collenchyma
bundles
(see fig. 14.15f)
(secondary growth
has started)
Fig 14.15 (a) (top left) Primary anatomy of the stem of a
parenchyma typical dicotyledon, Helianthus annuus (sunflower). (b) (top
right) Anatomy of the stem of a typical monocotyledon, Zea
mais (maize). (c) (above left) Low power micrograph of part of
a TS of the stem of Helianthus. (d) (above right) Low power
micrograph of part of a TS of the stem of Zea. (e) (facing
page, top left) High power micrograph of a TS of a vascular
bundle from a stem of Helianthus. (f) (facing page, top right)
epidermal High power micrograph of a TS of a vascular bundle from a
hair stem of Zea (g) (facing page, centre) Micrograph of an LS of
the stem of Helianthus (h) (facing page, bottom) Micrograph
of an LS of the stem of Zea
aa
A mypericycle of
ad pericycle of
. .
eye sclerenchyma fibres

sclerenchyma
metaxylem
vessels
+ protoxylem
# lysigenous cavity
ye formed at
7/ {breakdown of
~ protoxylem
40
Ph va 6... sclerenchyma
ai 7 fibres
spiral vessels
annular vessel
reticulate vessel
491
The cohesion-tension theory (or cohesion theory) of
14.4 Ascent of water in the xylem water movement adequately accounts for these observa-
tions. According to this theory, evaporation of water from
Xylem in flowering plants contains two types
the cells of a leaf is responsible for raising water from the
of water-transporting cell, the tracheid and the vessel,
roots. Evaporation results in a reduced water potential in
whose structures as seen in the light microscope are
the cells next to the xylem as described in section 14.3.
discussed in section 8.2.1, together with the appearance of
Water therefore enters these cells from the xylem sap
vessels as seen with the scanning electron microscope (fig
which has a higher water potential, passing through the
8.11). The structure of the secondary xylem (wood) is dealt
moist cellulose cell walls of the xylem vessels at the ends of
with in section 21.6.6.
Xylem, together with phloem, forms the vascular or
the veins, as shown in fig 14.7.
The xylem vessels are full of water and as water leaves
conducting tissue of higher plants. Vascular tissue consists
them a tension is set up in the columns of water. This is
of bundles of tubes called vascular bundles whose structure
transmitted back down the stem all the way to the root by
and arrangement in the primary stems of dicotyledonous
cohesion of water molecules. Water molecules have high
plants (dicots) and monocotyledonous plants (monocots)
cohesion, that is tend to ‘stick’ to each other, because,
are shown in fig 14.15.
being polar, they are electrically attracted to each other
and are held together by hydrogen bonding (section 5.1.2).
14.19 Summarise in table form the major
They also tend to stick to the vessel walls, a force called
differences in primary structure between dicot and
adhesion. The high cohesion of water molecules means that
monocot stems. .
a relatively large tension is required to break a column of
14.20 W ;the overall shape of the. water, that is a water column has a high tensile strength.
_ following tissues in three dimensions: (a) epidermis, The tension in the xylem vessels builds up to a force
(b) xylem, (c) pericycle
of dicots and (d) pith? capable of pulling the whole column of water upwards by
means of mass flow, and water enters the base of the
The fact that water can move up the xylem may be columns in the roots from neighbouring root cells. It is
demonstrated by immersing the cut end of a shoot in a essential that the xylem walls should also have high tensile
dilute solution of a dye, such as eosin. The dye rises in the strength if they are not to buckle inwards, as happens when
xylem and spreads through the network of veins in the sucking up a soggy straw. Lignin and cellulose both provide
leaves. Sectioning and examination with a light microscope this strength. Evidence that the contents of xylem vessels
reveals the stain to be in the xylem. are under high tension comes from measuring diurnal
Better evidence that xylem conducts water is given by changes in the diameters of tree trunks using an instrument
‘ringing’ experiments. These were among earlier experi- called a dendrogram. The minimum diameters are re-
ments done before radioactive isotopes made the tracing of corded during daylight hours when transpiration rates are
substances through living organisms much easier. In one highest. The minute shrinkage of each xylem vessel under
type of ringing experiment an outer ring of bark, including tension combines to give a measurable shrinkage in
phloem, is removed and, in the short term, this does not diameter of the whole trunk.
affect the upward movement of water. However, lifting a Estimates of the tensile strength of a column of xylem sap
flap of bark, removing a section of xylem, and replacing the vary from about 3 000-30 000 kPa, the lower estimates
flap of bark leads to rapid wilting. being the more recent. Water potentials of the order
Any theory for water movement up the xylem has to required to generate enough tension to raise water, about
account for the following observations. —4 000 kPa, have been recorded in leaves, and it seems
(1) Xylem vessels are dead tubes with narrow lumens likely that xylem sap has the required tensile strength to
ranging in diameter from 0.01 mm in ‘summer wood’ to withstand this tension, though there may be a tendency for
about 0.2 mm in ‘spring wood’. the columns to break, particularly in vessels of relatively
(2) Large quantities of water are carried at relatively high large diameter.
speeds, up to 8 m h"' being recorded in tall trees and Critics of the theory point to the fact that any break in a
commonly in other plants at 1 m h"!. column of sap should stop its flow, the vessel tending to fill
(3) To move water through such tubes to the height ofatall with air and water vapour, a process known as cavitation.
tree requires pressures of around 4 000 kPa. The tallest Shaking, bending and shortage of water can all induce
trees, the giant sequoias or redwoods of California cavitation. It is well known that the water content of tree
(conifers and therefore possessing only tracheids, not trunks gradually decreases during summer as the wood
vessels) and Eucalyptus trees or blue gums of ‘Austra- becomes filled with air. This is made use of in the lumber
lia, can reach heights greater than 100 m. Water will industry because such wood floats more easily. However,
rise in fine capillary tubes due to its high surface breaks in water columns do not greatly affect water flow
tension, a phenomenon called capillarity, but could rise rates. The explanation may be that water flows from one
only about 3 m in even the finest xylem vessels by this vessel to another, or by-passes air-locks by moving through
method. neighbouring parenchyma cells and their walls. Also, it is
492
calculated that only a small proportion of the vessels need favour low transpiration rates, including dim light and high
be functional at any one time to account for the observed humidity. It is common in many rain forest species and is
flow rates. In some trees and shrubs water moves only frequently seen at the tips of the leaves of young grass
through the younger outer wood, which is therefore called seedlings.
sapwood. In oak and ash, for example, water moves mainly
through the vessels of the current year, the rest of the 7 14.21 Summarise the properties of xylem
sapwood acting as a water reserve. New vessels are added which make it suitable for the long-distance transport
throughout the growing season, mostly early in the season of water and solutes. _.
when flow rates are higher.
A second force involved in water movement up the
xylem is root pressure. This can be observed and measured 14.5 Uptake of water by roots
when afreshly cut root stump continues to exude sap from
its xylem vessels. The process is inhibited by respiratory The primary structures of typical dicot and
inhibitors such as cyanide, lack of oxygen and low monocot roots are shown in fig 14.16.
temperatures. The mechanism probably depends on active
secretion of salts or other solutes into the xylem sap, thus | _ 14.22 Summarise in table form the major
lowering its water potential. Water then moves into the differences
in primary structure between typical dicot _
xylem by osmosis from neighbouring root cells.
The positive hydrostatic pressure of around 100-200 kPa
__and monocot roots.
(exceptionally 800 kPa) that is generated by root pressure Water is absorbed mainly, but not exclusively, by the
is usually not sufficient alone to account for water younger parts of roots in the regions of the root hairs. As a
movement up the xylem but it is no doubt a contributing root grows through the soil, new root hairs develop a short
factor in many plants. It can be sufficient, however, in distance behind the zone of elongation and older hairs
)
slowly transpiring herbaceous plants, when it can cause die. These hairs are tubular extensions of epidermal cells
guttation. Guttation is the loss of water as drops of liquid (fig 14.16) and greatly increase the available surface area
from the surface of a plant (as opposed to vapour in for uptake of water and mineral salts. They form a very
transpiration). It is favoured by the same conditions that intimate relationship with soil particles.
(a) root hair root hair b)

rN
piliferous layer outgrowth ofa single cell as dicot
epidermis modified to produce piliferous layer
root hairs.One cell thick, no cuticle, as dicot but retained
soon lost from dicots (often absent
in prepared slides) exodermis
as dicot
exodermis
protective layer one to several cells cortex
thick, a type of hypodermis, as dicot
sometimes like a root endodermis
in structure and function, with endodermis
suberised walls. May replace as dicot
epidermis in older parts Ca \
cortex root)
parenchyma, starch may be stored
ne
endodermis as dicot but does not remain
one cell thick, cells possess meristematic
Casparian strip of suberin (fig T
14.175), may form a starch sheath e phloem
.
alternates with xylem
pericycle
one to several cells thick
parenchyma. In dicots it remains exarch, in more than eight
meristematic, producing lateral groups, 1.e. polyarch
roots and contributing to LS metaxyle
secondary growth m
phloem : pith
alternates with xylem parenchyma, may or may not be
present
protoxylem
exarch, i.e. nearer outside than
metaxylem.Annular and spirally
thickened vessels, 2-8 groups in cambium
: ;
triarch, eer Saeatan
dicots, calleddiarch,
Be (this example is fea tae Gong be ents
ro in this case) =
(b) Primary anatomy of the root of

Fig 14.16 (a) Primary anatomy of the root of a typical dicotyledon, Ranunculus (buttercup).
(d) overleaf
a typical monocotyledon, Zea mais (maize). (c) and
493
exodermis Lat
(epidermis lost) =. you Fe 7%
piliferous
cortex of
layer
parenchyma #
ae BROS s ry : —__ exodermis
_ of fibres
phloem. oa
cortex of
parenchyma
endodermis with
U-shaped thickening
passage cell in pericycle of
endodermis parenchyma
endodermis
pith of metaxylem protoxylem ~phloem
parenchyma
(c) Low power micrograph of a TS of the root of Ranunculus (d) Low power micrograph of a TS of the root of Zea
Fig 14.17a is a diagrammatic representation of the such as cell A in fig 14.17a. As water leaves cell A, the
pathway taken by water across a root. A water potential water potential of cell A decreases and water enters it from
gradient exists across the root from higher potential in the cell B by osmosis or through the symplasm in exactly the
piliferous layer to lower potential in the cells adjacent to same way as described for cells A and Bin the leaf (section
the xylem. This gradient is maintained in two ways: 14.3.2). Similarly the water potential of cell B then
(1) by water moving up the xylem, as described, setting up decreases and water enters it from cell C and so on across
tension in the xylem and thus lowering the water the root to the piliferous layer.
potential of its sap; The soil solution is normally fairly dilute, so it has a
(2) the xylem sap has a higher osmotic pressure than the higher water potential than cells of the piliferous layer,
dilute soil solution. which include the root hairs. Water therefore enters the
Water moves across the root by pathways similar to those root from the soil by osmosis.
in the leaf, namely apoplast, symplast and vacuolar
pathways.
14.5.1 Symplast and vacuolar pathways 14.23 Arrange th
3)
soilsolution, xylem sap, cell
A,
As water moves up the xylem in the root, it is 2 cell. (Use the symbol > (greater thar
replaced by water from neighbouring parenchyma cells,
(a) cell wall piliferous

layer <——______—_cortex endodermis
cytoplasm
vacuole pericycle
root hair air + water
xylem vessels
APOPLAST PATHWAY
(interrupted at endodermis)
thin film of water,
tightly bound if ——> SYMPLAST PATHWAY
clay particle ~——)> VACUOLAR PATHWAY
Fig 14.17 (a) Diagrammatic representation of water ; and ion movement across a root shown in TS. . ThiThickness o ;
exaggerated for clarity. Cells A, B, C are referred to in the text. The apoplast pathway is of greatest importance neCone
and solutes. The symplast pathway is less important, except for salts in the reg ion of the endodermis. Movement along the
vacuolar pathway is negligible. (b) and (c) facing page
494
(b) Casparian strip seen in TS appears aS passage cell (c) TS
as a ‘dot’ with light microscope outer tangential
YOUNG OLD wall
inner tangential
extra suberi
be In wall
radial wall transverse wall
Casparian strip
encircles cell extra secondary passage cell lacks
thickening extra thickening 3D tangential wall
consisting of
more suberin
3D appearance of a 3D appearance of three Fig 14.17 (b) Structure and function of root
single endodermal cell eae endodermal ~~ endodermis showing Casparian strip in young
cells age ne
endodermal cells and deposition of extra suberin in
YOUNG OLD older endodermal cells, with exception of ‘passage
movement of water and solutes, cells’. (c) Naming of walls. The transverse and radial
e.g. mineral salts walls are anticlinal (at right-angles to the surface of
the root) and the tangential wall is periclinal (parallel).
14.5.2 Apoplast pathway Mineral elements exist in the form of ions in salts, and in
solution the ions dissociate and move about freely. In
The apoplast pathway operates in much the
attempting to explain the uptake and movement of mineral
same way as in the leaf (section 14.3.1). However, there is
ions, the following facts must be taken into account.
one important difference. When water moving through (1) Cell membranes, including the plasma membrane and
spaces in the cell walls reaches the endodermis its progress tonoplast, are not truly semi-permeable but dif-
is barred by the waterproof substance called suberin which ferentially permeable, allowing to varying extents the
is deposited in the cell walls in the form of bands called passage of substances other than water, such as ions.
Casparian strips. These strips prevent apoplastic move- (2) Active transport can occur across cell membranes. This
ment of water (fig 14.17b) and therefore water and salts requires energy in the form of ATP, made during
must pass through the plasma membrane under the respiration, and can lead to an accumulation of ions
cytoplasmic control of the endodermal cell. In this way, it is against a concentration gradient (section 7.2.2).
believed, control by living cells is exercised over the (3) There is a continuous system of cell walls, the
movement of water and mineral salts from soil to xylem. apoplasm, extending inwards from the piliferous layer
Such control is necessary to regulate salt movement and of the root. Water, and any solutes it contains, enters
may be a protective measure against entry of toxic the system from the soil by mass flow and toa lesser
substances, fungal pathogens, and so on. It is interesting to extent by diffusion.
note that when endodermal cells are plasmolysed, their (4) Water moves through the apoplasm as part of the
cytoplasm remains attached to the Casparian strip even transpiration stream.
when shrinking away from the rest of the cell wall. As roots Fig 14.18 shows the uptake of potassium ions by young
get older suberisation in the endodermis often gets more cereal roots which had previously been thoroughly washed
extensive as shown in fig 14.17b. This blocks the normal in pure water. After 90 min the respiratory inhibitor
exit of water and mineral salts through the inner tangential potassium cyanide was added to the solutions.
walls. However, plasmodesmata may stay as pores in these
walls, and ‘passage cells’ in which no extra thickening
occurs also remain to allow water and solute movement.
The relative importance of apoplast, symplast and
3 potassium cyanide added |
vacuolar pathways is not known.
14.6 Uptake of mineral salts and their SS

transport across roots
As part of their nutrition, plants require uptake/yg
Potassium
sample
root
per
certain mineral elements in addition to the carbohydrates
10 20 30 40 50 60 70 80 90 100 110 120 130
made in photosynthesis. The uses of these elements are Time/min
described in table 9.10. In higher plants minerals are taken
up from the soil or surrounding water by roots. Uptake is Fig 14.18 Absorption of potassium ions by young cereal
plants in aerated solution
greatest in the region of the root hairs.
495
endodermis, ions must pass by diffusion or active transport
14.24 (a) Describe the uptake of potas- through the plasma membranes of endodermal cells,
sium ions at 0 °C and 25 °C. entering their cytoplasm and possibly their vacuoles. Thus
(b) Explain the differences described, and the effect the plant monitors and controls which types of ions
of potassium cyanide (KCN). eventually reach the xylem.
Similar results to those in fig 14.18 can be obtained 14.29 Howcould you demonstrate, using
with isolated tissues, those of storage organs, such as a radioactive ion and autoradiography, ‘that the
carrot, being commonly used. The data shown in fig endodermis is a barrier to the
14.19 confirm the inhibition of respiration by potassium throug)cell a
cyanide.
Ions can also move through the symplast pathway. Once
Z transfer y potassium they are taken into the cytoplasm of one cell, they can
s from pure cyanide move through the symplasm without having to cross further
= & || water to rate of | added
eo potassium respiration membranes. The symplasm extends from the piliferous
SE | chloride
go solution ; layer right through to the xylem. Fig 14.17a summarises the
aS potassium
possible ways in which ions can cross the root.
o€ chloride
aS content The final stage in the movement of mineral salts across
o4
n
the root is the release of ions into the xylem. To achieve

SO
~ a, this, ions must leave living cells at some stage, crossing
0 60 120 180 240 back through a plasma membrane. This could be by
Time/min
diffusion or active transport.
Fig 14.19 Rate of respiration and uptake of potassium
chloride by carrot discs. (Based on data by Robertson &
Turner (1945))
14.7 Translocation of mineral salts
through plants
14.25 Fig 14.19 shows that the rate of The pathway of mineral salts across the root to
respiration of carrot discs increases when they are _ the xylem, described above, is the first stage in their
__ transferred from pure water to potassit translocation. Once in the xylem, they are distributed
solution. From the results shown, ae throughout the plant by the transpiration stream, in which
they move by mass flow. Movement of the mineral
/ 44. 26 Why does the rise in pota: elements in the xylem can be demonstrated by ringing
experiments like those already described, in which removal
chloride content stop when KCN is added? of tissues external to the xylem, such as phloem, has no
. 14.27 In an experiment similar to. that effect on upward movement of ions. Analysis of the xylem
“described in fig 14.18, but involving phosphate — sap also reveals that although some of the nitrogen travels
uptake, 16% of the phosphate taken up by barley as inorganic nitrate or ammonium ions, much ofit is carried
roots over a short period could be washed out after in the organic form of amino acids and related compounds.
sy to pure wate again. Explain. Some conversion of these ions to amino acids must
14.28 Cot ach the xylem entirely therefore take place in the roots. Similarly small amounts
by meansofthe 0 pathway? of phosphorus and sulphur are carried as organic com-
pounds.
To summarise so an fhe uptake of ions by roots is a Thus, although xylem and phloem are traditionally
combination of passive uptake, whereby ions move by mass regarded as conducting inorganic and organic materials
flow and diffusion through the apoplasm, and active respectively, the distinction is not clear-cut. In addition, a
uptake, or active transport, whereby ions can be taken up small amount of exchange between xylem and phloem is
into cells against a concentration gradient using energy common and phloem carries significant quantities of
from respiration. mineral elements away from organs other than the root, as
Active transport is selective and dependent on respira- discussed below.
tion, whereas diffusion is non-selective and not dependent The chief sinks, that is sites of utilisation, for mineral
on respiration. Each cell of the root cortex is bathed in a elements are the growing regions of the plant, such as the
solution similar in composition to that of the soil solution as apical and lateral meristems, young leaves, developing
a result of passive uptake. Thus there is a large surface area fruits and flowers, and storage organs. Unloading of solutes
for ion uptake. from xylem occurs at the fine vein endings and entry of
Ions moving in the apoplasm can only reach the solutes into cells can take place by diffusion and active
endodermis, where the Casparian strip prevents further uptake. Transfer cells (section 14.8.6) may sometimes be
progress as described in section 14.5.2. To cross the involved.
496
14.7.1 Recirculation and remobilisation Fig 14.20 Movement of organic solutes in a green plant
Very often the path from roots to sinks via photosynthetic tissue
xylem or phloem is not the end of translocation of mineral (mainly leaves)
elements. Generally speaking, xylem makes the initial
delivery of a given element to an organ, and then phloem
carries away the element for continued translocation up or
down the plant if it is not required by that organ. This
recirculation can be investigated with radioactive isotopes
and is shown to be a common phenomenon. In one
experiment, the roots of a maize plant were separated into storage centres other growth centres
(stem, roots, bulbs, > (meristems, younger
two beakers of nutrient solution, one containing radio- tubers etc.) leaves, flowers and
active phosphorus. Within 6h the phosphorus had fruits)
travelled up the stem in the xylem, back down in the
(losing food) or as sinks (gaining food) at different times.
phloem and had been detected in the second beaker. This is
Typically, about 90% of the total solute carried in the
part of the evidence which suggests that phosphorus
phloem is the carbohydrate sucrose, a disaccharide. This is
circulates continuously and rapidly within the xylem and
a relatively inert and highly soluble sugar, playing little
phloem of plants. Other elements are less mobile. Sulphur,
direct role in metabolism and so making an ideal transport
for example, is mainly removed during its first circulation,
sugar, being unlikely to be used in transit. Once at its
and calcium is notoriously immobile, being virtually
destination it can be converted back to the more active
trapped in any organ where it is deposited because it has
monosaccharides. It can be present in very high concentra-
poor mobility within the phloem.
tions, up to 25% mass to volume in the phloem of plants
Often an element will stay in an organ for some time and
such as sugarcane.
then be remobilised, that is leave the organ for some other
It should be noted that phloem also carries certain
part of the plant after having served some useful function. mineral elements in various forms, particularly nitrogen
This occurs during sequential senescence (ageing) of and sulphur in the form of amino acids, phosphorus in the
leaves, when older dying leaves export much of their form of inorganic phosphate ions and sugar phosphates,
mineral content to younger leaves. Similarly, prior to
and potassium ions. Small amounts of vitamins, growth
abscission of leaves by deciduous trees and shrubs useful substances such as auxins and gibberellins, artificially
minerals can be conserved by remobilising them for storage applied chemicals, viruses and other components may also
elsewhere. Development of flowers, fruits and seeds, and be present. The importance of phloem in recirculation and
storage organs also involves remobilisation. Directional remobilisation of mineral elements is in section 14.7.1.
control of the movement of nutrients is under the influence Evidence for the circulation of carbon within the plant
of plant growth substances, particularly cytokinins (section can be obtained by supplying leaves with carbon dioxide
15.2.7). The elements most readily mobilised are phos- containing the radioactive isotope “C. Radioactive carbon
phorus, sulphur, nitrogen and potassium. dioxide is fixed in photosynthesis and “C passes into an
organic solute such as sucrose. Its movement around the
14.8 Translocation of organic solutes plant can then be traced by various techniques used to
in phloem locate radioactive isotopes, such as autoradiography,
In those multicellular plants where certain application of a Geiger counter to the plant surface, or
extraction of the isotope from different parts. Ultimately,
parts of the plant, such as roots, are some distance from the
both phloem and xylem will be intimately associated in the
sites of photosynthesis, there is a need for a transport
circulation of carbon. For example, carbon in the form of
system to circulate the products of photosynthesis. In
sucrose may reach the roots and there be used in the
vascular plants phloem is the tissue which carries these
conversion of nitrates to amino acids. The latter, contain-
organic products away from the leaves, the main organs of
ing the carbon, can then travel up the shoots in the xylem.
photosynthesis, to other parts. Phloem consists of sieve
elements, companion cells, parenchyma, fibres and Experiment 14.6: To investigate the pattern of
sclereids. Its structure, as revealed by the light microscope, distribution of the products of photosyn-
is described in section 8.2.2. The sieve elements are thesis in a pea plant
arranged end to end to form sieve tubes, each element
being separated from the next bya sieve plate. Fig 14.20 The experiment and data below are based on the
summarises the relationship between autotrophic cells
Independent Television schools’ programme An investiga-
tion of photosynthesis and assimilate transport. Providing
producing organic food and those receiving the food. Note
the terms of licence granted by the Independent Television
from fig 14.20 that movement of organic solutes must be up
Companies to local authorities are complied with, this
and down in the same plant, that is bidirectional. This
with movement in the xylem, which is only programme could be usefully recorded on a video-
contrasts
cassette recorder.
upwards. Note also that storage organs act either as sources
497
In the experiment below, carbon dioxide containing the (p)
radioactive isotope of carbon, '*C, is supplied to photo-
synthesising pea plants (Pisum sativum). '*C is a useful
isotope because it has a long half-life (5 570 years) and
therefore retains its radioactivity throughout the experi-
ment. (Compare ''C which has a half-life of 20.5 min.) It is
also relatively safe to handle because it emits only weak
radiation (low energy particles).
“CO, is used by the plant in photosynthesis in exactly
the same way as the usual carbon dioxide ('*CO,) and is
incorporated into the products of photosynthesis (assimi-
lates). The movement of these assimilates can be followed
in the experiment because they contain 'C; they are said
to be ‘C-labelled.
Method
Fig 14.21 outlines the stages of an experiment in which
“CO, is fed either to a lower leaf or an upper leaf of
separate pea plants with one pod. After feeding and
allowing 24 h for transport of '*C-labelled assimilates from
the leaves to other parts of the plant, the pattern of
distribution of activity is revealed either by autoradio-
graphy (fig 14.21) or by measuring the amount of '*C in
each plant part. If autoradiography is done first, the plants
can subsequently be cut up and used for the second
determination which involves combustion.
(a)
(c)
<a
\ wee of
Sellotape
498
Fig. 14.21(a)
Peaplant with selected leaf in a well-illuminated
bag into which '*CO, is introduced .Total
exposure time is 15-30 min.
Leaf removed from bag, and plant left

for 24 h to allow transport of '4C-labelled
assimilate (mainly '*C-sucrose) from the
leaves to other parts of the plant
Whole plant Quantitative determination

autoradiography of '*C content of different
plant parts
Autoradiography of '*C-labelled plants plant is cut up into its component parts
Fig. 14.21(b)
plant after drying and mounting on apical 'C-treated other stem pod (if roots
paper, the arrow indicates the leaf that regionof leaf leaves present)
was exposed to '*CO, shoot
Fig. 14.21(c)
autoradiograph obtained by placing
X-ray film over dried plant and
leaving in a dark room for four
days; the B-radiation emitted by
the '*C exposes the X-ray film, the each part is combusted with O,
blackest areas being those with the in a furnace; '*C-containing
most !*C compounds release '*CO, which
is absorbed into a
scintillation fluid for
scintillation counting
radioactive sample + scintillation fluid. As

'4C decays, the energy of the }-particles
radiated causes the scintillation fluid to
fluoresce; the sample is placed in an
instrument called a scintillation counter
where the tiny flashes of light emitted are
amplified to generate an electrical signal
and a count rate is obtained which is
proportional to the amount of '4C present
Fig 14.21 (opposite and above) Stages in an experiment to

investigate the pattern of distribution of “C-labelled
assimilates in a pea plant
499
Results (3) The distances travelled can be very large. The tallest
Fig 14.21 shows the results of autoradiography. Table 14.8
trees, such as Eucalyptus, may be over 100 mtall. The
shows the results of counting the '*C in each plant part. leaves of Eucalyptus trees are located mainly near the
top of the trunk, so assimilates must travel the length of
Table 14.8 Radioactivity of different parts of a pea plant the stem and often a considerable distance through the
24 h after feeding with “CO,. roots.
(4) The amount of phloem is not great. In a tree trunk, the
Upper leaf Lower leaf
treated/counts treated/counts
functional phloem tissue is a layer only about the
Plant part min !* min ‘* thickness of a postcard around the circumference. It
forms the innermost layer of the bark of woody stems
apical region of shoot 1123 739 and roots, the older phloem becoming stretched and
“C-treated leaf 11325 11372 dying as the plant grows and its circumference
other leaves 234 168
stem 819 1 160 increases.
pod 9055 4 937 (5) The sieve tubes are very fine, not more than 30 um in
roots 842 2 700 diameter. This is comparable with a very fine human
hair. At regular intervals the tubes are spanned by sieve
* Corrected for background radiation which is always present as a result
of cosmic radiation. plates with pores of even smaller diameter. The smaller
the diameter of the tubes and pores, the greater is their
14.30 (a) From table 14.8 calculate the resistance to the passage of fluid, and the greater the
percentage distribution of radioactivity in each plant force required to move it.
and thereby compare directly the patterns of “C-
assimilate distribution in the two plants. What are the 14.32 How many sieve plates per metre
main similarities and differences in the pattern of '*C would be encountered by a sucrose molecule moving
export from the upper and lower leaves? through a sieve tube whose sieve elements were
(b) Consider the significance of these similarities and 400 um long?
differences in relation to the growth of the plant and
its parts. (It may help you if you draw a simple (6) Apart from sieve plates, sieve tubes have other
diagram of a pea plant with about eight leaves, with a structural features which must be taken into account.
pod positioned by the third leaf from the top, then —
indicate the direction and degree of assimilate —
movement from both an upper and lower leaf tothe
pod and to the root system.) a
14.8.1 Features of phloem translocation

Although an adequate hypothesis for xylem
translocation has been established, there is still controversy
about the mechanism of phloem translocation. Before
considering possible mechanisms for phloem translocation
it is useful to list some outstanding facts which any oe cell wall
hypothesis has to account for, and which make the problem . Pip so) middle. 7. >
such a difficult one to solve. lamella
(1) The quantity of material moved can be very large. It is
estimated, for example, that as much as 250 kg of sugar eon e ze
can be conducted down the trunk of a large tree during e, wD & pat Re Pig .. callose | ee 2
a growing season.
(2) The rate of flow is high, commonly 20-100 cm h"'. middle lamella
of sieve plate
Maximum rates in excess of 600 cm h | have been
recorded.
sieve tube .
14.31 If sucrose were moving at
100 cmh"! through sieve tubes where sieve ele-
ments were 200 um long, how long would ittake a Key
CC companion cell
given molecule of sucrose to pass through one sieve P_ sieve pore
element? PI plastid
SP sieve plate
a)
Translocation values of 10-25 g dry mass h7! cm~” Pp phloem protein
of sieve tube cross-sectional area are commonly re . a
obtained for dicotyledonous stems. Fig 14.22 Electron micrograph of a mature sieve element
500
STEVE TUBE ELEMENT or SIEVE ELEMENT
when mature no nucleus, no ribosomes, no Golgi
apparatus, small mitochondria, little cytoplasm;
diameter 10-50 pm, length 150-1 000 um
mitochondrion
small and few present
smooth endoplasmic cell wall
reticulum
% Ley ys /
sieve area small vacuole
cytoplasm thin peripheral layer, free occasionally / / /
bounded by a plasma membrane 4h yh present in the side @Y free ribosomes
Just inside cell wall walls, similar, / @) throughout cytoplasm
to sieve plate / Y
cell wall
Y Y Golgi apparatus
plastid hts oes

A ribosomes
starch grain
rough endoplasmic
reticulum
endoplasmic reticulum
typically stacked
plasmodesma -—large
nucleus
phloem protein may or may not be

present in sieve pores in life small vacuole
sieve pore
mitochondrion
callose
a polymer of glucose
COMPANION CELL
plasma membrane dense cytoplasm,
numerous ribosomes
cellulose cell wall and mitochondria,
middle lamella very active
phloem protein
Wis,
Fig 14.23 Diagrammatic LS of sieve tube elements and a companion cell as seen with the electron microscope. If the sieve
tube is damaged, for example by a grazing animal, more callose is rapidly deposited, blocking the sieve plate and preventing
loss of valuable solutes from the sieve tube
14.8.2 Ultrastructure of sieve tubes

develop into sieve plates. These are formed when the
In contrast with xylem vessels, which are dead plasmodesmata of the end walls enlarge greatly to form
empty tubes with few, if any, internal obstructions, phloem sieve pores. A surface view of a sieve plate is shown in
sieve tubes are living and do apparently contain obstruc- fig 8.12. The effect of all the changes is to leave a tube-like
tions to the flow of solution, namely the sieve plates and, structure with a wide lumen and a very narrow, indistinct,
to a lesser extent, the cytoplasm. Since the mechanism of peripheral layer of living cytoplasm bounded by a plasma
movement is still unclear, it is important in the search for membrane.
evidence to study the structure of sieve tubes in more detail Closely associated with each sieve element are one or
than can be revealed by the light microscope. Fig 14.22 more companion cells, parenchyma cells which are derived
is an electron micrograph of a mature sieve element, and from the same parent cell as the neighbouring sieve
fig 14.23 is a diagram showing the main features of sieve element. Companion cells have dense cytoplasm with small
elements and their neighbouring companion cells. vacuoles, and the usual cell organelles. They are metaboli-
During development of a sieve element from a meriste- cally very active, as indicated by their numerous mitochon-
matic cell its nucleus degenerates, making it an unusual dria and ribosomes (fig 14.23). They show a very close
example ofa living cell with no nucleus; in this respect it is structural and physiological relationship with sieve ele-
like mammalian red blood cells. At the same time many ments, being essential for their survival because when
other profound changes take place, the results of which are companion cells die, so do sieve elements.
shown in fig 14.23. The cell walls at each end of the element In dicotyledonous and some monocotyledonous plants
501
sieve elements develop large quantities of a fibrous protein sections from plants fed with “CO, revealed radioactivity
called phloem protein (P-protein). This sometimes forms in the phloem. The introduction of radioactive tracers in
deposits large enough to be seen with the light microscope. the 1930s and 1940s provided a tremendous boost to work
Such deposits were once called ‘slime bodies’ or ‘slime on translocation.
plugs’, but the material is not carbohydrate in nature and Confirmation that movement is through the sieve tubes
therefore not a true mucilaginous slime. There is much comes from a neat type of experiment in which the ability
debate as to whether or not the fibres of protein are of aphids to feed on translocating sugars is made use of.
normally present in the sieve pores, where they are The aphid penetrates the plant tissues with its specially
sometimes, but not always, seen in the electron micro- modified mouthparts; these include extremely fine, tube-
scope. One of the great problems of preparing phloem like ‘stylets’ which are pushed slowly through the plant’s
tissue for electron microscopy is that the contents of sieve tissues to the phloem. They can be shown to penetrate
tubes are thought to be under high hydrostatic pressure, individual sieve tubes, as revealed by fig 14.25.
possibly as great as 3 000 kPa. Cutting a specimen for If the aphid is anaesthetised with carbon dioxide and the
fixation might therefore release this pressure and the body removed, leaving the stylets in the plant, the contents
sudden surge of sieve tube contents might result in phloem of the sieve tube will continue to be forced up the tube of
protein and the other contents being swept into and the mouthparts by the hydrostatic pressure in the sieve
plugging the sieve plates. Any phloem protein seen in the tube, and the oozing fluid can be collected by microcapil-
sieve pores might therefore be due to a surge artefact. A lary tubes. This technique has found a number of useful
number of attempts have been made to get round this applications, for example in estimating rate of flow through
problem. For example, wilting a plant before cutting it sieve tubes (rate of exudation from tube) and in analysing
should eliminate or reduce hydrostatic pressure. Electron their contents.
microscopy of wilted plants reveals the sieve pores are Finally, improvements in the sensitivity of film used in
sometimes plugged, sometimes unplugged and sometimes microautoradiography have enabled precise location of the
partially plugged. This and other techniques have so far weakly emitting isotope of hydrogen, tritium (*H), in sieve
failed to resolve the problem. tubes rather than other phloem cells. The isotope is
supplied as part of an amino acid or sucrose.
Experiments have also established that different mate-
14.8.3 Evidence for movement in phloem rials are carried up and down the phloem at the same time,
It is important, especially in view of the although it is probable that this bidirectional movement is
discussion so far, to be certain that organic solutes really in neighbouring sieve tubes rather than in the same sieve
are carried in the phloem sieve tubes. The earliest evidence tube.
for movement of sugars and other compounds in phloem
came from ringing experiments, in which aring of tissue
14.8.4 Mechanism of translocation in phloem
containing phloem was removed from the outer region of
the stem, leaving the xylem intact. Malpighi obtained The facts which any hypothesis must account
evidence in 1675 for ascent of water in wood and descent of for are summarised in section 14.8.1. A picture of active
food in ‘bark’. He removed rings of bark from trees (bark phloem has been presented in the previous sections in
contains the phloem) and found that the leaves did not wilt, which large quantities of material move at relatively rapid
but that growth below the ring was greatly reduced. speeds through very fine sieve tubes. Within the tubes are
Mason and Maskell, working with cotton plants in apparent obstructions, the sieve plates, and other structu-
Trinidad during the 1920s and 1930s, did many ringing ral features such as phloem protein for which no certain
experiments, one of which is described in fig 14.24. From roles have been found. Combine these facts with the
the results of the experiments shown in fig 14.24, Mason further fact that the system is delicate and easily damaged
and Maskell concluded that some lateral exchange of by interference, and it is not surprising that research
sugars can take place between xylem and phloem when workers have found it difficult to establish the mechanism
they are in contact and the phloem is interrupted (fig of translocation through sieve tubes.
12.24a) but that downward movement occurs in phloem (b Most workers now believe that a mass flow of solution
and c). occurs through sieve tubes, involving the bulk movement
Two simple types of experiment have been done to show of water and solutes in the same direction, unlike diffusion
the movement of sucrose in phloem. In 1945 a non- in which molecules and ions move independently of each
radioactive isotope of carbon, °C, was introduced into a other. Diffusion itself is too slow to account for the rates
plant as CO, and detected by mass spectrometry. A ring observed. The evidence for mass flow through sieve tubes is
of phloem was killed with a fine jet of steam and summarised below.
translocation of C-labelled sucrose through this section (1) When phloem is cut, sap can be induced to exude,
was shown to be prevented. Movement of mineral apparently by mass flow. This is sometimes utilised
elements in the xylem is not affected by such treatment. In commercially as a source of sugar. For example the
the second experiment, microautoradiography of stem sugar palm exudes 10 dm’ of sugar-rich sap per day.
502
METHODS one of four flaps, flaps cut as in (a), F
flaps tied back into vlc Peaiaeck as (a) but ringed at
place as shown in before tying back both ends of cuts
(b) with no vaseline flaps, therefore
xylem and phloem
16.5 cm cannot exchange
cut materials
~~. a
1.3cm
ring
(a) (b) (c)

RESULTS Carbohydrate found Carbohydrate found Little carbohydrate
in xylem and phloem mainly in phloem found in xylem or phloem
Fig 14.24 Ringing experiments on cotton plants carried out by Mason and Maskell
of an aphid inserted into

Fig 14.25 (a) An aphid with its feeding stylets inserted through a leaf epidermis. (b) Feeding stylets
a sieve tube
503
(2) The prolonged exudation of sucrose solutions from solutes are constantly being used at the sinks (C) and made
aphid stylets, as described in section 14.8.8, is evidence at the sources (A).
of hydrostatic pressure in the sieve tubes. The Miinch hypothesis is a purely physical explanation
(3) Certain viruses are moved in the phloem translocation and so does not explain why sieve tubes must be living and
stream, indicating mass flow rather than diffusion since metabolically active. It also does not explain the observa-
the virus is passive and not in solution. tion that leaf cells are capable of loading sieve tubes against
The main debate concerns the mechanism by which mass a concentration gradient, that is the fact that the OP of
flow is brought about, and the major hypotheses will be sieve tubes is greater than that of the leaf cells. The
discussed. hypothesis has therefore been modified to include an active
loading mechanism of solutes into the sieve tubes. The
Minch’s hypothesis or pressure flow osmotic and hydrostatic pressure gradient therefore starts
hypothesis in the tubes rather than in the photosynthetic cells. It is also
In 1930, Miinch put forward a purely physical hypothesis to believed that unloading at the sinks is an active process.
explain how mass flow might be brought about in sieve Evidence for Minch’s hypothesis
tubes. It can be illustrated by the model shown in fig 14.26.
In the model there is an initial tendency for water to pass (1) The hypothesis predicts that mass flow will occur
by osmosis into A and C, but the tendency is greater for A through sieve tubes and this seems to occur (the
because the solution in A is more concentrated than that in evidence is given above).
C. As water enters A, a turgor pressure (hydrostatic (2) The hypothesis requires the existence of an osmotic
pressure) builds up in the closed system A-—B-C, forcing gradient and high turgor (hydrostatic) pressure in the
water out of C. Mass flow of solution occurs through B phloem. These have been demonstrated in a number of
along the hydrostatic pressure gradient so generated. plants.
There is also an osmotic gradient from A to C. Eventually Evidence against Munch’s hypothesis
the system comes into equilibrium as water dilutes the
contents of A and solutes accumulate at C. The model can (1) The hydrostatic pressure gradients required to move
be applied to living plants. The leaves which make sugar solutes at the observed rates are relatively high.
during photosynthesis, thus raising the OP of the leaf cells, Assuming the sieve pores are completely open, a
are represented by A. Water, brought to the leaf in xylem gradient of 100 kPa m“' might be adequate, but it is
(D), enters the leaf cells by osmosis, raising their turgor still doubtful if such large gradients exist. If the sieve
pressure. At the same time, sugars are used in the sinks, tube pores are obstructed, such as with cytoplasm or
such as roots (C), for various purposes including respira- phloem protein, the required pressures become un-
tion and synthesis of cellulose. This lowers the OP of these realistic.
cells. A hydrostatic pressure gradient exists from leaves to (2) The Minch hypothesis does not explain the occurrence
roots, or, in more general terms, from sources to sinks, of sieve plates or phloem protein, both of which would
resulting in mass flow. Equilibrium is not reached because seem to be barriers to mass flow.
==—) mass flowof solution a manometer attached to B would

register a hydrostatic pressure
———+» movement of water
semi-permeable membrane semi-permeable membrane
solution of high OP, e.g. solution of low OP, e.g.

concentrated sucrose dilute sucrose solution
solution
water
Fig 14.26 Physical mode! to illustrate Munch's mass flow hypothesis of phloem translocation
Equivalents in living plant
A: source, such as leaf; B: phloem; C: sink, such as roots, meristems, fruits; D: xylem, cell walls and intercellular spaces
504
Electroosmosis (4) A potential difference (PD) builds up across the sieve
A mechanism involving electroosmosis was proposed plate because anions begin to accumulate above it. The
independently by Fensom in 1957 and Spanner in 1958. The net charge is negative above the plate and positive
phenomenon of electroosmosis is illustrated in fig 14.27. below it.
It depends upon the presence of an ionic solution in (5) Once acritical PD is reached, a local surge of H™ ions
which a fixed charged surface occurs. The surface becomes (protons) occurs from the wall of the sieve tube,
charged when the molecules of which it is composed lose through its plasma membrane and into the tube,
ions. For example, if hydrogen ions leave an insoluble reducing pH (increasing H* concentration).
molecule, such as a fibrous protein, when it is placed in (6) This more than balances the negative charge and
water, the positively charged hydrogen ions can move reverses the PD across the plate, temporarily making
freely in solution leaving, behind a negatively charged, the local environment above the plate positively
immobile molecule. The solution then carries a mobile charged.
positive charge which balances the fixed negative charge. If (7) This positive charge causes electroosmosis of K* ions
a potential difference is applied across the charged surface, through the sieve plate, past the fixed negative charge
the positive ions move towards the negative electrode of the P-protein. Movement of potassium and other
(cathode). Each ion is surrounded by a shell of water cations induces a mass flow of solution.
molecules (fig 5.5) which moves with the ions, together (8) A proton pump in the sieve tube membrane quickly
with other ions and uncharged molecules, such as sucrose, restores the high pH of the sieve tube by actively
in a mass flow of the whole solution past the fixed charge pumping out protons from the sieve tube. Energy for
(fig 14.27). Y this is supplied by ATP from the companion cells and
sieve tubes.
cathode anode
(9) The original PD begins to build up again.
The overall result is that translocation through sieve
tubes is by mass flow, as proposed by Minch, but is boosted
by electroosmosis at the sieve plates. The boosting is in the
form of brief pulses. It does not necessarily take place
original level of solution
simultaneously at all sieve plates and there may or may not
mass flow of be a particular sequence of ‘firing’ at different sieve plates.
solution
A pulsed, rather than smooth, flow may therefore occur
ionic solution through sieve tubes.
Evidence for the electroosmosis hypothesis

(1) A role for companion cells, in supplying ATP, is
established.
porous membrane or fine tubes or macromolecules;
they are fixed in position and have charged surfaces (2) High levels of potassium ions are found in sieve tubes.
(3) The hydrostatic pressures required are lower than
Fig 14.27 Electroosmosis. In the case illustrated, the fixed
those for Mtinch’s hypothesis.
surface is negatively charged and the solution carries a
corresponding net positive charge (4) A role is given for sieve plates: they provide a charged
porous structure allowing the flow of solutes by
In sieve tubes, fixed charged surfaces could be provided electroosmosis. There is also a possible role for phloem
by the cell walls, including the sieve plates, and the phloem protein, if this is found in the sieve pores.
protein (P-protein), all of which would be negatively Evidence against the electroosmosis hypo-
charged under the alkaline conditions usually found in thesis. The hypothesis is largely circumstantial, that is
phloem. Spanner’s hypothesis has been modified several there is no direct evidence for it. For example, the
times, but his ‘final restatement’, published in 1979,* can postulated changes in potential difference have not been
be outlined as follows. measured. Also, it has not been demonstrated that the
(1) Mass flow is initiated by the loading of sieve tubes and sieve tube walls act as a reservoir for hydrogen ions.
operates as described by Munch.
(2) P-protein is loosely packed in the sieve pores, possibly The relay hypothesis
as a result of mass flow. A major criticism of Spanner’s hypothesis is that massive
(3) There is a high concentration of potassium ions (K*) in expenditures of metabolic energy would be needed to
the sieve tube solution. These and other cations can maintain the observed flow rates, since pumping takes
pass through the sieve plate, but anions are repelled by place over the relatively small area of the sieve plates. In
the negative charge of the phloem protein (and cell 1979, Alexander Lang suggested that pumping could occur
walls). over a much larger surface area if it took place across the
t, 2, 107-21. membranes of adjacent, overlapping sieve tubes in the
* D. C. Spanner (1979) Plant, Cell and Environmen
505
region of the overlap. The pumping process could be complex, extensive plasmodesmata. The numerous
similar to that which is involved in loading and unloading mitochondria in their cytoplasm provide energy for this.
sieve tubes at source and sink. Movement of solutes would These modified cells are called transfer cells, and similar
still occur along a pressure gradient between source and examples of plant cells with wall ingrowths have since been
sink, as postulated by Miinch, but a much smaller overall found in many situations where short-distance transport
gradient would be required. This ‘relay mechansim’ is occurs, including xylem parenchyma. Similar cells can
therefore a mass flow mechanism which reconciles some of easily be seen in a toluidine blue-stained, hand-cut section
the conflicting evidence for and against Miinch’s hypo- of a Tradescantia node. They are not found in all plants, but
thesis. It is supported by the observation that solutes move are common in the pea family and some other families.
readily in and out of sieve tubes along their lengths, such as Fig 14.28 shows their appearance when viewed with an
between phloem and xylem. electron microscope.
Other hypotheses
It should be noted that other hypotheses for phloem
transport have been put forward, including some form of
cytoplasmic streaming, and the existence of cytoplasmic
strands (‘transcellular strands’) supported by the sieve
plates and extending long distances through the sieve tubes
suggests that solutions could in some way be pumped along
the strands. Like electroosmosis, these ideas are interest-
ing but as yet are supported by little, if any, experimental
evidence.
14.8.5 First-aid mechanisms — a possible

role for sieve plates, phloem protein
and plastids
One danger faced by all plants is damage from
being eaten by animals. If sieve tubes are ruptured, leakage
of high energy substances such as sucrose would be costly to
cell wall
the plant. Usually damaged sieve tubes are sealed within
protuberances chloroplasts
minutes by deposition of callose across the sieve plates, of transfer cell
blocking the sieve pores. This represents a possible role for
sieve plates. It has been suggested that in dicotyledons, the —~ plasmodesma
xylem vessels
phloem protein serves an even more rapid first-aid function
by blocking the sieve pores as soon as the tube is broken.
This is due to the pressure surge mentioned in section
14.8.2. In monocotyledons, where phloem protein is
Arrows indicate
usually scarce, the same role may be performed by plastids | . possible flaw of
found in sieve tubes. © solutes between xylem
and phloem
14.8.6 Loading sieve tubes

It has been shown that the sucrose concentra-
tion in sieve tubes in leaves is commonly between 10-30%,
Fig 14.28 Transverse section of a minor vein in a leaf of
whereas it forms only about a 0.5% solution in the Senecio vulgaris. The phloem region, in the top half of the
photosynthetic cells where it is produced. Loading of sieve picture, shows six parenchyma cells centred around two sieve
tubes therefore occurs against a concentration gradient. elements. Pairs of companion cells modified as A-type
Both symplast and apoplast routes are possible in the short transfer cells lie both above and below the sieve elements.
journey, 3 mm at most, from chloroplasts to sieve tubes. There is a B-type transfer cell to each side. The A- and the
B-types differ strikingly in the relative density of their
In 1968 a modified type of companion cell was reported cytoplasm, in the polarity of their zones of wall ingrowths, and
by Gunning and fellow workers. It has numerous internal in their connections with the sieve elements: plasmodesmata
protuberances of the cell wall, a result of extra thickening. between sieve elements and A-types are common, but are
This in turn results in an approximately tenfold increase in very rare between sieve elements and B-types. In the lower
surface area of the plasma membrane lining the wall. It is half of the picture two xylem elements occupy the centre,
while to each side there are parts of two large bundle sheath
thought that such cells are thus modified for active uptake cells. Arrows indicate some of the possible routes for intra-
of solutes from neighbouring photosynthetic cells and that veinal retrieval of solutes delivered to the apoplast in the
they actively load their adjacent sieve elements through xylem (see text). Magnification x 6 560
506
Transport in animals contact with the body tissues. Distribution of blood to the
tissues is able to be adjusted. The only entry and exit to this
system is through the walls of the blood vessels.
In the animal kingdom, the protozoa, coelenterates and Blood vessels are named according to their structure and
platyhelminths lack a specific system for the transport and function. Vessels conveying blood away from the heart are
distribution of materials. The organisms in these phyla called arteries. These branch into smaller arteries called
possess a large surface area to volume ratio, and diffusion arterioles. The arterioles divide many times into micro-
of gases over the whole body surface is sufficient for their scopic capillaries which are located between the cells of
needs. Internally, the distance that materials have to travel nearly all the body tissues. It is here that exchange of
is again small enough for them to move by diffusion or materials between blood and tissues takes place.
cytoplasmic streaming (section 7.2.4). Within the organ or tissue the capillaries reunite forming
As organisms increase in size and complexity so the venules which begin the process of returning blood to the
quantity of materials moving in and out of the body heart. The venules join to form veins. It is these blood
increases. The distance that materials have to travel within vessels that actually pass the blood back into the heart. The
the body also increases, so that diffusion becomes anatomy of each type of blood vessel is discussed in detail
inadequate as a means for their distribution. Some other later in section 14.12.
method of conveying materials from one part of the
organism to another is therefore necessary. This generally
takes the form of a mass flow system. 14.10 The development of transport
systems in animals
14.9 General characteristics of a In the protozoa circulation of materials is

circulatory system primarily achieved by protoplasmic streaming. Coelenter-
ates rely on movements of their body wall to create water
The purpose of a circulatory system is to currents in the enteron which circulate food, water and
provide rapid mass flow of materials from one part of the dissolved gases. The musculo-epithelial and flagellate cells
body to another over distances where diffusion would be of the endoderm assist in this activity. Platyhelminths have
too slow. On reaching their destination the materials must a very thin, flattened shape, enabling materials to be
be able to pass through the walls of the circulatory system exchanged within the organism and with its environment by
into the organs or tissues. Likewise, materials produced by diffusion.
these structures must also be able to enter the circulatory
14.10.1 Annelids
system.
Every circulatory system possesses three distinct charac- Annelids are coelomate animals. The pres-
teristics: ence of a coelom separates the body wall from the internal
(1) a circulatory fluid, generally called blood, organs and confers the advantage of independence of
(2) a contractile, pumping device to propel the fluid movement of internal structures such as the gut. However,
around the body, this may either be a modified blood this is countered by the need for some form of connecting
vessel or a heart; system between the two regions which enable food, gases
(3) tubes through which the fluid can circulate, called and waste substances to be transported between the
blood vessels. environment and the gut or vice versa. It is at the
Two distinct types of circulatory system are found in the coelomate level that a blood system evolved in order to
higher invertebrates and vertebrates. They are the open connect gut and body wall.
and closed vascular systems.
The earthworm — a closed blood vascular
The open vascular system (most arthro- system
pods, some cephalopod molluscs, tunicates). Blood is “There is a well-developed blood system in which blood
pumped by the heart into an aorta which branches into a circulates around the body through a system of closed
number of arteries. These open intoa series of blood spaces blood vessels. The largest blood vessel is the longitudinal,
collectively called the haemocoel. Blood under low pres- dorsal vessel which has muscular walls. It is situated above
sure moves slowly between the tissues, gradually percolat- the alimentary canal. Peristaltic contractions originate at
ing back into the heart via open-ended veins. Distribution the rear end of the vessel and drive blood forwards towards
of blood to the tissues is poorly controlled. the anterior end of the animal, backflow being prevented
The closed vascular system (echinoderms, by a series of valves. Each valve is formed from a fold of
cephalopod molluscs, annelids, vertebrates). Blood is endothelium within the blood vessel. The dorsal vessel is
pumped by the heart rapidly around the body under the main collecting vessel and receives blood from the body
sustained high pressure and back to the heart. It is confined wall, gut, nerve cord and nephridia. The names of the main
to aseries of specific vessels and not permitted to come into segmental vessels are given in figs 14.29 and 14.30.
507
Fig 14.29 (right) Segmental COLLECTION DISTRIBUTION
distribution of blood in Lumbricus in
regions behind the clitellum. (From
A. E. Vines & N. Rees (4th SS eee vessel
edition, 1972) Plant and animal
efferent gut
biology, Vol. |, Pitman) vessel afferent body
wall vessel
commissural
vessel
afferent gut
efferent nephridial vessel
vessel
afferent nephridial
Fig 14.30 (below) Main blood efferent body vessel
vessels in the earthworm wall vessel
ventral vessel
(Lumbricus terrestris). (From E. G.
Springthorpe (1973) An introduction
to functional systems in animals, aubneural lateral neural vessel
Longman) vessel
dorsal five pairs of commissural ing no haemoglobin, and serves to transport dissolved
‘worm hearts’
vessel vessel
foodstuffs and excretory materials and to circulate colour-
less amoeboid leucocytes.
The cockroach — an open blood vascular
system
There is only one blood vessel in the cockroach, the dorsal
blood vessel, and the posterior part of it is modified into a
distinct heart, whilst the anterior end is called the aorta.
ventral subneural
vessel vessel The heart lies in a large sinus which is a modified part of the
haemocoel called the pericardium (fig 14.31). There are 13
dilations or chambers in the heart, three are located in the
The dorsal vessel connects with the smaller, more thoracic segments and ten in the abdomen. Every chamber
contractile, ventral vessel via five pairs of contractile except the most posterior possesses a pair of lateral
‘pseudohearts’ located in segments 7-11. Each ‘pseudo- openings called ostia. Each ostium is fitted with a valve
heart’ possesses four valves which permit the blood to flow which permits blood to enter but not leave the chamber.
only towards the ventral vessel. Between adjacent chambers are more valves which prevent
The blood flows posteriorly in the ventral vessel. It is backward flow of the blood. Intersegmental alary muscles
distributed to the nephridia, nerve cord, gut and body wall are attached to the ventral wall of the heart, and when
by aseries of segmental blood vessels. In addition the nerve these muscles contract they increase the volume of the
cord is supplied with blood by the longitudinal, subneural heart and create a negative pressure. This draws in blood
vessel which runs below it. from the pericardium through the ostia. When the alary
Within the various organs of the worm, capillary muscles relax and the heart contracts, blood is propelled
networks enable materials to be exchanged between blood forwards into the aorta. This divides into several arteries
and tissues. Eventually the blood passes back to the dorsal which are open-ended. Consequently blood pours out of
vessel, when it can once again begin its journey through the them into blood sinuses where it bathes the organs of the
closed blood vascular system of the worm. insect directly. Eventually the blood percolates back into
The blood itself is red in colour, containing haemoglobin the pericardium and ultimately into the heart.
(section 14.13.1) dissolved in the plasma. It transports
oxygen, carbon dioxide, soluble excretory materials and heart ligament pericardial
foodstuffs around the body. Colourless amoeboid cells cavity
which have a defensive function also circulate in the blood. alary

muscles
pericardial
14.10.2 Arthropoda membrane gut
The coelom is drastically reduced in the

arthropods and its place taken by the haemocoel. This is a direction of
= ‘aVp
je blood flow
network of blood-filled spaces called sinuses in which the
internal organs are suspended. Gaseous exchange in most
Fig 14.31 Schematic transverse section to show open blood
arthropods is effected by the tracheal system (section 4.10),
vascular system of an insect. (After Ramsay (1968) A
and the blood vascular system is not used for transporting physiological approach to lower animals, Cambridge
respiratory gases. Arthropod blood is colourless, contain- University Press)
508
14.10.3 Vertebrate circulatory systems pressure developed in the surrounding pericardium. It is
All vertebrate circulatory systems possess a passed on to the thicker, more muscular ventricle. When
well-defined muscular heart, lying in a ventral position in this contracts, blood is pumped via the conus arteriosus into
the region of the pectoral girdle. The heart is responsible a ventral aorta. Valves between the atrium and ventricle
for conveying blood rapidly to all parts of the animal’s and the ventricle and conus arteriosus prevent backflow of
body. Arteries convey blood away from the heart, whilst blood. The conus arteriosus is highly elastic and easily
veins pass blood from the body to the heart. Another stretched by the pressure of the blood leaving the ventricle.
system, the lymphatic system, is also present and supple- In turn, its elastic walls act on the blood and force it
ments the activities of the circulatory system. forwards at a more uniform speed, thus enhancing a
Comparative embryological studies indicate that, in all continuous flow of blood in the ventral aorta.
vertebrate embryos, six lateral arterial arches emanate In the dogfish five pairs of afferent branchial arteries
from the ventral aorta and unite to form apair of lateral branch from the ventral aorta and carry deoxygenated
dorsal aortae which eventually merge into a single median blood to the gills (fig 14.33a). Within the gills the arteries
dorsal aorta (fig 14.32). Fish are the only vertebrates to divide many times into numerous fine capillaries. Even-
exhibit this arrangement in anything like its original form in tually they rejoin to form efferent branchial arteries. These
carry oxygenated blood and form aseries of four loops
the adult condition. In all other vertebrates the embryonic
which encircle the perimeter of each of the first four
pattern has been extensively modified.
internal branchial clefts. A single blood vessel from the
anterior part of the fifth branchial cleft opens into the
fourth loop. Four epibranchial vessels from either side of
the body convey the blood from the loops backwards and
(a) towards the midline where they join to form the dorsal
aorta. Anteriorly the dorsal aorta divides and joins the
internal carotid arteries, and supplies oxygenated blood to
the head. Posteriorly the dorsal aorta branches many times
ventral aorta
into arteries which supply the rest of the body with
NS ets aorta
oxygenated blood.
As blood flows through the heart only once during each
circuit of the body, the dogfish is said to possess a single
circulation (fig 14.33c). The blood pressure falls as it passes
dorsal aorta
through the gill capillaries and is low in the dorsal aorta.
The blood pressure falls even further after it has passed
atrio-ventricular ¢ sinu-atrial
atrium valves through the capillaries of the body organs. This is because
valves
conus ia of the resistance to its flow exerted by the various capillary
arteriosus N networks of the body. Consequently, return of blood to the
eee heart is very slow. To offset this apparent disadvantage,
ventral duct of large blood spaces called sinuses, which provide little
x» Cuvier
aorta + resistance to blood flow, are present. Raised anterior and
(b) posterior cardinal sinuses, together with laterally placed
sinus veins, return blood to the region of the heart where it
ventricle venosus
passes to the sinus via a pair of Cuvierian veins. Backflow
Fig 14.32 (a) Vertebrate embryo condition of the blood of blood in the veins is prevented by valves, and forward
vascular system. Six arterial arches branch from the ventral — flow enhanced by the body muscles squeezing these vessels.
aorta, eventually rejoining to form lateral aortae. (b) Embryonic In the posterior region of the fish there are blood vessels
heart called portal veins. These are blood vessels with capillary
beds at both ends. Blood from the small intestine passes to
the liver via the hepatic portal vein, and blood from the tail
to the kidney via renal portal veins. Thus the blood of the
dogfish having already passed through the capillaries of the
gills passes through two sets of capillaries of the portal
The dogfish systems — three sets of capillaries in all.
Dogfish blood contains oval-shaped, nucleated red
The dogfish heart (fig 14.33b), which is confined within a
blood cells and amoeboid leucocytes suspended in plasma.
pericardial cavity, is S-shaped and consists of two main
an atrium and a ventricle. Blood from the The lymphatic system is comprised of lymphatic vessels
chambers,
emptying into two dorsal longitudinal collecting ducts
general body circulation enters a thin-walled chamber, the
which finally return their contents (that is lymph) to the
sinus venosus, which precedes the heart. From here it
blood via the cardinal sinuses.
passes into the weakly muscular atrium by a negative
509
epibranchial artery
efferent branchial artery
afferent branchial artery
ventral aorta
dorsal aorta
(a) arterial circulation
sinu-atrial valve
atrium
Ss sinus venosus
atrio-ventricular
valve
Ss
ventricle
pericardium
conus arteriosus
(b) heart — two main chambers, one (c) general circulation

atrium and one ventricle — single circulation
Fig 14.33 Dogfish arterial blood supply. Six arterial arches are reduced to five, the first gill slit is modified as a spiracle. Direct
branchial vessels have split up to form afferent, efferent and epibranchial arteries. This gives a greater surface area of the
blood system in the gill region for gaseous exchange
Amphibia — the frog blood in the atria mix. However, both atria contract at the
In the adult frog the heart and aortic circulation show same time and deliver their blood to the single ventricle.
modifications which can be correlated with its semi- The inner surface of the dorsal and ventral! walls of the
terrestrial mode of life. Lungs have developed and ventricle is built up into ridges which prevent mixing of the
replaced gills as respiratory organs, and a new circulation, blood to some extent. When the blood passes from the
the pulmonary circulation between the lungs and heart, has ventricle to the conus arteriosus, mixing is further
been established. There is no longer a need to retain all the prevented by a spiral valve which incompletely divides the
aortic arches to supply branchial arches, and some have conus into two distinct corridors. The dorsal passage,
been put to new uses whilst others have been lost. The third called the cavum pulmocutaneous, leads to the pulmonary
arch has become the carotid artery and transports blood arch which, besides running to the lungs, also sends
anteriorly to the head (fig 14.34@). The fourth is the branches to the skin, whilst the ventral cavum aorticum
systemic, conveying oxygenated blood to the rest of the takes blood to the carotid and systemic arches.
body via the dorsal aorta, whilst the sixth arch transports It is clear that there are two distinct circuits which the
deoxygenated blood from the heart to the lungs via the blood can take, one to the lungs and one to the body. This
pulmonary artery. The first, second and fifth arches have may be regarded as a partial double circulation, where
disappeared. blood, after passing from the heart to the lungs, is returned
The heart consists of three main chambers, two atria and again to the heart before passing to the body (fig 14.34c),
one ventricle (fig 14.34b). Blood which has been oxygen- The separation of oxygenated and deoxygenated blood in
ated at the lungs is returned to the left atrium via the this system is not entirely complete.
pulmonary vein, whilst deoxygenated blood from the body The blood vascular system of a frog is a very special
is passed to the right atrium via the sinus venosus by the system suitable for an amphibious animal. When the frog
anterior and posterior venae cavae. At no time does the is relatively inactive, oxygen diffuses into the capillaries
510
(a) arterial circulation
TADPOLE. FROG
head &
forelimbs
+
systemic ———
5 arch ______ oxygenated blood
—>
ue “= deoxygenated v
6 pul
__ pulmonary blood an
arch A
dorsal
aorta
first and second arterial arches three functional arches: 3, 4 and 6;

disappear; arches 3-6 supply gill capillaries short-circuited;
the gills with blood sixth arch supplies lungs
(b) heart — three-chambered, ——

pulmonary veins
two atria and one ventricle
anterior venae
cavae
right atrium -
__posterior vena
left atrium cava
sinus venosus
ventral
aorta
carotid
arch
iN “a (c) general circulation

posterior
arch ree | — partial double
trunk
circulation
pulmonary
arch conus
arteriosus ventricle
Fig 14.34 Blood system of a frog
under the skin and is transported to the right atrium of the complete separation of oxygenated blood in the left side
heart via the subclavian vein. The lungs are not used except from deoxygenated blood on the right side. The conus
under conditions of strenuous action. Thus normally the arteriosus has been replaced by three distinct arteries
left atrium receives little oxygenated blood. which form arches directly from the ventricle. They are the
Veins replace sinuses in the venous system of the frog. right systemic arch, which gives rise to the carotid arteries,
Blood is returned to the sinus venosus by anterior and the left systemic arch, and the pulmonary arch.
posterior venae cavae which receive blood from veins
leaving the major organs of the body. Renal and hepatic
portal veins are also present.
Intercellular channels containing lymph (section 14.12.1)
drain into lymphatic vessels. Eventually the lymph is
pumped into the venous system at two points, one anterior ventral aorta
and one posterior. A pair of small contractile lymph hearts
in each region drive the lymph into the veins. right systemic
arch
Frog’s blood consists of nucleated erythrocytes contain- left systemic
ing haemoglobin, and a variety of white cells suspended in arch
carries
the plasma. They include macrophages, lymphocytes, deoxygenated
granulocytes and monocytes (section 14.11.2). blood
pulmonary arch
Reptilia — the crocodile
Here the heart consists of four chambers, two atria and two
ventricles, the right side being more or less completely arterial circulation
separated from the left side (fig 14.35). Therefore the
Fig 14.35 Reptile arterial blood supply — the crocodile
crocodile possesses a double circulatory system with almost
511
Aves and Mammalia — general pulmonary arch. Within the venous system the posterior
vena cava replaces the renal portal system. Therefore
The heart is four-chambered in birds and mammals. The
blood from the posterior parts of the body is returned
right atrium and ventricle are completely separated from
directly to the heart, thus increasing the speed at which
the left atrium and ventricle (fig 14.36b). This means that
blood can be supplied back to the tissues.
oxygenated and deoxygenated blood are also kept comp-
letely separate. In order for blood in the right side of the
heart to get to the left side, it must first pass through the 14.11 Composition of mammalian blood
lungs (fig 14.36c). For blood from the left side to get to the
right side it has first to pass round the body. This double Blood is composed of cells bathed in a fluid
circulation ensures that oxygenated blood flowing to the matrix called plasma. The cells constitute about 45% by
tissues is under high pressure and not, as is the case of fish, volume of the blood, whilst the other 55% is represented by
under low pressure. As the blood passes through the lungs the plasma.
before it passes to the body, this ensures that it is well
oxygenated before it reaches the actively respiring organs. 14.11.1. Plasma
14.33 Why is it an advantage in birds and Plasma is a pale yellow liquid. It consists of
mammals for oxygenated blood to flow to the tissues 90% water and 10% of a variety of substances in solution
under high pressure? and suspension, some of which are normally maintained at
constant concentrations, whilst the concentrations of
The arterial system is further reduced in birds and others may fiuctuate within narrow limits according to their
mammals. Birds retain only the right half of the systemic rates of removal or supply from particular organs. The
arch, whilst mammals retain the left half (fig 14.36a). The major components of blood plasma together with their
third arch persists as the carotid arch and the sixth as the functions are shown in table 14.9.
(a)
arterial
circulation BIRD (b) (i) MAMMAL head &
forelimbs
oxygenated blood
carotid arch to
head and CEE == deoxygenated
forelimbs blood |
&
Nn
aD left systemic
pulmonary arch arch, blood to
to lungs body
right systemic pulmonary arch
arch, blood to to lungs
body
left systemic right systemic
arch disappears arch disappears
Z\
ee
pulmonary ee
right atrium :
\
NY V2
{ left atrium
ou
right ventricle left ventricle
(b) (ii) heart- four-chambered,

complete separation of (b) (iii) general circulation - posterior
right and left halves double circulation trunk
Fig 14.36 (a) Bird and (6) mammal arterial blood systems
512
Table 14.9 Components of blood plasma and their functions.
a —a se
Constituent Function
Ti
Constituents maintained at a constant concentration

Water Major constituent of lymph. Provides tissue cells with water. Conveys many dissolved
materials round the body. Aids maintenance of blood pressure and blood volume
Plasma proteins
Serum albumin Very abundant. Binds plasma calcium
Serum globulins:
a-globulin Binds thyroxine and bilirubin
B-globulin Binds iron, cholesterol and the vitamins A, D and K
y-globulin Binds antigens and is important in the body’s immunological reactions. Generally called
antibody. Also binds histamine
Prothrombin A catalytic agent which takes part in the blood-clotting process
Fibrinogen Takes part in blood-clotting process
Enzymes Participate in metabolic activities
Mineral ions
These include: All help collectively to regulate osmotic pressure and pH levels of the blood. They also exert
NaeaKe. Cay Mg’, rePOm, HPO, , a variety of other effects on the cells of the body; e.g. Ca" may act as a clotting factor,
PO act HCO:, so* regulate muscle and nerve cell sensitivity, influence the sol-gel condition within cells
Constituents that occur in varying concentrations

Dissolved products of digestion
Dissolved excretory products All are being constantly transported to and from cells within the body
Vitamins
Hormones
14.11.2 Blood cells In the adult, each erythrocyte has a life span of about
three months after which time it is destroyed in the spleen
Erythrocytes or liver. The protein portion of the erythrocyte is broken
These are the red blood cells. Characteristically in Man down into its constituent amino acids; the iron of the haem
they are small, enucleated and appear as circular biconcave portion is extracted and stored in the liver as ferritin (an
discs. Their average diameter is 7-8 um, which is approx- iron-containing protein). It may be re-used later in the
imately the same as the internal diameter of blood production of further erythrocytes or as a component of
capillaries. Their particular shape permits a larger surface cytochrome. The remainder of the haem molecule is
area to volume ratio than that of a sphere, and therefore broken down into two bile pigments, bilirubin and
increases the area which can be used for gaseous exchange. biliverdin. Both are ultimately excreted by way of the bile
Each cell is very thin, thus permitting efficient diffusion of into the gut.
gases from its surface inwards. Its membrane is pliable and Between 2-10 million erythrocytes are destroyed and
this property allows the erythrocyte to squeeze through replaced each second in the human body. The rate of
capillaries whose internal diameters are smaller than its destruction and replacement is determined by the amount
of oxygen in the atmosphere which is available for carriage
own.
The mechanism of production of erythrocytes is known by the blood. If the quantity carried is low, then the
marrow is stimulated to produce more erythrocytes than
as haemopoiesis and the tissue which gives rise to them is
the liver destroys. This is one of the ways in which
called haemopoietic tissue. In an infant, all bones contain
mammals acclimatise to the reduced oxygen content at high
haemopoietic tissue, whilst in adults the principal regions
altitudes. When the oxygen content is high, the situation is
of erythrocyte production are the bones of the pelvis, ribs,
reversed.
sternum, vertebrae, clavicles, scapulae and skull. There
are approximately five million erythrocytes per cubic mil-
White blood cells — leucocytes
limetre of blood. However this figure varies according to
the age, sex and state of health of each individual. These cells are larger than erythrocytes, and present in
An important characteristic of erythrocytes is the much smaller numbers, there being about 7 000 per cubic
presence of haemoglobin which combines reversibly with millimetre of blood. All are nucleated. They play an
oxygen to form oxyhaemoglobin in areas of high oxygen important role in the body’s defence mechanisms against
concentration, and releases the oxygen in regions of low disease. Although they are nucleated their life span in the
oxygen concentration. They also contain the enzyme bloodstream is normally only a few days. There are two
carbonic anhydrase which plays a role in carbon dioxide main groups of white blood cell, the granulocytes and the
transport (section 14.13.4). agranulocytes.
313
Table 14.10 Cellular components of blood (diagrams not drawn to scale).
Number o
Component Origin cells/mm Function Structure
Erythrocytes bone marrow 5 000 000 transport of oxygen

and some carbon
dioxide
Leucocytes bone marrow
(a) Granulocytes
(72% of total white blood cell
count) .
neutrophils (70%) 4900 engulf bacteria
eosinophils (1.5%) bone marrow 105 anti-histamine

properties
basophils (0.5%) 35 produce histamine

and heparin
(b) Agranulocytes (28%)
monocytes (4%) bone marrow 280 engulf bacteria d
lymphocytes (24%) bone marrow 1 680 production of antibodies

lymphoid tissue
spleen
Platelets bone marrow 250 000 initiate blood-clotting fe é

mechanism og
Basophils. They represent 0.5% of the white blood cell

Granulocytes (polymorphonuclear leuco- P ies -
population and produce heparin and histamine. The
cytes). These originate in the bone marrow but are
granules in these cells stain blue with basic dyes such as
produced by cells different from those that make erythro-
methylene blue.
cytes. Each cell contains a lobed nucleus and granular
cytoplasm (table 14.10). All are capable of amoeboid Agranulocytes (mononuclear leucocytes).
movement. Granulocytes can be further subdivided into These cells possess non-granular cytoplasm and either an
neutrophils, eosinophils and basophils. oval or bean-shaped nucleus. Two main types exist.
Neutrophils (phagocytes). These constitute 70% of the Monocytes (4%). These are formed in the bone marrow
total number of white cells. They are able to squeeze and have a bean-shaped nucleus. They are actively
between the cells of the capillary walls and enter the phagocytic and ingest bacteria and other large particles.
intercellular spaces. This process is called diapedesis. From They are able to migrate from the bloodstream to inflamed
here they move to infected areas of the body. They are areas of the body and act in the same manner. as
actively phagocytic and engulf and digest disease-causing neutrophils.
bacteria (section 14.13.5). Lymphocytes (24%). These are produced in the thymus
Eosinophils. They possess cytoplasmic granules which stain gland and lymphoid tissues from precursor cells which
red when the dye eosin is applied to them. Generally they originate from the bone marrow. The cells are rounded and
represent only 1.5% of the total number of white cells, but possess only a small quantity of cytoplasm. Amoeboid
their population increases in people with allergic condi- movement is limited. Their major function is to cause or
tions such as asthma or hayfever. It is thought that mediate immune reactions (such as antibody production,
eosinophils possess anti-histamine properties. The number graft rejection and tumour cell killing). The life span of
of eosinophils present in the bloodstream is under the these particular cells can vary from a matter of days in
control of hormones produced by the adrenal cortex. rodents up to ten years or more in humans.
514
14.11.3 Platelets (b)
Platelets are irregularly shaped membrane-
bound cell fragments, frequently enucleated and formed
direction of movement
from large bone marrow cells called megakaryocytes. They
function to initiate the mechanism of blood clotting. There
are about 250 000 platelets per cubic millimetre of blood.
microscope
slide
Experiment 14.7: Preparation of a blood smear

Materials
disposable towels Fig 14.37 (b) Preparation of a blood smear
cotton wool swabs
70% alcohol (8) Take another microscope slide, hold it at an angle of
Sterile disposable lancet 45° and touch the drop of blood with it. Then push the
two slides second slide along the first slide (fig 14.37b) in a
sealable container direction away from the drop of blood to make a blood
Leishman’s stain smear.
distilled water (9) Allow the smear to dry for several minutes so
glycerine permitting the cells to stick to the slide. This may be
cover-slip done by holding the slide above a bench lamp or
freshly prepared aqueous sodium chlorate solution waving it in the air.
(10) Add six drops of Leishman’s stain and leave for 60 s.
Method
(11) Add six drops of distilled water and mix with the stain
(1) Thoroughly wash both hands using soap and water. by rocking the slide.
Those giving blood samples must pay particular (12) Leave for 15 min and then rinse off the stain with
attention to washing the site chosen for sampling. Dry water.
the hands using only disposable towels. (13) Add a drop of glycerine to the preparation, followed by
(2) Shake the wrist of one hand downwards several times acover-slip, and view the smear under a microscope.
to force blood into the hand. (14) Erythrocytes will be stained pink, platelets and the
(3) Sterilise an area 5-10 mm from the lower corner of nuclei of white blood cells will be stained blue.
the nail of the little finger (fig 14.37a) using a cotton (15) Any blood spilt on the bench or elsewhere during the
wool swab that has been soaked in 70% alcohol and experiment must be wiped up immediately using a
allow it to dry. freshly prepared aqueous solution of sodium chlo-
(4) Remove a new sterile disposable lancet from its rate.
packet just prior to its use. DO NOT permit the sharp (16) Great care should be taken to avoid contamination of
end to touch anything. the skin with blood from another person. If this should
(5) Puncture the skin in the chosen area using the lancet occur, however, the contaminated area must be
and then place the lancet in a sealable container. cleaned immediately with sodium chlorate diluted
Lancets must be used once only. with water to ten times its volume, and then washed
(6) Transfer the drop of blood which appears onto one thoroughly with soap and water.
end of a sterile slide. Avoid any contact between the (17) At the end of the practical both hands must be washed
punctured skin and the slide. with soap and water and dried using disposable
(7) Using a fresh cotton wool swab, wipe the site of the towels.
puncture again with 70% alcohol and apply slight NB Because of the risk of contamination through
pressure if necessary to stop blood flow. Put all used broken skin, participation in this practical by people
swabs in the container with the lancets. with any kind of open wound should be avoided.
(a)
{ nail
@
Fig 14.37 (a) Point where blood may be taken from the
finger. It is easier to insert the lancet if the finger is crooked at
the top joint
515
Fig 14.38 (a) A blood smear showing red blood cells and three types of white cell. (b) A scanning electron micrograph of red
blood cells of a mammal
Table 14.11 Comparison in structure and function of an artery, capillary and vein (diagrams are not drawn to scale).
Artery Capillary ; Vein
Transport blood away from the heart Link arteries to veins. Site of exchange Transport blood towards the heart
of materials between blood and tissues
Tunica media thick and composed of No tunica media. Only tissue present is Tunica media relatively thin and only
elastic, muscular tissue squamous endothelium. No elastic fibres slightly muscular. Few elastic fibres
No semi-lunar valves No semi-lunar valves Semi-lunar valves at intervals along the
: length to prevent backflow of blood
Pressure of blood is high and pulsatile Pressure of blood falling and non- Pressure of blood low and non-pulsatile
pulsatile
Blood flow rapid Blood flow slowing Blood flow slow
Low blood volume High blood volume Increased blood volume
Blood oxygenated except in pulmonary Mixed oxygenated and deoxygenated Blood deoxygenated except in
artery blood pulmonary vein F
tunica externa of tunica externa of
collagen fibres collagen fibres
lumen lumen
endothelial cell
tunica intima of tunica media of smooth

tunica media of smooth tunica muscle fibres and a few
endothelium muscle and elastic fibres
intima elastic fibres
516
blood
cells
valve
thick wall
of artery
collagen fibres
wall
of
blood cells
vein
elastic
fibres and
smooth
muscles
Fig 14.39 (a) TS of an artery and a vein. (b) LS of a vein showing a valve
14.12 The mammalian circulatory system When these structures contract, blood is prevented from
flowing through the capillary network. Also present in
As blood circulates continuously round the certain regions of the body are arterio-venous cross-
body it passes through aseries of arteries, capillaries and connections, which act as short-circuit routes between
veins. Basically each artery and vein consists of three arterioles and venules and serve to regulate the quantity of
layers, an inner lining of squamous endothelium, the tunica blood which flows through the capillary beds according to
intima, a middle layer of smooth muscle and elastic fibres, the needs of the body.
the tunica media, and an external layer of fibrous Blood passes from the arterioles into capillaries, the
connective tissue possessing collagen-fibres, the tunica smallest of all blood vessels in the body. They form a vast
externa (table 14.11 and fig 14.39(a)). network of vessels pervading all parts of the body, and are
The large arteries in close proximity to the heart (that is
the aorta, subclavians and carotids) must be able to (a) =
contend with the high pressure of blood leaving the
ventricles of the heart. The walls of these vessels are thick
ae
ventricle
and the middle layer is mainly composed of elastic fibres. contracting \ panes
This enables them to dilate but not rupture during
ventricular systole (section 14.12.3). When systole ceases,
the arteries contract and promote an even flow of blood open valves
along their length (fig 14.40).
The arteries further away from the heart have a similar
structure but possess more smooth muscle fibres in the (b) closed valves
middle layer. They are supplied with neurones from the
sympathetic nervous system. Stimulation from this system
regulates the diameter of these arteries and this is ventricle
important in controlling the flow of blood to different parts
relaxed
of the body. SS elastic recoil

Blood passes from the arteries into smaller vessels called of vessel wall
arterioles. In all of these except the pulmonary arterioles, Fig 14.40 Diagram demonstrating how the arteries near the
the tunica media consists entirely of smooth muscle fibres heart assist in maintaining a continuous flow of blood in spite
supplied with neurones from the sympathetic nervous of a discontinuous flow received from the ventricles. (From
system (section 16.2). Many arterioles possess precapillary Clegg & Clegg (2nd edition, 1963) Biology of the mammal,
‘sphincters’ (figs 14.41 and 14.42) at their capillary ends. Heinemann Medical Books)
Sih
‘
so numerous that no capillary is more than 0.5 mm from Blood from the capillary beds drains into venules, whose
any cell. They are 7-10 um in diameter and their walls, walls consist of a thin layer of collagen fibres. They pass the
consisting solely of endothelium, are permeable to water blood into veins which eventually convey it back to the
and dissolved substances. It is here that exchange of heart.
materials between the blood and body cells takes place. A vein possesses less muscle and elastic fibres in its
<—_____blood flow (oxygenated)

arterio-venous
anastomosis precapillary
sphincter
venule
capillary bed
—¥ deoxygenated
arteriole | blood
venule
Fig 14.41 The possible routes that blood may take between arteriole, capillary bed and venule
IO aia
7 SB tat o
Pry
PF
eager
Pa
ha er J < EE
aS
mo
we
3 or
A
ws
1 ae
a!
seta
(a) OPEN (b) CLOSED
Fig 14.43 Action of semi-lunar vaive in a vein. (a) Upward

pressure of the blood forces the valve open and blood flows
towards the heart. (b) Backflow of blood closes valves,
blood therefore cannot flow away from the heart
Fig 14.42 Capillary bed showing arterioles and
capillaries
vein
upper valve
blood flow
towards the
muscle heart
lower valve valve
a (b) (o)
muscles relaxed, valves closed muscles contract, upper valve opens muscles relax, upper valve closes,
and blood is forced upwards, lower lower valve opens as a result of
valve remains closed muscle contraction elsewhere and
blood flows forwards
Fig 14.44 Diagram illustrating how muscle contraction around a vein aids one-way flow of blood towards the
heart
518
middle layer than an artery and the diameter of its lumen is 14.12.1 Formation of intercellular fluid
greater. Semi-lunar valves (fig 14.43) are present, being
formed from folds of the inner walls of the vein which are Intercellular, or tissue, fluid is formed when
permeated by elastic fibres. They function to prevent blood passes through the capillaries. The capillary walls are
backflow of blood thereby maintaining a unidirectional permeable to all components of the blood except the
blood flow. A number of veins are located between the erythrocytes and plasma proteins.
large muscles of the body (as in the arms and legs). When The osmotic pressure exerted by the plasma proteins is
these muscles contract they exert pressure on the veins and about 25 mm Hg” and this far exceeds the OP in the tissue
squeeze them flat (fig 14.44). This assists the venous flow to fluid. Under these conditions one would normally expect
tissue fluid to flow into the blood plasma. However, the
the heart. A general plan of the mammalian double
circulation is shown in fig 14.45. blood pressure at the arterial end of a capillary is about
32 mm Hg. Therefore fluid passes from the capillary into
the minute spaces between the cells to form the inter-
jugular vein carotid artery cellular fluid. It is through the tissue fluid that exchange of
HEAD & NECK
materials between blood and tissues occurs.
The blood cannot afford to constantly lose so much fluid
and therefore much of it is returned. This occurs in two
ways.
subclavian vein subclavian artery
FORELIMBS
(1) At the venous end of the capillary blood pressure has
<——_ ——— fallen to 12 mm Hg and therefore below the OP exerted
by the plasma proteins. Thus there is a net flow of tissue
anterior
| vena cava t fluid back into the capillary (fig 14.46).
(2) The rest of the intercellular fluid drains into blindly
ending lymphatic capillaries, and once inside these the
—> fluid is termed lymph. The lymphatic capillaries join to
dorsal aorta
form larger lymphatic vessels. The lymph is moved
through the vessels by contraction of the muscles
surrounding them, and backflow is prevented by valves
pulmonary
iz present in the major vessels which act in a similar
artery
pulmonary
fashion to those found in veins (fig 14.48).
; vein The lymphatic vessels of the legs join to those from
the alimentary canal to form the thoracic duct. This
empties the lymph into the blood system in the neck
region via the left subclavian vein. The right lymphatic
hepatic vein hepatic artery
duct drains lymph back into the bloodstream via the
_
right subclavian vein (fig 14.47).
Situated at intervals along the lymphatic system are
gastric artery lymph glands or nodes. Lymphocytes, in the course of
eo circulation through the blood and lymph, ‘rest’ and
posterior anterior mesenteric artery accumulate in the lymph nodes. They produce anti-
vena Cava
<—— bodies and are an important part of the body’s immune
posterior mesenteric artery} system. The nodes (fig 14.49) also filter out bacteria
<+— and foreign particles from the lymph, which are
ultimately ingested by phagocytes.
segmental lumbar segmental lumbar
veins arteries
e e 25.> TRUNK
<+— —
* In medicine, the unit of pressure commonly used is still millimetres of
mercury (mm Hg). For comparison with SI units, see section A4.3.3.
renal vein renal artery
(See ees eee | KIDNEYS |
a - <q -—-
genital vein genital artery

| Gonaps |
<j
<__
iliac artery
Fig 14.45 Mammalian double circulatory system. Principal
iliac vein POSTERIOR
& HINDLIMBS
blood vessels are shown but not capillary beds
519
arterial end venous end
of capillary
32
Hydrostatic
pressure/mm Hg
25
plasma colloidal OP
arterial end of venous end

capillary
: hydrostatic pressure hydrostatic pressure
3
blood flow| 32 mm Hg 12 mm H
plasma colloidal is i ema
osmotic pressure
25 mm Hg t
Ve
LOrGx (O
intercellular
fluid
tissue cell
() lymph
lymphatic capillary
Fig 14.46 Formation of tissue fluid and lymph. Some tissue fluid is reabsorbed at the venous end of the capillary whereas the
remainder is collected in lymphatic capillaries. (After K. Schmidt-Nielsen (1980) Animal physiology, 2nd ed., Cambridge
University Press)
14.12.2 |The mammalian heart

ee ee Adie tacae vein
The heart is situated between the two lungs
subclavian and behind the sternum in the thorax. It is surrounded by a
- vein conical-shaped sac, the pericardium, the outer part of
which consists of non-distensible white fibrous tissue,
axillary lymph whilst the inner part is made up of two membranes. The
nodes (armpit) fi
inner of the two membranes is attached to the heart whilst
the outer one is attached to the fibrous tissue. Pericardial
fluid is secreted between them which reduces the friction
mesenteric between the heart walls and surrounding tissues when the
thoracic duct lymph nodes heart is beating. The general inelastic nature of the
pericardium as a whole prevents the heart from being
gut overstretched or overfilled with blood.
There are four chambers in the heart, two upper
thin-walled atria and two lower thick-walled ventricles
inguinal lymph
nodes (groin) (fig 14.50). The right side of the heart is completely
separated from the left. The atria function to collect and
retain blood temporarily until it can pass to the ventricles.
The distance from atrium to ventricle is very small, hence
Fig 14.47 Human lymphatic system. (From E. G. the power of contraction of the atria does not have to be
Springthorpe 1973) An introduction to functional systems in very great. The right atrium receives deoxygenated blood
animals, Longman) from the general circulation of the body whilst the left
520
capsule
lymphatic
nodule
a
*
«
Pes
» ‘
fe
Fig 14.48 LS through lymph vessel showing an internal Fig 14.49 Section through a lymph gland
valve
aorta
pulmonary veins from
lungs
pulmonary artery
pulmonary veins from

vena cava lungs
left atrium
right atrium .
semi-lunar valves
tricuspid valve bicuepidalve
papillary muscle chordae tendinae
right ventricle left ventricle

J
interventricular depute) Fa
Fig 14.50 Section through mammalian heart
S21
atrium receives oxygenated blood from the lungs. The valves guard the point of entry of blood into these vessels
muscular wall of the left ventricle is at least three times as and prevent regurgitation of blood back into the ventricles.
thick as that of the right ventricle. This difference can be The walls of the heart are composed of cardiac muscle
correlated with the fact that the right ventricle has only to fibres, connective tissue and tiny blood vessels. Each
supply the pulmonary circulation whilst the left ventricle muscle fibre possesses one or two nuclei, myofilaments and
pumps blood through the much larger systemic circulation many large mitochondria. The fibres branch and cross-
of the body. Correspondingly the blood entering the aorta connect with each other to form a complex net-like
from the left ventricle is at a much higher blood pressure arrangement. This permits contraction waves to spread
(approximately 105 mm Hg) than the blood entering the quickly amongst the fibres and enhance the contraction of
pulmonary artery (16 mm Hg). the chambers as a whole. No neurones are present in the
wall of the heart (figs 14.51 and 14.52).
14.34 What other advantages arethere _
_ in supplying the pulmonary circulation with blood at a 14.12.3 The cardiac cycle
lower pressure than that of the systemic circulation? The cardiac cycle refers to the sequence of
events which take place during the completion of one
As the atria contract they force blood into the ventricles,
heartbeat. It is as follows.
and rings of muscle which surround the venae cavae and
(1) Deoxygenated blood, under low pressure, enters the
pulmonary veins at their point of entry into the atria
right atrium and oxygenated blood enters the left
contract and close off the veins. This prevents reflux of
atrium. These chambers gradually become distended.
blood into the veins. The left atrium is separated from the
Initially the bicuspid and tricuspid valves are closed,
left ventricle by a bicuspid (two-flapped) valve, whilst a
but as pressure in the atria rises, as they fill with
tricuspid valve separates the right atrium from the right
blood, it eventually exceeds that of the ventricles
ventricle. Attached to the ventricular side of the flaps are
and the valves are pushed open. Some of the blood
fibrous cords which in turn attach to conical-shaped
flows into the relaxed ventricles. This resting period
papillary muscles which are extensions of the inner wall of
of the heart chambers is called diastole (fig 14.53a).
the ventricles. These valves are pushed open when the atria
(2) When diastole ends, the two atria contract simul-
contract, but when the ventricles contract the flaps of each
taneously. This is termed atrial systole and results in
valve press tightly closed so preventing reflux of blood to
more blood being conveyed into the ventricles (fig
the atria. At the same time the papillary muscles contract
14.535). Almost immediately the ventricles contract.
so tightening the fibrous cords. This prevents the valves
This is called ventricular systole (fig 14.53c). When this
from being turned inside out. Pulmonary and aortic pocket
occurs the bicuspid and tricuspid valves are closed. The
cross-striations nucleus in ventricular pressure rises and soon exceeds the blood
of myofilaments sarcoplasm pressure in the aorta and pulmonary artery, forcing the
aortic and pulmonary valves open. Thus blood is
expelled from the heart into these elastic-walled
vessels. During ventricular systole blood is forced
against the closed atrio-ventricular valve and this
produces the first heart sound (‘lub’).
(3) Ventricular systole ends and is followed by ventricular
branching fibre
intercalated diastole (fig 14.53d). The high pressure developed in
disc
the aorta and pulmonary artery tends to force some
Fig 14.51 Structure of cardiac muscle blood back towards the ventricles and this closes the
aortic and pulmonary artery pocket valves. Hence back-
Fig 14.52 Photograph of cardiac muscle
flow is prevented. The impact of this backflow against
an
#85 wr wm *
myofibrils
Re
. spaces the valves causes the second heart sound (‘dub’):
showing striations . : befvcen
intercalated fibres filled ventricular systole =‘lub’
disc : with connective
issu d capillaries
ventricular diastole =‘dub’
The repeated recoil of the elastic arterial vessels as a
result of ventricular systole forces the blood into the
pulmonary and systemic circulations as a series of
pulses. As blood is propelled further and further away
from the heart, the pulses become less and less
pronounced until, in the capillaries and veins, blood
flows evenly (figs 14.55 and 14.56).
nt One complete heartbeat consists of one systole
Sh%
%
and one diastole and lasts for about 0.8 s (fig 14.54).
venae i pulmonary
Cavae
Ke vein
Atria
\AL left Ventricle

right
atrium
3 atrium
Mitral ae
valve
left
ventricle Aortic
right valve
ventricle
Isometric
phase left U
(a) Atria in state of diastole, and ventricle
filling with blood 120-120
Left side
of heart 100
( lie wall 80
. contracted
AV Pressure/
mm Hg al
oN {' bicuspid
valve
tricuspid
valve
(b) Atrial systole forces blood into

ventricles. Bicuspid and tricuspid
valves open. Sphincters of venae
cavae and pulmonary veins closed
right
Right 40 ventricle
side of
heart
Me
pulmonary /)7,
artery eli LA
Pressure/
—— mm Hg 0
filling
-20
100% Stroke
Ventricular 70% volume
volume
0%
(c) Atria relax, ventricles contract.
ECG
Blood propelled into aorta and
pulmonary artery
Buk
Heart
sounds
0 O1 02 03 04 0540.6 .07 0.8.09) 10014

Time/s
Fig 14.54 The sequence of events in the cardiac cycle starting with
the onset of atrial systole. Part of the subsequent cycle is shown. PA,
blood pressure in the pulmonary artery; RA, right atrial pressure; LA,
left atrial pressure. (From J. H. Green (1968) An introduction to human
physiology, Oxford University Press)
(d) Pocket valves of aorta and

pulmonary artery close. Atria begin
to refill. Ventricles in state of
diastole
Fig 14.53 Sequence of heart actions

involved in one complete heartbeat — the
cardiac cycle
a3
Volume/cm* Pressure/mm Hg Velocity/cm s~!
aorta 100 0 | ao
arteries [| 300 [_} 100-40 [=f 40-10
arterioles {|50 40-25 [| 10-0.1
capillaries fa 250 | 25-12 |< 0.1
venules [~} 300 [| 12-10 |< 0.3

veins ee 2200 []10-5 [] 0.3-5
vena cava Fa 300 |2 5-20
Fig 14.55 Distribution of blood volume, pressure and The cells of the S-A node maintain a differential ionic
velocity in the human vascular system. (From K. Schmiadt- concentration across their membranes of —90 mV. These
Nielsen (1980) Animal physiology, 2nd ed., Cambridge cells have a permanently high sodium conductance, that is
University Press)
to say that sodium ions continually diffuse into the cells.
This produces a depolarisation which leads to a propagated
Fig 14.56 Blood pressure throughout the human circulatory action potential (section 16.1.1) being set up in the cells
system. (From J. H. Green (1968) An introduction to human adjacent to the S-A node. As this wave of excitation passes
physiology, Oxford University Press) across the muscle fibres of the heart it causes them to
arteries arterioles veins
contract. The S-A node is known as the pacemaker of the
capillaries heart because each wave of excitation begins here and acts
as the stimulus for the next wave of excitation.
N oS
oo Once contraction has begun it spreads through the walls
of the atria via the network of cardiac fibres at the rate of
1 ms’. Both atria contract more or less simultaneously.
The atrial muscle fibres are completely separated from
Hg
pressure/mm
Blood those of the ventricles by the atrio-ventricular septum of
connective tissue, except for a region in the right atrium
<— Pulsatile__,»
4 Non-pulsatile_» ——
called the atrio-ventricular node (A-V node).
The tissues of the A-V node are similar to those of the
S-A node and supply a bundle of specialised fibres, the A-V
bundle, which provide the only route for the transmission
14.12.4 Mechanism of heart excitation and
of the wave of excitation from the atria to the ventricles.
contraction
There is a delay of approximately 0.15 sin conduction from
When a heart is removed from a mammal and the S-A node to the A-V node, thus permitting atrial
placed in well-oxygenated Ringer solution at 37 °C it will systole to be completed before ventricular systole begins.
continue to beat rhythmically for a considerable time,
without stimuli from the nervous or endocrine systems.
This demonstrates the myogenic nature of the heart, vena Cava aorta
that is it possesses its own inherent or ‘built-in’ mechanism
for initiating contraction of the cardiac muscle fibres.
sino-atrial node
atrio-ventricular
Myogenic control of heartbeat rate node |
left atrium
The stimulus for contraction of the heart originates in a right atrium

specific region of the right atrium called the sino-atrial node bundle of His
(or S-A node for short) close to the point of entry of the right ventricle I containing
Purkinje tissue
venae cavae. The S-A node is a vestige of the sinus venosus
seen in the heart of lower vertebrates. It consists of a small left ventricle
number of diffusely orientated cardiac fibres, possessing interventricular

few myofibrils, and a few nerve endings from the septum
autonomic nervous system. The S-A node initiates the
heartbeat, but the rate at which it beats can be varied by Fig 14.57 Position of the sino-atrial and atrio-ventricular
stimulation from the autonomic nervous system. nodes, and the bundle of His
524
14.12.5 Regulation of heartbeat rate
The intrinsic rate of the heartbeat is controlled
by the activity of the S-A nodes as described earlier. Even
®) ® when removed from the body and placed into an artificial
@ medium the heart will continue to beat rhythmically, albeit
0 0.2 0.4 0.6 0.8 more slowly. In situ, however, the body’s demands on its
Time/s circulatory system are constantly changing and the heart
rate has to be continuously adjusted accordingly. This is
Fig 14.58 An electrocardiogram (ECG) trace demonstrating
the change in electrical potential across the heart during one achieved by the dynamic and integrated activity of two
cardiac cycle. P, atrial depolarisation over the atrial muscle types of control system, one nervous and the other
and spread of excitation from the sino-atrial node equivalent chemical. This is a homeostatic response whose overall
to atrial systole; Q, R and S, ventricular systole; T, ventricular function is to maintain constant conditions within the
diastole begins bloodstream even though conditions around it are con-
stantly changing.
The A-V bundle is connected to the bundle of His, a strand The amount of blood flowing from the heart over a given
of modified cardiac fibres which gives rise to finer branches period of time is known as the cardiac output and depends
known as Purkinje tissue. Impulses are conducted rapidly upon the volume of blood expelled at each beat (the stroke
along the bundle at 5 ms"', ultimately proceeding to all volume) and the heart rate. These three variables are
parts of the ventricles. Both ventricles are stimulated to related by the expression
contract simultaneously, and the wave of ventricular
contraction begins at the apex of the heart and spreads cardiac output = stroke volume x heart rate
upwards squeezing blood out of the ventricles towards the and it is the cardiac output which is the important variable.
arteries which pass vertically upwards out of the heart One way of controlling cardiac output is by varying the
(figs 14.57 and 14.58). heart rate.
Certain characteristics of cardiac muscle make it suited
Nervous control of heart rate
to its role of pumping blood round the body throughout the
life of the mammal. Once cardiac muscle has begun to Within the medulla oblongata of the hindbrain are several
contract it cannot respond to any other stimulus until it regions concerned with cardiovascular control. In all
begins to relax. This is known as the refractory period. The regions, part of their function is to control the heart rate.
length of time that the cardiac muscle is in this condition is Two vagus nerves carrying parasympathetic fibres leave the
called its absolute refractive period (fig 14.59). This period cardio-inhibitory centre of the medulla oblongata and run,
is longer than that of other types of muscle, and enables the one on either side of the trachea, to the heart. Here nerve
fibres lead to the S-A node, A-V node and the bundle of
muscle to contract vigorously and rapidly without becom-
ing fatigued. It is thus impossible for the heart to develop a His. Impulses passing along the vagus nerve reduce the
state of sustained contraction called tetanus, or to develop heart rate. Other nerves, of the sympathetic system, have
an oxygen debt. their origin in the pressor region of the vasomotor centre of
the medulla, run parallel to the spinal cord and emerge
from the thoracic region to the S-A node. Stimulation by
these nerves results in an increase in the heart rate. It is the
integrated activity of the inhibiting and accelerating
Contraction effects, occurring within the medulla oblongata, that
controls the heart rate.
Sensory fibres from stretch receptors within the walls of
the aortic arch, the carotid sinuses and the vena cava run to
the cardio-inhibitory centre in the medulla. Impulses
received from the aorta and carotids decrease the heart
rate, whilst those from the vena cava stimulate the pressor
centre which increases the heart rate. As the volume of
psc blood passing to any of these vessels increases so does the
refractive refractive
normal period period
excitable normal distension of the vessels, and this increases the number of
impulses transmitted to the cardiovascular centres in the
medulla.
Time ———>
For example, under conditions of intense activity body
Fig 14.59 The refractory period of cardiac muscle. The muscles contract strongly and this increases the rate at
upper figure shows a record of the contraction of the muscle, which venous blood returns to the heart. Consequently the
the lower figure shows the varying excitability of the muscle to
of the vena cava is distended by large quantities of blood and the
stimuli. (From Clegg & Clegg (2nd edition, 1963) Biology
mammal, Heinemann Medical Books)
heart rate is increased. At the same time the increased
S29
Table 14.12 Non-nervous agencies affecting heart rate.
medulla ee ooo
oblongata
Non-nervous stimulus Effect on heart rate
cardiovascular
centres
High pH Decelerates
Low pH (e.g. high CO, levels, | Accelera tes
as is the case during active
sensory nerve sensory
ensory, nerve exercise)
(accelerating) (inhibiting) Decelerates
Low temperature
High temperature Accelerates
Mineral ions
Endocrine factors (e.g.
inhibitor nerve accelerator thyroxine, insulin, sex The rate is influenced directly
(vagus) nerve or indirectly
(sympathetic
hormones, adrenaline,
(parasympathetic
system) system) pituitary hormones)
aorta
vena Cava
where they pass to the cardiovascular centre via intercon-
necting pathways. Under such stimulation the cardio-
sino-atrial node
vascular centre responds accordingly (table 14.12).
It should be noted that this centre is influenced at any
given moment by a combination of nervous and non-
nervous agencies, and never by a single one. The activity of
the cardiovascular centre also fluctuates according to the
health and age of the individual.
——»—— Sensory innervation — nerves originate in the vena cava, aorta
and carotid sinuses and run to the heart rate centre (those from 14.12.6 Regulation of blood pressure
the carotid sinuses not shown in diagram)
——p~— — Motor innervation — impulses from the accelerator nerve Blood pressure depends on several factors:
stimulate the heart rate, impulses from the vagus decelerate it heart rate, strength of heartbeat, blood output (stroke
Fig 14.60 Neurones connecting the heart to the
volume) and resistance to blood flow by the blood vessels
cardiovascular system (peripheral resistance). Heart rate and stroke volume have
been discussed in the last section. Resistance to blood flow
blood flow to the heart places the cardiac muscle of the is altered by contraction (vasoconstriction) or relaxation
heart under tension. Cardiac muscle responds to this (vasodilation) of the smooth muscle in the blood vessel
tension by contracting more strongly during systole and walls, especially those of the arterioles. This peripheral
pumping out an increased volume of blood (the stroke resistance is increased by vasoconstriction but decreased by
volume). This relationship between the volume of blood vasodilation. Increased resistance leads to a rise in blood
returned to the heart and cardiac output was named after pressure, whereas a decrease produces a fall in blood
the English physiologist Starling and is known as Starling’s pressure. All such activity is controlled by a vasomotor
law. centre in the medulla oblongata.
The increased stroke volume stretches the aorta and Nerve fibres run from the vasomotor centre to all
carotids which in turn, via stretch reflexes, signal the arterioles in the body. Changes in the diameter of these
cardio-inhibitory centre to slow the heart rate. Therefore blood vessels are produced principally by variation in the
there is an automatic fail-safe mechanism which serves to activity of constrictor muscles. The dilator muscles play a
prevent the heart from working too fast, and to enable it to somewhat less important role.
adjust its activity in order to cope effectively with the Vasomotor centre activity is regulated by impulses
volume of blood passing through it at any given time coming from pressure receptors (baroreceptors) located in
(fig 14.60). the walls of the aorta and carotid sinuses in the carotid
arteries (fig 14.61). Stimulation of parasympathetic fibres
Non-nervous control of heart rate in these areas, caused by increased cardiac output,
There are a number of non-nervous stimuli which act produces vasodilation throughout the body and conse-
directly on cardiac muscle or on the S-A node. They are quent reduction in blood pressure, as well as a slowing of
briefly summarised in table 14.12. the heart rate. The converse occurs when blood pressure is
Many activities affect the cardiovascular centre in some low. In this case, a fall in blood pressure increases impulse
way or other, for example emotions, such as blushing or transmission along sympathetic fibres. This causes body-
turning white with anger, sights and sounds. In such wide vasoconstriction and a compensatory rise in blood
instances sensory impulses are transmitted to the brain pressure.
526
medulla
oblongata
vasomotor \____} ~\
ia
centre
large number few dilator few large number

of constrictor impulses constrictor] | |of dilator
impulses impulses impulses
external internal o>
carotid carotid
artery artery
carotid body — registers carotid sinus -

changes in CO,/O, monitors
levels and pH pressure
ree changes
carotid blood flow
artery
> many impulses
——p, few impulses VASOCONSTRICTION VASODILATION
Fig 14.61 Interrelationships between the carotid body, carotid sinus, vasomotor centre and general circulatory system
Chemical control of the vasomotor centre and therefore raises blood pressure. The significance of the
Blood arriving at the carotid bodies carrying a high control of heart rate and blood pressure is described further
concentration of carbon dioxide stimulates chemorecep- in section 18.1.5.
tors in these regions to transmit impulses to the vasomotor
centre (fig 14.61). Nerve fibres leaving the chemoreceptors 14,35 When an animal is wounded, its
synapse with fibres from the carotid sinus prior to passing to overall blood pressure rises, but the area in the -
the vasomotor centre. When the vasomotor centre is vicinity of the wound swells as a result of local
vasodilation. Why? a
stimulated in this way it sends impulses to the blood vessels
to vasoconstrict and therefore raises blood pressure. As 14.36 Outline the main adjustments that
increased carbon dioxide concentration in the body is occur to the heart rate and circulatory system just
usually brought about by increased activity by body tissues, before, during and after a 100 m race. —
the blood containing the carbon dioxide will be transported
more rapidly to the lungs where expulsion of carbon
dioxide in exchange for oxygen can take place more
quickly. 14.13 Functions of mammalian blood
Carbon dioxide can also directly affect the behaviour of
the smooth muscle of the blood vessel itself. When a tissue Mammalian blood performs many major func-
suddenly becomes very active, producing a large quantity tions. In the following list, the first five functions are carried
of carbon dioxide, the carbon dioxide acts directly on the out solely by the plasma.
blood vessels in the vicinity and stimulates them to dilate. (1) Transport of soluble organic compounds from the small
This increases their own blood supply thus allowing more intestine to various parts of the body where they are
oxygen and glucose to reach the active cells. It must be stored or assimilated, and transport from storage areas
remembered, however, that when the carbon dioxide to places where they are used.
leaves this localised area it will have the effect of promoting (2) Transport of soluble excretory materials from tissues
vasoconstriction elsewhere via vasomotor activity. This is a where they are produced to the organs of excretion.
good illustration of how dynamic and adjustable the (3) Transport of metabolic by-products from areas of
control of blood pressure and therefore circulation and production to other parts of the body.
distribution of blood can be. (4) Transport of hormones from the glands where they are
Other agencies, such as types of emotional stress (for produced to all parts of the body or certain target
example excitement, pain and annoyance), increase sym- organs. This facilitates communication within the
body.
pathetic activity and therefore blood pressure. Also when
the adrenal medulla is stimulated to produce adrenaline by (5) Distribution of heat from the deeply seated organs.
This serves to dissipate excess heat and to aid the
impulses from higher nervous centres this again increases
maintenance of a uniform body temperature.
the rate of heartbeat, promotes bodywide vasoconstriction
527
(6) Transport of oxygen from the lungs to all parts of the The amount of oxygen that can combine with haemoglo-
body, and carriage of carbon dioxide produced by the bin is determined by the oxygen tension. This is expressed
tissues in the reverse direction. as a partial pressure and is the fraction of oxygen found in
(7) Defence against disease. This is achieved in three ways: the air. It is still measured in millimetres of mercury. For
(a) clotting of the blood which prevents excessive example, atmospheric pressure at sea level is 760 mm Hg.
blood loss and entry of pathogens; Approximately one-fifth of the atmosphere is oxygen,
(b) phagocytosis, performed by the granulocytes which therefore the partial pressure of oxygen in the atmosphere
engulf and digest bacteria which find their way into at sea level ist x 760 =152 mm Hg. When the percentage
the bloodstream; oxygen saturation of blood is plotted against the partial
(c) immunity mediated by antibodies and/or lympho- pressure of oxygen an S-shaped curve, called the oxygen
cytes. dissociation curve, is obtained (fig 14.63).
(8) Maintenance of a constant blood osmotic pressure and Analysis of the curve indicates that for physiological
pH asa result of plasma protein activity. As the plasma reasons haemoglobin is completely saturated with oxygen
proteins and haemoglobin possess both acidic and basic at a point known as the loading tension. This is taken as the
amino acids they can combine with or release hydrogen tension when 95% of the pigment is saturated and coincides
ions and serve to minimise pH changes over a wide with a partial pressure of about 73 mm Hg in the example
range of pH values. given in fig 14.63. At higher partial pressures of oxygen
further uptake of oxygen can occur, but 100% saturation of
haemoglobin is rarely achieved. At an oxygen partial
14.3.1 Oxygen carriage pressure of approximately 30 mm Hg only 50% of the
It is the haemoglobin molecule, found in the haemoglobin is present as oxyhaemoglobin, and at a partial
erythrocytes, which is responsible for the transport of pressure of zero no oxygen is attached to the haemoglobin
oxygen round the body. Haemoglobin is a tetrameric molecule. Over the steep part of the curve, a small decrease
protein with a relative molecular mass of 68 000. It in the oxygen partial pressure of the environment will bring
possesses four haem prosthetic groups, responsible for the about a sizeable fall in the percentage saturation of
characteristic red colour of the blood, linked to four globin haemoglobin. The oxygen given up by the pigment is
polypeptide chains. A ferrous iron atom is located within available to the tissues.
each haem group, and each of these can combine loosely But why is the curve S-shaped? The explanation lies in
with one molecule of oxygen (fig 14.62). the way that haemoglobin binds to oxygen. It involves the
phenomenon of allostery (section 6.6). When an oxygen
Hb + 40, = HboO, molecule combines with the ferrous iron atom of a single
Combination of oxygen with haemoglobin, to form haem unit it distorts its shape slightly. This distortion is
oxyhaemoglobin, occurs under conditions when the partial registered by the whole molecule which changes its shape
pressure of oxygen is high, such as in the lung alveolar accordingly. Further structural alterations occur when
capillaries. When the partial pressure of oxygen is low, as in
the capillaries which supply metabolically active tissues,
the bonds holding oxygen to haemoglobin become unstable 100
and oxygen is released. This diffuses in solution into the OS) SS SSS
|
surrounding cells. |
|
|
CH,
§3s ||
I 5 |
CHa CH, eS0 a loading
5p | tension
oe oe =o !| |
/ CH=CH, S eae |
2 unloading or
half-saturation |
tension |
J H | |
Ba 0
|
he Ve ee
|
ee
0 50 100
ILC CH,
NX sS We Oxygen tension/mm Hg
Fig 14.63 Diagram to explain the terminology applied to

C H,eer CH, oxygen dissociation curves of oxygen-carrying pigments.
| | Loading tension is the tension at which 95% of the pigment is
CH, CH,
saturated with oxygen; unloading tension is the tension at
| |
COOH COOH which 50% of the pigment is saturated with oxygen. (From
Florey (1966) An introduction to general and comparative
Fig 14.62 A single haem molecule physiology, W.B. Saunders & Co.)
528
oxygen molecules attach to the second and third haem
groups, each one facilitating much faster uptake of oxygen 14.37 What is the physiological signifi- —
than the preceding one. When the last haem monomer is cance of the Bohr effect?
ready to pick up oxygen it does so several hundred times 14.38 Consider fig 14.65. The oxygen
faster than the first. dissociation curve of the foetus is to the left of that of
The converse reaction occurs when oxyhaemoglobin is its mother. Why is this so?
exposed to regions where the partial pressure of oxygen is 14.39 The oxygen dissociation curve of
low, as in actively respiring tissues. The first oxygen the South American llama, which lives in the High
molecule is released to the tissues very rapidly, but the Andes at an altitude of about 5000 m above sea
second, third and fourth molecules are given up much less level, is located to the left of most other mammals (fig ©
readily and only at a very reduced partial pressure of 14.66). Why is this so?
oxygen.
In regions with an increased partial pressure of carbon
dioxide the oxygen dissociation curve is shifted to the right.
This is known as the Bohr effect or shift (fig 14.64).
oxygen pressure/kPa Oxygen pressure/kPa

0 een 4 ee 12 0 4 8 12
100_ __ Carbon
Carbon dioxide
dioxide presyure/mmHe
Hg (PPT
Rhs eel eo ee)
100
80
80
foetal maternal
60
60
40
saturation
Percentage Saturation/%
20 40
0
a aneieen eee ae cies 20
Fig 14.64 (above) Oxygen dissociation curves of

haemoglobin illustrating the Bohr effect at different pressures — 80
20 40 60
of carbon dioxide. (From Garth Chapman (1967) The body
fluids and their functions, Studies in Biology No. 8, Arnold) Oxygen pressure/mm Hg
Oxygen partial pressure/kPa
ee
0 4 8 12
Fig 14.65 (above) Oxygen dissociation
100 curves of foetal and maternal blood of a
goat
80
60
f
40
Saturation/%
oo other mammals
— llama
|
20
Fig 14.66 (left) Oxygen dissociation

Al curves of the llama and other mammals
20 40 60 80 100 120
Oxygen partial pressure/mm Hg
529
100 Fig 14.67 Comparison of haemoglobin and
myoglobin oxygen dissociation curves.
myoglobin | Myoglobin remains 80% saturated with
oxygen until the partial pressure of oxygen
80 a falls below 20 mm Hg. This means that
myoglobin retains its oxygen in the resting
cell but gives it up when vigorous muscle
60 / haemoglobin
activity uses up the available oxygen
supplied by haemoglobin
40
Saturation/%
0 40 80 120
Oxygen partial
pressure/mm Hg
14.13.2 | Myoglobin 14.13.4 Carriage of carbon dioxide

The myoglobin molecule is widely distributed Carbon dioxide is carried by the blood in three
in animals and is particularly common in skeletal muscle different ways.
tissues of mammals. It displays a great affinity for oxygen
In solution (5%). Most of the carbon dioxide
and its oxygen dissociation curve is displaced well to the
carried in this way is transported in physical solution. A
left of haemoglobin (fig 14.67). In fact it only begins to re-
very small amount is carried as carbonic acid (H,CO;).
lease oxygen when the partial pressure of oxygen is below
20 mm Hg. In this way it acts as a store of oxygen in resting Combined with protein (10-20%). Carbon
muscle, only releasing it when supplies of oxyhaemoglobin dioxide combines with the amine (NH,) group of haemo-
have been exhausted. Myoglobin is very similar to the globin to form a neutral carbamino-haemoglobin com-
haemoglobin subunits with respect to both amino acid pound. The amount of carbon dioxide that is able to
sequence and three-dimensional structure, but the myoglo- combine with haemoglobin depends on the amount of
bin molecules do not associate to form tetramers and so oxygen already being carried by the haemoglobin. The less
cannot show cooperative oxygen binding. The two proteins the amount of oxygen being carried by the haemoglobin
have presumably evolved from a common ancestral molecule, the more carbon dioxide that can be carried in
molecule. this way:
H
Z”
HHbNH, +CO, —————®» HHbN—C
14.13.3 Carbon monoxide and haemoglobin ou
haemoglobin
The affinity of the ferrous ions in haemoglobin carbamino-haemoglobin
for carbon monoxide is several hundred times as great as it As hydrogencarbonate (85%). Carbon
is for oxygen. Therefore haemoglobin will combine with dioxide produced by the tissues diffuses passively into the
any carbon monoxide available in preference to oxygen to bloodstream and passes into the erythrocytes where it
form a relatively stable compound called carboxyhaemo- combines with water to form carbonic acid. This process is
globin. If this occurs oxygen is prevented from combining catalysed by the enzyme carbonic anhydrase found in the
with haemoglobin, and therefore the transport of oxygen erythrocytes and takes less than one second to occur.
round the body by the blood is no longer possible. In Man, Carbonic acid then proceeds to dissociate into hydrogen
collapse follows quickly after exposure to carbon monoxide and hydrogencarbonate ions:
and, unless the victim is removed from the gas, asphyxia-
tion is inevitable. Removal from the gas must be followed CO, + H,O = H,CO, = H* +HCO,
by administering a pure oxygen-carbon dioxide mix since When the erythrocytes leave the lungs their oxyhaemo-
the oxygen tension in air is insufficient to replace the globin (represented as HbO,) is weakly acidic and
carbon monoxide attached to the haemoglobin. associated with potassium ions. This may be represented as
530
KHbO,. In areas of high carbon dioxide concentrations (as
at the tissues), oxygen is easily given up by oxyhaemoglo- 14.40 Summarise how carbon dioxide in
bin. When this happens the haemoglobin becomes strongly the blood is expelled as gaseous carbon dioxide by
basic. In this state it dissociates from the potassium ions the lungs.
and readily accepts hydrogen ions from carbonic acid
forming haemoglobinic acid (H.Hb). The potassium ions
associate with hydrogencarbonate ions to form potassium 14.13.5 Defensive functions of the blood
hydrogencarbonate. Every mammal is equipped with a complex
KHbO, sae a KE Unt 0, system of defensive mechanisms which are designed to
enable it to withstand attacks by pathogens, and to remove
H* +HCO, +KHb = H.Hb + KHCO, foreign materials from its system. Three defensive mechan-
(haemoglobinic
acid)
isms are discussed here:
(1) clotting of blood both contributing to
By accepting hydrogen ions, haemoglobin acts as a buffer (2) phagocytosis wound healing
molecule and so enables large quantities of carbonic acid to (3) immune response to infection.
be carried to the lungs without any major alteration in
blood pH. Clotting
The plasma membrane of an erythrocyte is relatively “When a tissue is wounded blood flows from it, and
impermeable to the passage of sodium and potassium ions, coagulates to form a blood clot. This prevents further blood
however a cation pump operates and expels large numbers loss and entry of pathogenic micro-organisms and is of clear
of sodium ions into the plasma. The majority of hydro- survival value to the animal concerned. It is just as
gencarbonate ions formed within the erythrocyte diffuse important that blood in undamaged vessels does not clot.
out into the plasma along a concentration gradient and The highly complex series of reactions that take place in
combine with sodium to form sodium hydrogencarbonate. order for coagulation to be achieved serves at the same
The loss of hydrogencarbonate ions from the erythrocyte is time to prevent it from occurring unnecessarily. The whole
balanced by chloride ions diffusing into the erythrocyte clotting process depends on at least 12 clotting factors
from the plasma. Thus electrochemical neutrality is working in harmony with each other. Only the main factors
maintained. This phenomenon is called the ‘chloride shift’. are described in this account.
Potassium hydrogencarbonate, formed in the erythrocyte, Blood escaping from a superficial wound is exposed to
is also capable of dissociating, and some of the chloride ions the air, and mixes with substances oozing from the
which enter the erythrocyte combine with potassium ions damaged cells and ruptured platelets. Thromboplastin, a
to form potassium chloride, whilst the hydrogencarbonate lipoprotein released from injured tissues, together with
ions diffuse out. When hydrogencarbonate leaves the clotting factors VII and X (plasma enzymes) and calcium
erythrocyte, the excess H* ions which remain decrease the ions, catalyses the conversion of inactive plasma protein
pH within the erythrocyte causing the dissociation of prothrombin to thrombin. Thrombin is a proteolytic
potassium oxyhaemoglobin (KHbO,) into oxygen and enzyme that hydrolyses the large soluble globular plasma
potassium haemoglobin (fig 14.68). protein molecule, fibrinogen, into smaller units which then
When the erythrocytes reach the lungs the reverse associate to form a meshwork of long tangled needle-like
process occurs. fibres of insoluble fibrillar fibrin (fig 14.69). When
PLASMA
CO>. FO mS HCO eH +HCO,,

tissue
fluid CO,+ HHbNH,=———= HHbNHCOOH
(haemoglobin) (carbamino- NaHCO,
haemoglobin)
TISSUE PLASMA
FLUID
Nat
tissue O,+KHb + <=; NaCl
fluid Cis
ERYTHROCYTE
KHbO, |
(oxyhaemoglobin)
chloride
shift
MITOCHONDRION
Fig 14.68 Carbon dioxide carriage by the plasma and erythrocyte and its role in the release of oxygen at the tissues
531
platelet Fig 14.69 Major features involved in the
prea OOS Ske clotting process
ne oe
wound OR ee thromboplastin
sustained to air
tissue
damage
erothrombin it ep thrombin
plasma (made
in liver)
i fibrinogen in ——___» fibrin

plasma Ca?* plasma (made in (clot)
enzymes liver)
Fig 14.70 (below) A red blood cell enmeshed in fibrin in a blood clot
red blood cell
meshwork of fibrin
fibrinogen has been removed from the plasma, the fluid oe te Ts bacterium
which remains is called serum. Blood cells become trapped
in the meshwork (fig 14.70) and a blood clot is formed. It neutrophil
dries to form a scab which acts to prevent further blood
loss, and as a mechanical barrier to the entry of pathogens.
(a) migration of neutrophil
Because the clotting process is so elaborate, it means that towards bacterium
the absence or low concentration of any of the essential
clotting factors could produce excessive bleeding. Such a
condition is known as haemophilia. For example, if an
essential factor necessary for the action of thromboplastin
is absent or only present in minute amounts, the individual
(b) adherence of bacterium
will bleed profusely from any minor cut. Haemophilia is to neutrophil
due to an inheritable gene mutation and is transmitted on
an X chromosome. The condition is usually seen only in CaN phagosome
males, with females being carriers, since the abnormal Fo forming
allele is carried on the X chromosome and is recessive to lysosome
the normal allele on the other X chromosome (section

23.651): (c) phagocytosis
Clotting does not occur in undamaged blood vessels
because the lining of the vessels is very smooth and does phago-
lysosome
not promote platelet or cell rupture. Also present are lysosome
substances which actively prevent clotting. One of these is
heparin, present in low concentrations in the plasma and
produced by most cells found in the connective tissues and (d) digestion
the liver. It serves to prevent the conversion of prothrom-

bin into thrombin, and fibrinogen to fibrin and is widely
used clinically as an anticoagulant.
If a clot does form within the blood circulation it is called
a thrombus and leads to a medical condition known as (e) absorption of digested
portions of bacterium
thrombosis. This may happen if the endothelium of a blood
vessel is damaged and the roughness of the damaged area Fig 14.71 Phagocytosis of a bacterium by a neutrophil
promotes platelet breakdown and sets in motion the
clotting process. Coronary thrombosis, a thrombus de- spleen and lymph nodes, large resident phagocytes, usually
veloping in the coronary artery of the heart, is particularly termed macrophages, are present. Their role is to engulf
dangerous and can lead to a swift death. toxic foreign particles as well as microbes, and to retain
Phagocytosis them for long periods of time, often permanently. In this
way infection can often be localised. The macrophages
This function is generally carried out by neutrophils. These together with the neutrophils form the body’s reticulo-
are amoeboid cells which are attracted to areas where cell endothelial system.
and tissue damage has occurred. The stimulus for this
migration appears to be some of the chemicals liberated by Inflammation
the ruptured blood cells and tissues. The neutrophils are When an area of the body is wounded, the localised
able to recognise any invading bacteria. This capability is reaction of the tissue surrounding the wound is to become
enhanced by plasma proteins called opsonins, which swollen and painful. This is called inflammation and is due
become attached to the surfaces of the bacteria and in some to the escape of chemicals, including histamine and
way make them more recognisable. Then the bacteria are 5-hydroxytryptamine, from the damaged tissues. Collec-
engulfed in an amoeboid fashion and a phagosome is tively they cause local vasodilation of capillaries. This
formed (fig 14.71). Small lysosomes fuse with the phago- increases the amount of blood in the area and raises the
some, forming a phagolysosome. Lysozyme and other temperature locally. Permeability of the capillaries is also
hydrolytic enzymes, together with acid, are poured into the increased, permitting escape of plasma into the surround-
phagolysosome from the lysosomes and the bacteria are ing tissues and a consequent swelling of the area, a
digested. Ultimately the soluble products of bacterial condition known as oedema. This plasma contains bacter-
digestion are absorbed into the surrounding cytoplasm of iocidal factors, antibodies and neutrophils, all of which
the neutrophil. help to combat spread of infection. Fibrinogen is also
Neutrophils are able to squeeze through the walls of present to assist blood clotting if necessary, and the excess
blood capillaries, a process called diapedesis, and move tissue fluid tends to dilute and negate any potential toxic
about in the tissue spaces. In organs such as the liver, irritants.
So
Wound healing 14.14 The immune system
Towards the end of the inflammatory phase, cells called
fibroblasts appear and secrete collagen. This is a fibrous Immunity has been defined by Sir Macfarlane
protein and becomes linked to polysaccharide to form a Burnet as
meshwork of randomly arranged fibrous scar tissue. ‘the capacity to recognise the intrusion of material
Vitamin C is important for collagen formation; without it foreign to the body and to mobilise cells and cell products
hydroxyl groups cannot be attached to the collagen to help remove that particular sort of foreign material
molecule and so it remains incomplete. After about 14 days with greater speed and effectiveness’.
the disorganised mass of fibres is reorganised into bundles
arranged along the lines of stress of the wound. Numerous Basic definitions
small blood vessels begin to ramify through the wound.
An antibody is a molecule synthesised by an animal in
They function to provide oxygen and nutrients for the cells response to the presence of foreign substances for which it
involved in repairing and healing the wound.
has a high affinity. Each antibody is a protein molecule
Whilst these processes are going on within the wound,
called an immunoglobulin (formerly globulin). Its structure
the epidermis around it is also engaged in repair and
consists of two heavy H-chains of 50 000-60 000 relative
replacement activity. Some epidermal cells migrate into
molecular mass, and two light L-chains of 23 000 relative
the wound and ingest much of the debris and fibrin of the
molecular mass. Functionally the antibody has a constant
blood clot which has formed over the wound. When the
and variable part, the variable part acting something like a
epidermal cells meet, they unite to form a continuous layer
key which specifically fits into a lock (fig 14.72). Each
under the scar. When this is complete the scab sloughs off
organism can produce thousands of antibodies with
thus exposing the epidermis to the surrounding atmos-
different specificities, recognising all kinds of foreign
phere.
substances.
Summary of events The name antigen or immunogen is given to the foreign
(1) Wound occurs, blood flows. material that elicits antibody formation. Antigen usually
(2) Clotting process occurs. takes the form of a protein or polysaccharide structure on
(3) Inflammation occurs. the surface of microbial organisms or as a free molecule.
(4) White cells migrate into wound. They absorb foreign Two systems of immunity have been developed by
matter and bacteria, and remove cell debris. mammals, a cell-mediated immune response and a humor-
(5) Fibroblasts enter the wound and synthesise collagen al response.
which is built up into scar tissue. The division of labour in the immune system is caused by
(6) Epidermal cells remove any final debris in the wound, the development of two types of lymphocytes, the T and B
and also begin to dismantle the scar. cells. Both types arise from precursor cells in the bone
(7) Epidermis creates a new skin surface in the area of the marrow. The influence of the thymus gland is essential in
wound. making the T cells immunologically competent, and it is
(8) Scab sloughs off. thought that the placenta, foetal liver and bone marrow
If the wound is small, phagocytosis is usually sufficient to exert a similar influence over the development of the B
cope with any pathogenic invasion. However, if there is cells. Within each type there is an enormous capacity for
considerable damage, the immune response of the body is recognising each of the millions of antigens that exist.
put into action. When an antibody-antigen reaction occurs it serves to fix
antigen-binding site
variable regions where the

amino acid sequence is
different in different molecules
light chain
heavy chain
‘hinge’
constant regions
where the amino disulphide bond
acid chain is Fig 14.72 Immunoglobulin antibody
the same in all molecule. Antigens are bound between the
molecules of the light and heavy chains of the variable
same type regions. (From M. Cooper & A. Lawton
(1972) The development of the immune
system, Scientific American)
534
and nullify the action of the antigen, thereby preventing it bone marrow
from acting upon the body in a harmful way.
Cell-mediated response. T cells possessing stem cells
membrane receptors which recognise antigen are stimu-
lated to proliferate and produce a clone of T cells. These stimulate B
thymus
cells then either combat micro-organisms and/or effect the cells to
increase
rejection of foreign tissues. antibody
lymphocytes thymocytes production
Humoral immune response. B cells recog- A
nise antigen in a similar way to T cells. However their thymosin lymphokines
|
response is different. They are stimulated to proliferate immunologically |
and form a plasma cell clone. The plasma cells synthesise competent T ES
lymphocytes
and liberate antibodies into the blood plasma and tissue
fluid. Here the antibodies adhere to the surfaces of bacteria
and speed up their phagocytosis, or combine with and
lymphoid spleen
neutralise toxins produced by micro-organisms. tissue
clones clones
14.14.1. The thymus gland and the
development of T cells general circulation
The thymus gland is situated in the thorax
Fig 14.73 T-cell differentiation and activity
under the sternum and close to the ventral side of the heart.
It begins to function during the embryonic period of the fo
individual and is at its most active at the time of, and just FT celts constantly leave the thymus and pass to the lymph
after, birth. After the period of weaning it decreases in size nodes and spleen. Here, if an antigen is recognised by a T
and soon ceases to function. cell, the T cell divides to form a clone of cells, all of which
Evidence that the thymus gland is important in the can identify and react with the antigen (fig 14.74). An
development of the immune response can be demonstrated individual T lymphocyte combats foreign cells by produc-
by the following experiments. ing a receptor for antigen which is built into its plasma
(1) Removal of the gland from a newborn mouse results in membrane. When this sensitised lymphocyte recognises a
death from a chronic deficiency of lymphocytes in its complementary antigen, it attaches itself to it, rather like a
tissue fluid and blood. key fitting into a lock, and destroys it. T cells regularly
(2) Tissue from another mouse grafted onto an ex- leave the lymphoid tissues to circulate in the blood and
perimental newborn mouse with the gland removed is tissue fluid. Their constant circulation increases their
unable to recognise and react with antigens. chances of meeting and combating antigens. /
_ (3) If the thymus gland is removed from a much older
V
mouse, this mouse suffers no adverse effects.

14.14.2 Bcell production
The stem cells of the bone marrow, which give rise to T
lymphocytes, must pass through the tissue of the thymus Stem cells that will eventually become B
gland before they can become fully functional. The lymphocytes must undergo further differentiation outside
mechanism employed for the maturation of T lymphocytes the bone marrow. This may be in the liver, spleen or lymph
has yet to be clearly understood, but it is known that the nodes.
thymus secretes a hormone called thymosin which may When surface receptors (immunoglobulins) on the B
promote T lymphocyte maturation. However, the role of lymphocytes detect complementary antigens, the B lym-
the thymus as an endocrine organ has yet to be clearly phocytes are stimulated to divide repeatedly and differenti-
established. The spleen and other lymph glands possess ate into plasma cell clones and memory cells (fig 14.74).
small numbers of lymphocytes at birth. The plasma cells, being genetically identical with each
The cortex of the thymus gland is packed with lympho- other, proceed to synthesise large quantities of antibody of
cytes, and those resident in the gland are called thymocytes, the same kind.
in order to distinguish them from the T lymphocytes These antibody-producing cells live for only a few days,
circulating in the blood and body fluids (fig 14.73). Most but during that time they are able to synthesise and secrete
thymocytes are immature, but a few are capable of reacting nearly 2 000 identical antibody molecules per second.
with antigen. When they do, they proliferate and manufac- The lymphatic memory cells are important in activating
ture complex molecules called lymphokines which aid in the body’s response to a second infection of the antigen.
the attack and elimination of foreign particles. Mature, Very little is known about memory cells, only that they
antigen-specific T lymphocytes are also required to help B exist and that in some way they enable an individual who
lymphocytes mature and produce their antibody. has been exposed to an antigen once to respond more
®
535
bone marrow
promptly and vigorously in a second encounter. This is
called the secondary response and it results in a massive
outpouring of antibody which quickly neutralises the
thymus lymphoid tissue harmful effect of the antigen (fig 14.76). Thus immunity is
achieved. However, it must be remembered that immunity
~ C09 to one antigen does not protect the individual against any
other antigens. Each time a new antigen gains entry into
+~ er, ‘
~S%on
the body, the appropriate complementary antibody must
peripheral peace
/~
peripheral
B
be produced if disease is to be prevented.
V
lymphocyte egen lymphocyte
14.14.3 Classes of immunoglobulins and
memory
cells their biological activities
activated
i plasma ee There are five classes of immunoglobulin
(B-immunocytes)
cells (T-immunocytes) antibodies. Each class is determined by the type of heavy
chain in its molecule, and demonstrates different activities.
cell-mediated antibodies
The light chains in the immunoglobulins are specified as
reactions either kappa or lambda. Their structural details need not
be discussed here. Table 14.13 indicates some of the
CELL-MEDIATED HUMORAL
IMMUNITY IMMUNITY activities of the immunoglobulins.
Fig 14.74 Comparison of cell-mediated and humoral
immunity
each prospective lymphocyte has genes coding for
only one specific antibody
a
prospective
© © © ® © @
immature lymphocyte
from own body reacts
with antigen during
embryonic development
rendered incapable rendered incapable

of making antibody of making antibody
against own antigen against own antigen
(cell killed) (cell killed)
after birth
mitosis
mitosis
clone ofB
immunocytes
memory
cell
production of
antibody
4
Fig 14.75 Clonal selection theory of antibody production
536
Table 14.13 Classes of immunoglobulins and their biological
immunological memory (fig 14.76), and is responsible
activities. for the body’s secondary response which generally
ll ————————————
ee confers immunity on an animal against the specific
Immuno- Heavy antigen.
globulin chain Activity
ee
IgM mu First class of antibody to appear in the
14.14.5 Types of immunity
serum after injection of antigen. It initiates
the body’s primary response Natural passive immunity
IgG gamma 1 Principal antibody in serum. Initiates the Preformed antibodies from one individual are passed into
body’s secondary response another individual of the same species. This only affords
temporary protection against infection, for as the anti-
bodies do their job, or are broken down by the body’s
IgA alpha 1 Major class of antibody in external secre-
», 2 tions such as saliva, tears, bronchial and
natural processes, their number diminishes and protection
intestinal mucus. Operates as body’s first is slowly lost. For example, antibodies from a mother can
line of defence against bacterial and viral cross the placenta and enter her foetus. In this way they
antigens provide protection for the baby until its own immune
IgD delta Little secreted. It is membrane-bound, but system is fully functional. Passive immunity may also be
its function is as yet unknown conferred by colostrum, the initial secretion of the
IgE epsilon _ Possibly involved in reactions to allergens. mammary glands, from which antibodies are absorbed
Functions otherwise unknown from the intestines of the baby.
Acquired passive immunity
14.14.4 The clonal selection theory of Here antibodies which have been preformed in one
antibody formation individual are extracted and then injected into the blood of
This theory was developed by Jerne, Burnet, another individual which may or may not be of the same
Talmage and Lederberg in the 1950s and is generally species. For example, specific antibodies used for combat-
accepted as a working model for antibody production (fig ing tetanus and diphtheria are cultured in horses and later
14.75). Its essential points are as follows. injected into Man. They act prophylactically to prevent
(1) There exists in every individual a very wide range of tetanus and diphtheria respectively. This type of immunity
lymphocytes, each of which is capable of recognising is again short-lived.
only one specific antigen. Natural active immunity
(2) The specificity of an antibody (which is a protein) is
The body manufactures its own antibodies when exposed
determined by its amino acid sequence, which in turn is
to an infectious agent. Because memory cells, produced on
determined by the lymphocyte’s own unique DNA base
exposure to the first infection, are able to stimulate the
sequence. Therefore the ability to synthesise a specific
production of massive quantities of antibody when exposed
antibody is predetermined before the cell ever meets
to the same antigen again, this type of immunity is most
the antigen.
effective and generally persists for a long time, sometimes
(3) Small amounts of antibody are produced by each cell as
even for life (fig 14.76).
it matures and some of it becomes interlocked into its
plasma membrane where it acts as a receptor site for its Pp rimary response secondary response
complementary antigen.
(4) It is thought that if an immature lymphocyte meets with
an antigen corresponding to its antibody during first injection second injection
of antigen of antigen
embryonic life, then the lymphocyte is killed. There-
fore an animal will not synthesise antibody against its
own macromolecules and becomes self-tolerant.
(5) When antigen locks into the receptor site of a mature
lymphocyte, somehow the genetic apparatus of the
lymphocyte is stimulated to produce antibody. Contact
with the antigen is necessary for the lymphocyte to
differentiate and divide into a clone of plasma cells Antibody
inserum
concentration
synthesising antibody, and into memory cells.

(6) As all plasma cells are genetically identical, they all (aa 10 20 30 60 70 80 90
make the same antibody. Days
(7) The memory cells persist after the disappearance of the Fig 14.76 Primary and secondary response to an initial and
antigen, and in some way retain the capacity to be later dose of antigen. The secondary response is more rapid
stimulated by antigen if it reappears. This is called and intense than the first
337
Acquired active immunity Table 14.14 Blood groups.
ooo
This is achieved by injecting small amounts of antigen, A B AB

Blood group O
called the vaccine, into the body of an individual. The
whole process is called vaccination or immunisation. The Percentage of population 46% 42% 9% 3%
small dose of antigen is usually safe because the antigen is Agglutinogen = A B Att.
a+b ‘b a =
either killed or attenuated. This ensures that the individual Agglutinin
does not contract the disease itself, but is stimulated to
manufacture antibodies against the antigen. Often a
second, booster injection is given and this stimulates a
much quicker production of antibody which is long lasting
and which protects the individual from the disease for a
considerable time. Several types of vaccine are currently in
use.
(1) Toxoids. Exotoxins produced by tetanus and diphtheria
bacilli are detoxified with formaldehyde, yet their
antigen properties remain unimpaired. Therefore vac-
cination with the toxoid will stimulate antibody
production without producing symptoms of the dis-
ease.
(2) Killed organisms. Some dead viruses and bacteria are
able to provoke a normal antigen—antibody response
and are used for immunisation purposes.
(3) Attenuated organisms. Modified but living organisms
are injected into the body. They are able to multiply
without producing disease. Attenuation may be
achieved by culturing the organisms at higher tempera-
tures than normal or by adding specific chemicals to the - noagglutination as agglutination
culture medium for long periods of time. Attenuated

vaccines for tuberculosis, measles, rubella and Fig 14.77 Interactions between human blood groups. Cell
agglutinogens are denoted by capital letters, agglutinins by
poliomyelitis are now in general use. small letters
14.14.6 Blood groups will happen to the cells of the donor. If there is a likelihood
of them being agglutinated by the recipient’s plasma
When a patient receives a blood transfusion it
antibodies then transfusion should not take place.
is imperative that he receives blood that is compatible with
Fig 14.77 indicates the consequences of mixing different
his own. If it is incompatible, a type of immune response
blood groups together. Individuals with blood group O are
occurs. This is because the donor’s red cell membranes
termed universal donors because their blood can be given
possess mucopolysaccharides known as agglutinogens
which act as antigens and react with antibodies (aggluti- to anyone. It possesses cells which will not be agglutinated
nins) in the recipient’s plasma. The result is that the donor’s by the recipient’s plasma agglutinins. Although group O
cells are agglutinated (the cells link or attach to each other possesses a and b agglutinins there will be very little
when the antigens on the surface of their cells interact with agglutination of the recipient’s cells because the donated
the antibodies). Two agglutinogens exist and they are plasma is diluted so much by the recipient’s blood that it is
named A and B respectively. The complementary plasma ineffective in its agglutination activity. Individuals with
agglutinins are named a and b, and are present in the group AB can receive blood from anyone and are called
universal recipients. However, they can only donate to
plasma all of the time. They are not produced in response
to the donor’s agglutinogen as is the case in the immune blood group AB.
reactions already studied. A person with a specific
14.14.7 The rhesus factor
agglutinogen in the red cells does not possess the
corresponding agglutinin in the plasma. For example, Of the total population, 85% possess red cells
anyone with agglutinogen A in the red cell membranes has containing an agglutinogen called the rhesus factor and are
no agglutinin a in the plasma and is classified under blood termed rhesus positive. The remainder of the population
group A. If only B agglutinogens are present the blood lack the rhesus agglutinogen and are therefore regarded as
group is B. Should both agglutinogens be present the blood rhesus negative. Rhesus negative blood does not usually
group will be AB, whilst if no agglutinogens are present the contain rhesus agglutinins in its plasma. However, if rhesus
blood group is designated O (table 14.14). positive blood enters a rhesus negative individual the
When transfusion occurs it is important to know what recipient responds by manufacturing rhesus agglutinins.
538
The practical importance of this observation is made 14.14.8 Transplantation
obvious when a rhesus negative mother bears a rhesus
positive child. During the later stages of the pregnancy, Replacement of diseased tissues or organs by
fragments of the rhesus positive cells of the foetus may healthy ones is called transplantation and is a technique
enter the mother’s circulation and cause the mother to used increasingly in surgery today. However, when foreign
produce rhesus agglutinins. These can infiltrate the foetus tissue is inserted into or onto another individual it is
and destroy foetal red cells. Normally the agglutinins are rejected by the recipient because it acts as an antigen, so
not formed in large enough quantities to unduly affect the stimulating the immune response in the recipient. The
first-born child. However, subsequent rhesus positive following terms are used for the different kinds of
children can suffer chronic destruction of their red cells. A transplantation.
thesus baby is usually premature, anaemic and jaundiced, Autograft — tissue grafted from one area to another on the
and its blood needs to be completely replaced by a same individual.
transfusion of healthy blood. This treatment may now be Isograft — a graft between two genetically identical
undertaken whilst the baby is still in the womb. individuals such as identical twins.
Allograft — a tissue grafted from one individual to another
Protection against the rhesus individual of the same species but of different genetic
positive—negative reaction constitution.
(1) Consider a rhesus negative mother of blood group O, Xenograft — a graft between individuals of different species
such as from pig to Man.
carrying a rhesus positive child of any blood group
other than O. Should foetal cells enter the maternal Only allografting will be discussed here.
As blood is a fluid tissue, then simple blood transfusion
circulation, the mother’s a and b agglutinins will
can be regarded as an allograft. Here rejection results in
destroy them before the mother has time to manufac-
agglutination of the donor’s red cells as discussed earlier.
ture anti-rhesus agglutinins.
When rejection of tissue, such as skin, occurs the following
(2) If an intravenous injection of anti-rhesus agglutinins,
sequence of events takes place.
called anti-D, is given to a rhesus negative mother
(1) The skin allograft initially develops blood vessels in the
within 72 h of her giving birth, sensitisation of the
first 2-3 days and generally looks healthy.
rhesus negative mother by rhesus positive foetal cells is
(2) During the next six days its vascularisation decreases,
prevented. Apparently the anti-D attaches itself to the
and a great number of T lymphocytes and monocytes
foetal cells which are in the mother’s circulation and
(sections 14.14.1 and 14.11.2) gather in the vicinity of
affects them in such a way that they are not recognised
the graft.
by the mother’s antibody forming cells and hence the
(3) Two days later the graft cells begin to die and the graft
antibody process in the mother is not set in motion
is eventually cast off.
(fig 14.78).
Prevention of graft rejection
Foetal blood Maternal blood There are several means of preventing graft rejection
currently in use, listed as follows.
(1) Tissue matching — this is an obvious and necessary
precaution to take prior to any surgery.
(2) Exposure of bone marrow and lymph tissues to
Rhesus
disease aD X-irradiation tends to inhibit blood cell production and
(ea « aD
red cell therefore slows down rejection.
(3) Immunosuppression — here the principle is to use
agents which inhibit the entire activity of the immune
foetal cells s
destroyed — anaemia 5 system. When this occurs graft rejection is delayed, but
do3S D aD See
injection of the main problem with this technique is that the patient
= ae aD? anti-D within becomes susceptible to all other kinds of infections. It
72 h of birth
has also been shown that immunosuppression may
make the patients more prone to develop cancer.
Prevention aD aD
of Rhesus
If the disadvantages of non-specific immunosuppression
are to be overcome then ways must be found of suppressing
Rh
disease aD
immunocyte
only the T cells response to the antigens of the graft. In this
way the rest of the patient’s immune system would remain
unable to recognise foetal
cells so no antibody
unimpaired and continue to function normally. The most
be
promising approach is to treat the patient (or his bone
Fig 14.78 The basis of Rhesus disease anditsprevention.
marrow) with antibody that recognises and destroys the T
(From Burnet (1978) Genes, dreams and realities, Pelican) lymphocytes responsible for the graft rejection.
539
14.14.9 The interferon system of virus particles. Thus interferon, itself, is not anti-viral
but brings about an ‘anti-viral state’ which involves cellular
Interferon is a generic term which it is now changes that then inhibit virus replication. The production
realised applies to a number of proteins with similar of interferon can be stimulated by a whole variety of factors
properties. Human interferons have been divided into apart from virus attack, such as some inactivated viruses,
three groups a, 6 and y depending on which type of cell double-stranded RNA, man-made double-stranded oligo-
they are produced by. The interferon a group consists of nucleotides and bacterial endotoxins. It would seem that
several species. They are all of similar molecular mass interferon is produced as a result of an ‘abuse’ of the cell.
(about 20 000). Interferon is biologically extremely active. Mouse
The most widely studied property of interferon is its interferon has an activity of 2 x10’ units per milligram of
ability to ‘interfere’ with the replication of viruses. It is protein, where one unit will bring about a 50% reduction in
produced in mammalian and avian cells in response to viral virus production. This means that as little as one molecule
attack and appears to be an effective anti-viral agent on per cell may be necessary to render acell resistant to virus
most types of cell against all viruses to a greater or lesser infection.
extent. The interferons have a wide range of other effects
When a cell is attacked by a virus, a number of things including inhibiting cell growth. Recent results show that
happen. The virus will multiply, but at the same time the this effect may mean that they will be useful as anti-cancer
host cell is stimulated to produce interferon (fig 14.79). agents under certain circumstances. There are also effects
This is released from the cell and when it comes into on the immune system and marked changes in the
contact with adjacent cells it renders them resistant to virus properties of cell membranes.
attack. To do this interferon triggers a series of events Thus it would seem that the interferon system may play
which lead to an inhibition of viral protein synthesis, viral an important role in the protection of the body against
RNA transcription and in some cases assembly and release viruses.
cell rendered
peas
_--~ interferon
a release resistant to
virus
interferon
production_
7
. / .
interferon virus
stimulation production
I Beene
interferon
release
cell rendered
resistant to
virus
virus attack
Fig 14.79 Interferon production and activity
540
Chapter Fifteen
Coordination and control in plants

Plants, like animals, need some form of internal coordina- Table 15.1 Examples of tropisms.
tion if their growth and development is to proceed in an
orderly fashion, with suitable responses to their environ- Stimulus Type of tropism Examples
ment. Unlike animals, plants do not possess nervous
systems and rely entirely on chemical coordination. Their Light Phototropism Shoots and coleoptiles
responses are therefore slower and they often involve positively phototropic
Some roots negatively photo-
growth. Growth, in turn, can result in movement of an tropic, e.g. adventitious roots
organ. In this chapter plant movements will be examined of climbers like ivy
before studying the various ways in which plants coordinate Gravity Geotropism Shoots and coleoptiles
their activities. negatively geotropic
Roots positively geotropic
Rhizomes, runners, dicotyle-
donous leaves diageotropic*
15.1 Plant movements Lateral roots, stem branches
plagiogeotropic*
It is a characteristic of all but a few unicellular Chemical Chemotropism Hyphae of some fungi
positively chemotropic, e.g.
plants that they do not show locomotion (movement of the Mucor
entire organism). However, movements of individual plant Pollen tubes positively
organs are possible and are modified by the sensitivity of chemotropic in response to
the plant to external stimuli. Movements induced by chemical produced at micro-
pyle of ovule
external stimuli fall into three categories: tropisms (tropic
movements), nasties (nastic movements) and taxes (tactic Water Hydrotropism Roots and pollen tubes
(special kind of positively hydrotropic
movements). chemotropism)
Solid surface Haptotropism Tendrils positively hapto-
15.1 Whatis the basic reason for the fact or touch (thigmotropism) tropic, e.g. leaves of pea
Central tentacles of sundew, an
that animals show locomotion whereas plants do insectivorous plant positively
not? haptotropic (section 9.12.2)
Air Aerotropism Pollen tubes negatively
(oxygen) (special kind of aerotropic
15.1.1 Tropisms chemotropism)
A tropism is a movement of part of a plant in
response to, and directed by, an external stimulus. The *diageotropism: growth at 90° to gravity, that is horizontal growth.
plagiogeotropism: growth at some other angle to gravity, that is not
movement is almost always a growth movement. Tropic horizontal or directly towards or away from gravity.
responses are described as positive or negative depending
on whether growth is towards or away from the stimulus
respectively. Some examples of tropisms are shown in table 15.1.2 Taxes
i he A taxis is a movement of an entire cell or
organism (that is locomotion) in response to, and directed
15.2 Complete a fourth column to table by, an external stimulus. As with tropisms they can be
15.1 to show for each response how it is advan- described as positive or negative, and can be further
classified according to the nature of the stimulus. Note that
tageous to the plant involved.
this kind of movement is not confined to plants. Examples
15.3 How could growth be modified to are given in table 15.2.
cause a tendril to coil round a solid object?
15.4 Design an experiment to demon-
strate the preferred light intensity of Euglena or
Phototropism and geotropism will be discussed in more Chlamydomonas in phototaxis.
detail later in this chapter (sections 15.2.1 and 15.2.2).
541
Table 15.2 Examples of taxes. The ‘sleep movements’ (nyctinasty) of certain flowers
ee———————————————— and leaves, whereby they open and close in response to
light intensity (photonasty) or temperature (thermonasty),
Se
Stimulus Taxis Examples

teed ceed So = aea egee are nastic because they are merely triggered by external
Light Phototaxis Positive: Euglena swims to- stimuli and are directed internally. Some flowers, such as
wards light, chloroplasts
move towards light crocus and tulip, close at night by the lower sides of petals
Negative: Earthworms, growing more rapidly (hyponasty). Opening is caused by
blowfly larvae, woodlice and more rapid growth of the upper sides (epinasty). Many
cockroaches move away from leaves or leaflets, particularly of leguminous plants, such as
light clover and Mimosa, have structures called pulvini. A
Chemical Chemotaxis Positive: Sperms of liver- pulvinus is a swelling at the base of a petiole or leaflet which
worts, mosses and ferns swim
possesses large parenchyma cells. Rapid turgor pressure
towards substances released
by the ovum, motile bacteria changes in these cells result in the pulvinus acting as a hinge
move towards various food and bringing about movement.
substances Haptonastic movements, where the stimulus is touch, are
Negative: Mosquitoes avoid among the most fascinating of plant movements, since they
insect repellent
include rapid and elaborate responses. The well-known
Air (oxygen) Aerotaxis Positive: motile aerobic ‘sensitive plant’ Mimosa pudica is sensitive to touch as well
(special kind of bacteria move towards oxygen
chemotaxis)
as a variety of other stimuli. It exhibits normal sleep
Positive: Planula larvae of
movements but responds very rapidly to shock (seismo-
Gravity Geotaxis
some coelenterates swim nasty) such as a sharp blow, injury, or sudden change in
towards sea bed temperature or light intensity. If leaflets at the tip are
Negative: Ephyra larvae of shocked, they fold upwards in seconds. If the stimulus is
some coelenterates swim strong or sustained, successive pairs of leaflets fold up and
away from sea bed
the stimulus eventually passes through the whole leaf,
Magnetic Magnetotaxis Certain motile bacteria resulting in the petiole drooping (fig 15.2). The stimulus
field
will pass in the reverse direction if the stem is stimulated.
Resistance Rheotaxis Positive: Planaria move
Fig 15.2 shows the three sites at which pulvini are found.
against water current, moths
and butterflies fly into the The stimulus is thought to be transmitted by a hormone
wind moving through the xylem; electrical changes are associ-
ated with its passage but there is no nervous system.
Insectivorous plants show some of the most elaborate
15.5 Fig 15.1 illustrates the distribution of movements in the plant kingdom, some of which are
motile bacteria 10 min after being placed under a haptonastic.
cover-slip with a filament of a green alga. (a) Name a
genus of a suitable alga for the experiment. (b) Put 15.1.4 Kinesis
forward a hypothesis to account for the final A type of locomotory response not so far
distribution of the bacteria. (c) How could you check — mentioned is kinesis. Since this is virtually confined to the
your hypothesis? animal kingdom it is discussed in chapter 16 with animal
behaviour.
cover-slip ee Plant growth substances

filament of
a green alga Chemical coordination in animals is controlled
by hormones, chemicals working in minute concentrations
at sites some distance from their sites of synthesis. Plants
are coordinated by chemicals which do not necessarily
Fig 15.1 Distribution of motile bacteria on a slide
move from their sites of synthesis and hence, by definition,
should not always be termed hormones. In view of this and
15.1.3 Nasties
because their effects are usually on some aspect of growth,
A nasty is a non-directional movement of part they are called growth substances. It is also important to
of a plant in response to an external stimulus. The direction stress that the precise mechanisms of action of the plant
of movement is determined by the structure of the growth substances so far discovered are far from clear and
responding organ. Movement is the result of growth or a that analogies with the better understood animal hormones
turgor change and small movements are typically amplified may be misleading. Plant growth substances are certainly
by the particular positioning of the responding cells. essential for plant development, but to what extent they act
542
petiole drooping
petiole leaflets folded
leaflets in upwards
normal
position
Fig 15.2 Response of the ‘sensitive plant’ (Mimosa pudica) to shock. (left) before (right) after
as ‘triggers’ for changes in growth or as ‘integrating discussed separately, and then key stages in the life cycle of
chemicals’, modifying processes triggered by other un- a plant will be discussed to emphasise the fact that growth
known events, is not certain. It should be borne in mind substances often work in association with each other to
that growth can be divided into the three stages of cell achieve their effects.
division, cell enlargement and cell differentiation (spe-
cialisation) and that these have particular locations in
plants (section 21.6). It can be expected, therefore, that the 15.2.1 Auxins and phototropism
action and distribution of different plant growth substances
will reflect this. Five major types of growth substance are Discovery of auxins
recognised: auxins, gibberellins, cytokinins, abscisic acid The discovery of auxins was the result of investigations into
and ethene (ethylene). Generally speaking, cytokinins are phototropism that began with the experiments of Charles
associated with cell division, auxins and gibberellins with Darwin and his son Frances. Using oat coleoptiles as
cell enlargement and differentiation, abscisic acid with convenient material (fig 15.3), they showed that the growth
resting states (such as lateral buds) and ethene with of shoots towards light was the result of some ‘influence’
senescence (ageing). being transmitted from the shoot tip to the region of growth
In this chapter, each type of growth substance will first be behind it. Some of their experiments are summarised in
first leaf
Soe coleoptile — hollow,
through tip colourless, protective
of coleoptile sheath
first leaf— rolled and

coleoptile green
ground level
coleoptile
stem apex
grain
node
g-
mesocotyl
(c) (d)
Fig 15.3 Germination of a typical grass seedling: (a) (b) and (c) stages in germination, (d) section of coleoptile at stage (b)
543
fine black sand
untreated control tip removed Opaque cap
direction of light
iC (7
(a) (b) (c) (d)
Fig 15.4 Darwin's experiments on phototropism using oat coleoptiles. (a), (b), (c) and (d) separate experiments showing
treatment (left) and result (right)
fig 15.4, the diagrams representing results obtained from In 1928 the Dutch plant physiologist Went finally proved
many seedlings. the existence of a chemical transmitter. His aim had been to
intercept and collect the chemical as it passed back from the
15.6 (a) List carefully the conclusions you tip and to demonstrate its effectiveness in a variety of tests.
could draw from experiments (a)-(d) in fig 15.4, He reasoned that a small diffusing molecule should pass
given that the curvature was due to growth in the freely into a small block of agar jelly, whose structure is
region behind the tip. (b) Why was experiment (c) such that relatively large spaces exist between its mole-
-necessary, bearing in mind the result from experi- cules. Fig 15.6 illustrates some of his experiments.
ment (b)?
15.9 What would you : conclude from the
If the tropic response is analysed in terms of the following: results shown in fig 15.6?
stimulus — receptor — transmission — effector —
response, then the largest gap in our knowledge remains 15.10 What result would you expect if the
the nature of the transmission. In 1913 the Danish plant treated block had been placed on the right side of the
physiologist Boysen-Jensen added to our knowledge. Fig decapitated coleoptile in experiment (6)?
15.5 summarises some of his experiments.
A further experiment of note carried out by Went is
15.7 What extra information is provided illustrated in fig 15.7. In control experiments the tip was
by Boysen-Jensen’s experiments? exposed to uniform light or darkness before transfer of agar
15.8 If these experiments were repeated blocks, and the degree of curvature induced by blocks A
in uniform light, draw diagrams to show what results and B was the same. Unilateral illumination of the tip,
you would expect. Give reasons for your answers. however, resulted in unequal distribution of the chemical
in blocks A and B (fig 15.7). Not only does this support the
untreated mica plate — thin,

control impermeable barrier
direction of light
Fig 15.5 Boysen-Jensen’s experiments

on phototropism using oat coleoptiles,
(a), (b) and (c), separate experiments
(a) (b) showing treatment (left) and result (right)
544
transfer tip to agar
block and leave for
several hours
then transfer block

to freshly decapitated ». growth substance
coleoptiles
—>
untreated agar block

take normal remove tip untreated agar block
.coleoptile
very little very little

growth growth
(a) control for (a) (b) control for (b)

Fig 15.6
> Went's experiments, i (a) and (b),Y separate experiments showin 1g treatment i
(left) and result (right). C
experiments are shown alongside. All treatments were carried out in darkness or en tea al
coleoptile tip degree of curvature proportional
to amount of chemical
direction
of light
ea 8 less curvature
than A
thin metal |
plate |A |
A, B- two
agar blocks
Fig 15.7 Went’s experiment showing effect of unilateral light on distribution of the chemical (auxin)
conclusions from Boysen-Jensen’s experiments about the studied and that others appear to be more complex. Also,
effect of light on the distribution of the chemical, but it there is little evidence for the development of auxin
shows howa test for measuring the amount of the chemical gradients in the critical period. before the response is
present, that is a bioassay, can be set up. A bioassay is an measured.
experiment in which the amount of a substance is found by
measuring its effects in a biological system. Went showed Eas Hight
that the degree of curvature of oat coleoptiles was directly
proportional to the concentration of the chemical (at
normal physiological levels). zone of y movement of auxin
The chemical was subsequently named ‘auxin’ (from the elongation
Auxin moves laterally away
Greek auxein, to increase). In 1934 it was identified as from light. It is not
indoleacetic acid (IAA). IAA was soon found to be widely inactivated by light.
Greater concentration of
distributed in plants and to be intimately concerned with degradation auxin on the shaded side
cell enlargement. Fig 15.8 summarises present beliefs of auxin by stimulates cell elongation
enzymes thereby causing curvature
concerning the movement of IAA during unilateral
illumination of coleoptiles. It should be pointed out, Fig 15.8 Hypothesis for effect of unilateral illumination on
however, that the coleoptile is the simplest system so far distribution of auxin in a coleoptile
545
Structure of IAA 6 petri dishes + lids
The structure of IAA is shown in fig 15.9. 5 x 5cm? graduated pipettes
25 cm? measuring cylinder or 10 cm® graduated pipette
CH, COOH coleoptile cutter (fig 15.10)
paint brush
stock IAA solution (1 g dm“°) (IAA is not readily soluble
N
in water and is therefore first dissolved in ethanol:
dissolve 1g of IAA in 2cm?® ethanol and dilute to
H
900 cm? with distilled water. Warm the solution to 80 °C
indole acetic acid
3 ;
and keep at this temperature for 5 min. Make up to 1 dm°
(ethanoic acid) with distilled water. Adjust quantities according to final
Fig 15.9 Structure of IAA (indoleacetic acid) volume required.)
2% sucrose solution
Other chemicals with similar structures and activity were distilled water
soon isolated, and some such substances have been
making a whole class of plant growth Method
synthesised,
substances called auxins. Some of these are discussed (1) Take six test-tubes and six petri dishes and label
below in section 15.2.5. them A-F.
(2) Add 18 cm? of 2% sucrose solution to each test-tube.
Synthesis and distribution of auxins (3) Using a clean 5cm*® pipette, add 2cm® of IAA
Auxins are made continuously in the shoot apex and young solution to tube A and mix the two solutions
leaves. Movement away from the tip is described as thoroughly.
basipetal (from apex to base of the organ) and polar (in one (4) Using a fresh pipette transfer 2 cm? of solution from
direction only). It moves, apparently by diffusion, from cell tube A to tube B and mix the contents of tube B
to cell and is eventually inactivated and degraded by thoroughly.
enzymes. Long-distance transport can also occur via the (5) Using a fresh pipette each time, transfer 2 cm? from
vascular system (mainly phloem) from shoots to roots. A tube B to tube C, mix, then 2 cm? from tube C to tube
little auxin is probably made in roots. The effects of D, mix, then 2 cm? from tube D to tube E.
different auxin concentrations on shoot growth can be (6) Add 2 cm? distilled water to tube F.
investigated by means of an experiment such as experiment (7) ‘Transfer the solutions from tubes A-F to petri dishes
fort A-F.
(8) Take 60 germinated oat seedlings and cut 10 mm
lengths of coleoptile, starting about 2 mm back from
Experiment 15.1: To investigate the effects of the tips. Use a double-bladed cutter with the blades
indoleacetic acid (IAA) on the growth of oat held exactly 10 mm apart by a series of washers, two
coleoptiles nuts and two bolts (see fig 15.10). If the tips of the
The aim of the experiment is to investigate the effect of coleoptiles are placed in a line, several lengths can
various concentrations of [AA on growth of oat coleoptiles. be cut simultaneously.
Growth is affected by white light and therefore cutting and (9) Using a paint brush, transfer 10 lengths of coleoptile
transferring of coleoptiles during the experiment should be to each dish avoiding cross contamination of the
carried out under red light or in the minimum amount of solutions (the greater the number of coleoptiles used,
light possible. Sucrose solution is used in the experiment the more statistically valid the results).
as energy will be required for growth, and sucrose is an (10) Place a lid on each and incubate the dishes at 25 °C
energy source. The apical tip (3 mm) of each coleoptile is for three days in the dark.
removed in order to prevent natural auxins produced by (11) Remeasure the lengths of coleoptiles as accurately
the coleoptile from having an effect on growth. as possible.
Materials washers nut

germinating oat seedlings with coleoptiles at least
1.5 cm long (Soak 100 oat grains in water overnight, cut cut bolt
place the soaked seeds on damp paper towelling in a
dish, cover the dish with aluminium foil and place in the I |
dark to germinate (five days in an incubator at 20 °C). In

—
razor
blades
order to obtain the 60 coleoptiles required for each coleoptiles
experiment, at least 100 grains should be soaked to <<—> arranged side
10mm by side
allow for germination failure.)
6 test-tubes in a test-tube rack Fig 15.10 Cutting 10 mm lengths of coleoptiles
546
20) S
rotating arm cork disc radicle
& 150 saturated

3 atmosphere
Ss
& pin
electric
2 100 shoots tight-fitting
on dei
Ss
&
oO
2oO
es) ees cotton wool or filter paper
roots
Fig 15.12 (above) Klinostat showing broad beans after

0 several days growth with rotation
<
cS Fig 15.13 (below) Effect of gravity on distribution of auxin
-
from a horizontal coleoptile tip
= 50
v coleoptile tip agar block
on
s laid horizontally
cS
o
= 100
a LOSS OR On LO 1052) alo}! l 10 100 1000
_ Auxin concentration/parts per million percentage of total auxin in each block
NB logarithmic scale for auxin concentration determined by Went’s bioassay
Fig 15.11 Effect of auxin concentration on growth responses

of roots and shoots. Note that concentrations of auxin which
stimulate shoot growth inhibit root growth
surface of an intact coleoptile would stimulate greater cell
(12) Ignoring the largest and smallest figures for each elongation here, and hence upward growth.
dish, calculate the mean (average) length. Decapitation of a root tip removes its sensitivity to
(13) Plot a graph of mean length (vertical axis) against IAA gravity, but it is not so easy to demonstrate movement of
concentration in parts per million (horizontal axis). auxins in roots because very low concentrations are
present, and these do not give convincing results in the
1 5.11 : What is the concentration in parts bioassay described. An interesting result though is
per million (ppm) of IAA in each petri dish (1 g dm~® obtained from the experiment shown in fig 15.14.
= 1000 ppm)? higher auxin
concentration
on left side
(14) Comment on the results and compare them with of block
fig 15.11. More accurate results can be obtained
by combining class results. COLEOPTILE
OR SHOOT
15.2.2 Auxins and geotropism
It is a common observation that roots are
positively geotropic, that is grow downwards, and shoots decapitated coleoptile
are negatively geotropic, that is grow upwards. That agar block or shoot
gravity is the stimulus responsible can be demonstrated by
coleoptile tip
using a piece of equipment called a Klinostat (figel5.12): placed horizontally
As the chamber rotates, all parts of the seedling receive, in
turn, equal stimulation from gravity. A speed of four
revolutions per hour is sufficient to eliminate the one-sided eeee8 auxin ROOT
effect of gravity and to cause straight shoot and root

growth. A non-rotating control shows the normal response
to gravity, with shoot growing up and root growing down. It decapitated root
is important to ensure even illumination during the TREATMENT RESULT
experiment (or to carry it out in darkness) so that there can
be no directional response to light. Fig 15.14 Effect of uneven auxin distribution on growth of
decapitated coleoptile and root
The involvement of auxins in geotropism is demons-
the
trated by the experiment shown in fig 15.13, which uses
techniques introduced by Went. Auxin moves out of the 15.12 What can you conclude from the
horizontally placed coleoptile tip but moves downwards as experiment shown in fig 15.14?
it does so. The greater auxin concentration on the lower
547
The gravity-sensing mechanism
The question now arises as to how the gravity stimulus is
SHOOT
detected. Darwin showed that removal of the root cap, the
group of large parenchyma cells that protect the root tip as
it grows through the soil, abolishes the geotropic response.
ROOT A section of the root cap reveals the presence of large
starch grains contained in amyloplasts within the cells
(fig 15.16).
high auxin concentration high auxin concentration It was suggested as long ago as 1900 that these cells act as
stimulates growth on lower inhibits growth on lower
side so shoot grows upwards side so root grows downwards statocytes, that is gravity receptors, and that the starch
grains are statoliths, structures which move in response to
Fig 15.15 Hypothesis for redistribution of auxin in a gravity. The so-called ‘starch-statolith hypothesis’ pro-
horizontally placed seedling
poses that sedimentation of the starch grains through the
cells occurs so that they come to rest on the lower sides of
Observations of this type led to the hypothesis summarised the cells with respect to gravity (fig 15.16). In some
in fig 15.15, which suggests that the opposite responses of unknown way this affects the distribution of growth
roots and shoots are due to different sensitivities to auxin. substances which are known to be produced sometimes in
Modifications of the hypothesis in the light of recent the root apex, sometimes in the root cap and sometimes in
findings are discussed later in this section. both. There is much evidence to support this hypothesis.
The different sensitivity of roots to auxin (fig 15.11) All plant organs which are sensitive to gravity contain
could also explain the negative phototropism shown by statocytes. They are found, for example, in the vascular
some; the higher accumulation of auxin on the shaded side bundle sheaths of shoots. Plants from which the starch
would cause inhibition of growth with the result that cells grains have been removed by certain treatments lose their
on the light side would elongate faster and the root would sensitivity to gravity, but regain it if allowed to make more
grow away from light. starch.
An important aspect of plant growth regulation has thus
been revealed. It is not only the nature of the growth 15.13 How is this mechanisi m of
2
substance which is relevant (qualitative control) but the detection similar to that in an
amount of that substance (quantitative control).
Oy s
os
gee mitochondria
Fig 15.16 Electron

micrograph of section of root
cap showing amyloplasts with «
starch grains located at the
bottoms of the cells
Modern hypotheses on geotropism loosening, possibly by an enzyme with a low pH optimum.
In coleoptiles the gravity response seems to be mediated by The ability to maintain a high osmotic potential inside the
auxins as described above, but with most shoots a geotropic cell, and availability of water to enter the cell and generate
response is still obtained if the tip is removed, and there is turgor pressure, are also necessary.
still doubt as to whether movement of auxin is involved. In
roots, auxin redistribution does occur, but probably not 15.2.4 Other effects of auxins
dramatically enough to account for the observed changes in
Apart from stimulating cell elongation and
growth rates. Transmission of a growth inhibitor from the
hence shoot growth, auxins have a number of other
root cap to the zone of elongation has been shown, but this
important roles in the plant, summarised in table 15.4.
is not necessarily auxin. Several groups of workers have
Further details of their roles in differentiation, apical
been unable to find auxin in the root caps of maize dominance, abscission and fruit growth are given later
seedlings, a common experimental plant. Instead, abscisic under the appropriate headings.
acid, a well-known growth inhibitor, has been found.
Ethene, another growth inhibitor, could also be involved.
15.2.5 Commercial applications of auxins
Finally, gibberellins (growth promoters) have been found
in higher concentrations than normal in the rapidly growing Discovery of IAA led to the synthesis by
sides of both shoots and roots when they are geotropically chemists of a wide range of active compounds with similar
stimulated. structure. Synthetic auxins have proved commercially
useful in a variety of ways. They are cheaper than IAA to
15.14 What can you conclude from the produce, and often more physiologically active because
experiments shown in fig 15.17? Controls, using plants generally do not have the necessary enzymes to
untreated agar, showed no curvature. When IAA was break them down. Table 15.3 gives some examples with
used instead of abscisic acid no significant curvature their structures and summaries of their uses. Chlorine
was obtained. substitutions in the structures often increase activity. Fig
15.18 shows the effect of treating tomatoes with a
fruit-setting auxin.
decapitated maize root
(a) fag he
agar block containing
abscisic acid (used also
in (b), (c) and (d) below) strong
downward
been curvature
arrier
(b) Ha
7 <8 slight upward

(os dias ale ae curvature
barrier
—— ak S strong upward
(d) ps oe curvature
Fig 15.17 Effect of abscisic acid on geotropic response to

decapitated roots. (Based on experiments by Pilet, 1975. )
15.2.3 Mode of action of auxins

The effect of auxins on cell enlargement is now
reasonably well understood. During cell extension the rigid
cellulose framework of the cell wall must be loosened.
Extension then occurs by a combination of osmotic
swelling as water enters the cell and by the laying down of
new cell wall material. The orientation of the existing
cellulose microfibrils probably helps to determine the
direction of extension. ‘Wall loosening’ is induced by acid
conditions, and by auxins. In 1973 four different groups of Fig 15.18 The three large trusses were set by spraying with
workers all demonstrated that auxins stimulate hydrogen beta naphthoxy acetic acid. The small one in the bottom
pH
ion (proton) secretion. This causes a lowering of left-hand corner was not sprayed — and produced only one
wall normal size tomato
outside the cell (increase in acidity) and hence
549
Table 15.3 Commercial applications of auxins.
Type of Auxin Examples with structures Uses
Indoles and NAA (naphthalene acetic acid) Fruiting — help natural fruit set; sometimes cause fruit setting in
hthyls absence of pollination (parthenocarpy)
ee Cee Pe (compare with IAA fig 15.9) Rooting hormone — promote rooting of cuttings
Indole propionic acid

plea a0
N
Phenoxyacetic 2,4-D (2,4-dichlorophenoxyacetic acid) Selective weedkillers — kill broad-leaved species (dicotyledons).
acids* 7A OL COOH Used in cereal crops and on lawns. Also in conifer plantations
for scrub clearance (conifers unaffected). 2,4-D/2,4,5-T mix used in
Vietnam war by US as the defoliant ‘Agent Orange’
Cl Cl
Potato storage — inhibit sprouting of potatoes
2,4,5-T (2,4,5-trichlorophenoxyacetic acid) Fruiting — prevent premature fruit drop (retard abscission)
as above but extra chlorine atom in 5 position
of ring
MCPA (2-methyl-4-chlorophenoxyacetic acid)
as 2,4-D but methyl group (CH,) in 2 position
of ring instead of chlorine
Benzoic acids 2,3,6-trichlorobenzoic acid Powerful weedkillers
Cl
2 : COOH
: Cl
cl
2,4,6-trichlorobenzoic acid
as above except chlorine atom in 4 position
instead of 3 position of ring
*Cause plants to outgrow themselves. Growth distorted. Respiration excessive. 2,4,5-T is being phased out owing to concern about its contaminant,
dioxin, which is the most toxic substance known to Man. Dioxin causes, for example, cancer, foetal abnormalities and a particularly severe form of acne
called chloracne.
15.2.6 Gibberellins Structure of gibberellins

Discovery of gibberellins All are terpenes, a complex group of plant chemicals
related to lipids; all are weak acids and all contain the
During the 1920s a team of Japanese scientists at the
‘gibbane’ skeleton (fig 15.19).
University of Tokyo was investigating a particularly
damaging worldwide disease of rice seedlings, caused by gibbane skeleton
the fungus Gibberella (now called Fusarium). Infected
Fig 15.19
seedlings became tall, spindly and pale and eventually died
Structures of
or gave poor yields. By 1926 a fungal extract had been gibbane skeleton
isolated which induced these symptoms in rice plants. An and gibberellic
active compound was crystallised by 1935 and a further two acid (GA3)
by 1938. These compounds were called gibberellins, after
gibberellic acid (GA,)
the fungus. Language barriers and then the Second World
eeetags
War delayed the initiation of work in the West, but
immediately after the war there was competition between
British and American groups to isolate these chemicals. In
1954 a British group isolated an active substance which they
called gibberellic acid. This was the third, and most active,
gibberellin (GA;) isolated by the Japanese. Gibberellins H,
ec COOH CH,
were isolated from higher plants during the 1950s, but the
chemical structure of GA; was not completely worked out Synthesis and distribution of gibberellins
until 1959 (fig 15.19). Now more than SO naturally Gibberellins are most abundant in young, expanding
occurring gibberellins are known, all differing only slightly organs, being synthesised particularly in young apical
from GA,. leaves (possibly in chloroplasts), buds, seeds and root tips.
550
They migrate after synthesis in a non-polar manner, that is
up or down the plant from the leaves. They move in phloem complete digestion of itssubstrate? (d) Why is
and xylem. a-amylase so important in cereal seeds?
15.16 Explain the role of storage proteins
Effects of gibberellins
in the aleurone layer by reference to fig 15.21.
Like the auxins, the principal effect of gibberellins is on
stem elongation, mainly by affecting cell elongation. Thus
genetically dwarf varieties of peas and maize are restored
to normal growth and dwarf beans can be converted into
runner beans (fig 15.20). Stem growth of normal plants is Experiment 15.2: To test the following two
promoted. Further information, relating to interaction hypotheses, (a) that gibberellin can stimulate
breakdown of starch in germinating barley
with auxins, is given in section 15.3.
grains and (b) that gibberellin is produced in
One of the classic effects of gibberellins, which has been the embryo
much studied in an attempt to understand their mechanism
of action, is the breaking of dormancy of certain seeds, The presence of amylase in barley seeds can be detected
notably of cereals. Germination is triggered by soaking the by placing a cut seed on the surface of agar containing
seed in water. After imbibing water the embryo secretes starch. If the surface is moist, amylase will diffuse from the
gibberellin which diffuses to the aleurone layer, stimulating seed and catalyse digestion of the starch. Addition of
synthesis of several enzymes, including o-amylase (fig iodine to the agar would stain remaining starch blue-black,
15.21). These catalyse the breakdown of food reserves in revealing a clear ‘halo’ of digestion around the seed. The
the endosperm and the products of digestion diffuse to the size of this circular zone gives a rough indication of how
embryo, where they are used in growth. much amylase was present. In practice, sterile handling
techniques are an important precaution because contami-
15.15 (a) What is the substrate of a- nating bacteria and fungi may also produce amylase.
With careful experimental design, further hypotheses
amylase? (b) What
is the product of the reaction it
_ catalyses? (c) What other enzyme is required to may be tested (see, for example, Coppage, J. & Hill, T. A.
(1973) J. Biol. Ed. 7, 11-18).
|
, fluence of gibberellic [
i acidt (GA) on the growth of variety t
Meteor dwarf pea. . TiThe plant ton on the left received no
i
typical aah habit. The remaining plants were treated with GA; the dose per plant in microgra is shown.
ms
BA Metae
With doses up to 5 micrograms there is increased growth of the stems with increase in GA dosage. This is the principle of the
dwarf pea assay of gibberellins
py!
Fig 15.21 Role of gibberellin in
mobilising food reserves of barley
grain during breaking of dormancy
husk or fruit coat |
(pericarp) fused with
seed coat (testa)
aleurone layer — three cells

thick in barley, contains
protein
starch ___ maltose
maltase
maltose ___ glucose
starchy endosperm
scutellum (absorptive organ)

AMINO
acids
———&
_
coleoptile + shoot
gibberellin embryo
[==> diffusion synthesis
es biochemical
reaction
root
water ve
Materials (per student) and ‘non-embryo’ halves can be separated (see below)
white tile and stored under dry conditions in a fridge for a
scalpel maximum of 2-3 days. Wheat grains are naked and do
forceps not need dehusking, and may be used as a substitute for
50 cm® beaker barley.
labels or chinagraph pencil Method
iodine/potassium iodide solution
(1) Take two starch agar, and two starch—gibberellin agar
sterile distilled water in sterile flasks (3)
5% sodium hypochlorite solution or commercial sterilis- plates appropriately labelled +GA or —GA. These
ing fluid, e.g. Milton’s or 70% alcohol have been sterilised. Label one of each type ‘embryo’
two starch agar plates (sterile): 1% agar containing and the other ‘non-embryo’.
0.5% starch poured to a depth of about 0.25 cmin sterile (2) Cut at least two dry barley grains transversely in half
Petri dishes (fig 15.22) on atile, thus separating into ‘embryo’ and
two starch—gibberellin agar plates: as above but add ‘non-embryo’ halves.
gibberellin (GA,) to the agar before autoclaving at the
concentration of 1 cm* of 0.1% GA, solution per 100 cm?
agar (final concentration 10 ppm GA,) (Gibberellin does ah
not dissolve readily in water and is best dissolved in embryo
ethanol; some of the GA, is destroyed during autoclav- |
ing but the seeds are sensitive to concentrations as low
as 10°° ppm GA,.)
dehusked barley grains: to dehusk barley grains, soak |
them in 50%(v/v) aqueous sulphuric acid for 3-4 h and
then wash thoroughly (about ten times) in distilled water;
violent shaking of the grains in a conical flask removes
‘non-embryo half?
most of the husks; grains should be used immediately, ‘embryo half?
since soaking starts germination. Alternatively, ‘embryo’ _ Fig 15.22 Cutting a barley grain for experiment 15.2
352
(3) Sterilise the halves in 5% sodium hypochlorite solution often results in short (dwarf), sturdy plants with deep green
for 5 min. Then wash in three changes of sterile leaves and sometimes greater pest and disease resistance.
distilled water in sterile flasks. They take up less space and may in the future lead to higher
(4) Using forceps sterilised by rinsing in 70% alcohol, yields per acre; also they are less inclined to blow over.
place the halves immediately in the relevant dishes,
cut face downwards, with minimal lifting of lids as 15.2.7 Cytokinins
follows:
Discovery of cytokinins
—GA +GA —GA +GA During the 1940s and 1950s efforts were made to perfect
embryo 3 embryo 3 non-embryo 3 non-embryo } techniques of plant tissue culture. Such techniques provide
an opportunity to study development free from the
(5) Incubate for 24-48 h at 20-30 °C. influence of other parts of the organism, and to study
(6) Test for presence of starch in each dish by flooding effects of added chemicals. While it proved possible to
surfaces of agar with I,/KI solution. Draw the final keep cells alive, it was difficult to stimulate growth. During
appearance of each dish. Discuss the results. the period 1954-6 Skoog, working in the USA, found that
coconut milk contained an ingredient that promoted cell
15.17 Using starch agar it is often pos- division in tobacco pith cultures. Coconut milk is a liquid
sible to demonstrate an association of amylase endosperm (food reserve) and evidence that it contained
activity with fingerprints. Suggest reasons for this growth substances had already been obtained in the 1940s
association. — when it had been studied as a likely source of substances to
promote embryo growth. In a search for other active
15.18 What further experiment could you
substances a stale sample of DNA happened to show
do, given the facilities, to prove that gibberellin
similar activity, although a fresh sample did not. However,
causes synthesis of new amylase rather than
autoclaving fresh DNA produced the same effect and the
activating pre-existing amylase?
active ingredient was shown to be chemically similar to the
15.19 Howcould you prove that amylase base adenine, a constituent of DNA (section 5.6). It was
synthesis takes place in the aleurone layer? called kinetin. The term ‘kinin’ was used by Skoog for
15.20 How might the effect of gibberellin substances concerned with the control of cell division.
on barley seeds be used as a bioassay for gibberellin Later the term cytokinin was adopted, partly from
activity? -. cytokinesis, meaning cell division, and partly because kinin
has an entirely different meaning in zoology (it is a blood
polypeptide). The first naturally occurring cytokinin to be
Other effects of gibberellins chemically identified was from young maize (Zea mais)
Further effects of gibberellins on flowering, fruit growth grains in 1963 and hence called zeatin. Note again the
and dormancy and their involvement in photoperiodism chemical similarity to adenine (fig 15.23).
and vernalisation are discussed later under the appropriate
headings. Their effects are summarised in table 15.4.
Mode of action of gibberellins
cat
The mechanism of action of gibbereilins remains unclear.
In cereal grains GA, has been shown to stimulate synthesis
of new protein, particularly a-amylase, and is effective in CH,OH
such low concentrations (as little as 10° ug cm”) that it
must be operating at a profound level in cell metabolism,
such as the ‘switching’ on or off of genes which takes place HN CH,
during cell differentiation (section 22.7). No conclusive
evidence that this is so has yet been obtained, and higher kinetin (6-furfuryl adenine) -
a synthetic cytokinin
concentrations are required for its other effects. In cell N
elongation it is dependent on the presence of auxins. NH, Le
Commercial applications of gibberellins N N N N
Gibberellins are produced commercially from fungal

H
cultures. They promote fruit setting and are used for zeatin — a natural cytokinin
growing seedless grapes (parthenocarpy). GA; is used in N N
the brewing industry to stimulate o.-amylase production in H
barley, and hence promote ‘malting’. A number of adenine — related to cytokinins
synthetic growth retardants act as ‘anti-gibberellins’, that is
they inhibit the action of gibberellins. Application of these Fig 15.23 Structures of kinetin, zeatin and adenine
553
Table 15.4 Roles of plant growth substances in plant growth and development.
SSS So ———
eS
Process affected Auxins Gibberellins
Stem growth Promote cell enlargement in region behind apex. Promote cell enlargement in presence of auxin. Also
Promote cell division in cambium. promote cell division in apical meristem and
cambium. Promote ‘bolting’ of some rosette plants
Root growth Promote at very low concentrations. Inhibitory at Usually inactive
higher concentrations, e.g. geotropism?
Root initiation Promote growth of roots from cuttings and calluses Inhibitory
Bud (shoot) initiation Promote in some calluses but sometimes antagonistic Promote in chrysanthemum callus. Sometimes
to cytokinins and inhibitory. Sometimes promote in promote in intact plant if apical dominance broken
intact plant if apical dominance broken (see below).
Leaf growth* Inactive Promote
Fruit growth Promote. Can sometimes induce parthenocarpy. Promote. Can sometimes induce parthenocarpy.
Apical dominance Promote, i.e. inhibit lateral bud growth Enhance action of auxins
Bud dormancy* Inactive Break
Seed dormancy* Inactive Break, e.g. cereals, ash
Flowering* Usually inactive (promote in pineapple) Sometimes substitute for red light. Therefore
promote in long-day plants, inhibit in short-day
plants
Leaf senescence Delay in a few species Delay in a few species
Fruit ripening
Abscission Inhibit. Sometimes promote once abscission starts or Inactive
if applied to plant side of abscission layer
Stomatal mechanism Inactive Inactive
Process Affected Cytokinins Abscisic acid Ethene
Stem growth Promote cell division in apical Inhibitory, notably during Inhibitory, notably during
meristem and cambium physiological stress, e.g. drought, physiological stress
Sometimes inhibit cell expansion _ waterlogging
Root growth Inactive or inhibit primary root Inhibitory, e.g. geotropism? Inhibitory, e.g. geotropism?
growth
Root initiation Inactive or promote lateral root —
growth
Bud (shoot) initiation Promote, e.g. in protonemata of —
mosses
Leaf growth* Promote —
Fruit growth Promote. Can rarely induce —
parthenocarpy
Apical dominance Antagonistic to auxins, i.e. promote —
lateral bud growth
Bud dormancy* Break Promotes, e.g. sycamore, birch Breaks
Seed dormancy* Break Promotes
Flowering* Usually inactive Sometimes promote in short-day Promotes in pineapple
plants and inhibit in long-day
plants (antagonistic to gibberellins)
Leaf senescence Delay Sometimes promotes —
Fruit ripening Promotes
Abscission Inactive Promotes
Stomatal mechanism Promotes stomatal opening Promotes closing of stomata under Inactive?
conditions of water stress (wilting)
Sil Nt atin: Titi Tia aa.
*Light and temperature are also involved — see photoperiodism and vernalisation.
NB The information presented in this table is generalised. The growth substances do0 no not necessarily always
have the effects attributed to them and
variation in response between different plants is common. It is best to pay closest attention to the positive
effects. Those stressed in bold type are the most
important for the A-level student.
554
Synthesis and distribution of cytokinins senescing leaves. A natural programme of senescence may
Cytokinins are most abundant where rapid cell division is therefore involve movement of cytokinins from older
occurring, particularly in fruits and seeds where they are leaves to younger leaves via the phloem.
associated with embryo growth. Evidence suggests that in Cytokinins are also implicated in many stages of plant
mature plants they are frequently made in the roots and growth and development (table 15.4 and section 15.3).
move to the shoots in the transpiration stream (in xylem).
Cytokinins may be re-exported from leaves via the phloem. Mode of action of cytokinins
The similarity of cytokinins to the base adenine, a
Effects of cytokinins
component of the nucleic acids RNA and DNA, suggests
Cytokinins, by definition, promote cell division. They do that they may have a fundamental role in nucleic acid
so, however, only in the presence of auxins. Gibberellins metabolism. Some unusual bases, derived from transfer
may also play a role, as in the cambium. Their interaction RNA molecules, have been shown to have cytokinin
with other growth substances is discussed later in section activity, raising the possibility that cytokinins are involved
1322. in transfer RNA synthesis. Whether this is true or not, it
One of the intriguing properties of cytokinins is their does not necessarily account for their role as growth
ability to delay the normal process of ageing in leaves. If a substances, and further evidence is still being sought.
leaf is detached from a plant it will normally senesce very
rapidly, as indicated by its yellowing and loss of protein, Commercial applications of cytokinins
RNA and DNA. However, addition of a spot of kinetin will Cytokinins prolong the life of fresh leaf crops such as
result in a green island of active tissue in the midst of cabbage and lettuce (delay of senescence) as well as
yellowing tissue. Nutrients are then observed to move to keeping flowers fresh. They can also be used to break
this green island from surrounding cells (fig 15.24). dormancy of some seeds.
15.21 Study fig 15.24 and then answer 15.2.8 Abscisic acid
the following.
(a) What difference is there in the fate of applied Discovery of abscisic acid
amino acid between an old leaf and a young leaf? Plant physiologists have more recently obtained evidence
(b) Why should there be this difference? that growth inhibitors, as well as growth promoters like
(c) What is the effect of kinetin on distribution of auxins, gibberellins and cytokinins, are important in the
radioactive amino acid in old leaves? normal regulation of growth. It had long been suspected
that dormancy was caused by inhibitors when a group at the
Even when kinetin is applied to dying leaves on an intact University of Aberystwyth, led by Wareing, set about
plant a similar, though less dramatic, effect occurs. It has trying to find them in the late 1950s. In 1963, an extract
been shown that levels of natural cytokinins decrease in from birch leaves was shown to induce dormancy of birch
Kinetin Kinetin
A hat
Old
Young
i ; ineti upon translocation

ffect of kinetin Rad
j acidid iin tobacco leaves. . Radioactiv
[ of an amino e amino acid was supplied as
photograph ic film. In the resulting autoradiographs, the
oe tendafter a periodaftranslocation the leaves were exposed to
areas containing the amino acid appear black
535
buds. The leaves had been treated with short days to mimic 15.2.9 Ethene (Ethylene)
approaching winter. Pure crystals of an active substance Discovery of ethene as a growth sub-
were isolated from sycamore leaves in 1964. The substance stance
was called dormin. It turned out to be identical to a
compound isolated by another group in 1963 from young It was known in the early 1930s that ethene gas speeded up
cotton fruit. This accelerated abscission and was called ripening of citrus fruits and affected plant growth in various
abscisin II (abscisin I is a similarly acting, but chemically ways. Later it was shown that certain ripe fruits, such as
unrelated and less active compound). In 1967 it was agreed bananas, gave off a gas with similar effects. In 1934
to call the substance abscisic acid (ABA). It has been found yellowing apples were shown to emit ethene and it was
in all groups of plants from mosses upwards and a substance subsequently shown to emanate from a wide variety of
that plays a similar role, lunularic acid, has been found in ripening fruits and other plant organs, particularly from
algae and liverworts. wounded regions. Trace amounts are normal for any
organ.
Structure of abscisic acid
Structure of ethene
Like the gibberellins, ABA is a terpenoid and has a
See fig 15:26:
complex structure (fig 15.25). It is the only growth
substance in its class. H H
CH,
C=C C,H,
wae se
Fig 15.26 Structure of ethene (ethylene)

OH
COOH
O CH,
Synthesis and distribution of ethene
Fig 15.25 Structure of abscisic acid As mentioned above, ethene is made by most or all plant
organs. Despite being a gas it does not generally move
freely through the system of air spaces in the plant because
Synthesis and distribution of ABA
it tends to escape more easily from the plant surface.
ABA is made in leaves, stems, fruits and seeds. The fact However, movement of the water-soluble precursor of
that isolated chloroplasts can synthesise it again suggests a ethene from waterlogged roots to shoots in the xylem has
link with the carotenoid pigments, which are also made in been demonstrated.
chloroplasts. Like the other growth substances, ABA
moves in the vascular system, mainly in the phloem. It also Effects of ethene
moves from the root cap by diffusion (see geotropism). Ethene is known chiefly for its effects on fruit ripening and
the accompanying rise in rate of respiration (the climac-
Effects of ABA
teric) which occurs in some plants (section 15.3.5). Like
Table 15.4 summarises the effects of ABA on growth and ABA it acts as a growth inhibitor in some circumstances
development. It is a major inhibitor of growth in plants and and can promote abscission of fruits and leaves. Its effects
is antagonistic to all three classes of growth promoters. Its are summarised in table 15.4.
classical effects are on bud dormancy (including apical
dominance), seed dormancy and abscission (see section Commercial applications of ethene
15.3.4) but it also has roles in wilting, flowering, leaf Ethene induces flowering in pineapple and stimulates
senescence and possibly geotropism. It is associated with ripening of tomatoes and citrus fruits. Fruits can often be
Stress, particularly drought. In wilting tomato leaves, for prevented from ripening by storage in an atmosphere
example, the ABA concentration is 50 times higher than lacking oxygen; ripening can subsequently be regulated by
normal and ABA is thought to bring about closure of application of ethene with oxygen. The commercial
stomata. High concentrations stop the plant growing compound ‘ethephon’ breaks down to release ethene in
altogether. plants and is applied to rubber plants to stimulate the flow
of latex.
Mode of action of ABA
This is unknown.
Commercial applications of ABA

15.3 Synergism and antagonism
ABA can be sprayed on tree crops to regulate fruit drop at Having studied individual growth substances it
the end of the season. This removes the need for picking has become clear that they generally work by interacting
over a long time-span. with one another, rather than each controlling its own
556
PML eUtter beri esa8) intermediate kinetin: auxin ratio low kinétin: auxin ratio
shoot growth callus growth root growth
Fig 15.27 Cultures of tobacco callus. The culture medium in each case contains IAA (2 mg dm-3). The culture 0.2 mg dm-*
kinetin (centre) continues growth as a callus; with a lower kinetin addition (0.02 mg dm-*) it initiates roots and with a higher kinetin
addition (0.5 mg dm-%) it initiates shoots
specific aspect of growth. Two kinds of control emerge. In formation whereas a high kinetin to auxin ratio promoted
the first, two or more substances supplement each other’s lateral buds which grew into leafy shoots. Undifferentiated
activities. It is often found that their combined effect is growth would occur if the growth substances were in
much greater than the sum of their separate effects. This is balance (fig 15.27).
called synergism and the substances are said to be
synergistic. The second kind of control occurs when two 15.3.3 Apical dominance
substances have opposite effects on the same process, one Apical dominance is the phenomenon where-
promoting and the other inhibiting. This is called antagon- by the presence of a growing apical bud inhibits growth of
ism and the substances are said to be antagonistic. Here the lateral buds. It also includes the suppression of lateral root
balance between the substances determines response. growth by growth of the main root. Removal of a shoot
Some of the better understood phases of plant growth apex results in lateral bud growth, that is branching. This is
and development can now be studied and the importance of made use of in pruning when bushy rather than tall plants
synergism and antagonism demonstrated. are required.
15.3.1 Shoot growth 15. 23 (a)What plant.grW substance isis

The effect of gibberellins on elongation of made in the shoot apex? (b) Design an experiment to
show whether itis responsible for apical dominance.
stems, petioles, leaves and hypocotyls is dependent on the
presence of auxins.
It is interesting to note that auxin levels at the lateral buds
are often not high enough to cause inhibition of growth.
15.22 How, could you demonstrate this
Auxins exert their influence in an unknown way, possibly
experimentally?
by somehow ‘attracting’ nutrients to the apex. In cocklebur
it appears that the fall in auxin level in the stem after
15.3.2 Cell division and differentiation decapitation permits the lateral buds to inactivate the high
Cytokinins promote cell division only in the levels of ABA that they contain. Gibberellins often
presence of auxins. Gibberellins sometimes also play a enhance the response to IAA. Kinetin application to
role, as in the cambium when auxins and gibberellins come lateral buds, however, often breaks their dormancy, at
from nearby buds and leaves. The interaction of cytokinins least temporarily. Kinetin plus IAA causes complete
with other growth substances was demonstrated in the breaking of dormancy. Cytokinins are usually made in the
classic experiments of Skoog in the 1950s, already roots and move in the xylem to the shoots. Perhaps,
therefore, they are normally transported to wherever auxin
mentioned. His team showed the effect of various
is being made, and they combine to promote bud growth.
concentrations of kinetin and auxin on growth of tobacco
Apical dominance is a classic example of one part of a
pith callus. A high auxin to cytokinin ratio promoted root
a
(a)
gradient of auxins and gibberellins
from apical bud which ‘attracts’
cytokinins
>ytokinins ¢ and food
00
(a)
i epidermis
periderm
stem
axillary
bud
leaf
dormant lateral bud with petiole

node low cytokinin, auxin and
gibberellin levels; may xylem vascular
food, e.g. sucrose, also possess a growth
exported from leaf
phloem f strand
inhibitor, e.g. abscisic
acid
abscission layer oe
cytokinins and
mineral salts from protective layer abscission
zone
roots to apex
periderm (outer layer is cork)

(b) removal of apical bud
separation layer shown as it

is just beginning to separate
lateral bud becomes active

and produces own cytokinin,
food from leaf auxin and gibberellin;
growth inhibitor, if
present, inactivated
cytokinins and
mineral salts
corky outer layer of

Fig 15.28 Possible involvement of plant growth substances periderm forms scar
in apical dominance, (a) in presence of apical bud, (b) after
removal of apical bud
raised dot indicates
former vascular strand
plant controlling another via the influence of a growth

substance. This is called correlation (fig 15.28).
15.3.4 Abscission
Abscission is the organised shedding of part of
the plant, usually a leaf, unfertilised flower or fruit. At the
base of the organ, in a region called the abscission zone, a
layer of living cells separates by breakdown of their middle
lamellae, and sometimes breakdown of their cell walls. Fig 15.29 Abscission zone ofa leaf, (a) during abscission,
This forms the abscission layer (fig 15.29). Final shedding (b) after abscission
of the organ occurs when the vascular strands are broken
mechanically, such as by the action of wind. A protective
layer is formed beneath the abscission layer to prevent
infection or desiccation of the scar, and the vascular strand evergreen species, abscission is spread over the whole year
is sealed. In woody species the protective layer is corky, and the leaves are usually modified to prevent water loss.
being part of the tissue produced by the cork cambium, It has been shown that as a leaf approaches abscission, its
namely the periderm (section 21.6). output of auxin declines. Fig 15.30 summarises the effect of
Abscission of leaves from deciduous trees and shrubs is auxin on abscission. It is worth noting that once abscission
usually associated with the onset of winter, but in the has been triggered, auxins seem to accelerate the process.
tropics it often occurs with the onset of a dry season. In Abscisic acid (ABA) acts antagonistically to auxin by
both cases it affords protection against possible water promoting abscission in some fruits. Unripe seeds produce
shortage, leaves being the main organs through which auxins, but during ripening auxin production declines and
water is lost by transpiration. In winter, for example, soil ABA production may rise. In developing cotton fruits, for
water may be unavailable through being frozen. In example, two peaks of ABA occur. The first corresponds to
558
abscission of easily be induced! Gibberellins have the same effect in
petiole
some plants, such as the tomato, including some that are
not affected by auxins, for example cherry, apricot and
peach. Developing seeds are not only a rich source of
auxins and gibberellins, but also of cytokinins (section
me
15.2.7). These growth substances are mainly associated
with development of the embryo and accumulation of food
remove blade IAA in lanolin
reserves in the seed, and sometimes in the pericarp (fruit
se paste wall) from other parts of the plant.
Fruit ripening is really a process of senescence and is
often accompanied by a burst of respiratory activity called
the climacteric. This is associated with ethene production.
Vee |) The subsequent roles of ethene and ABA in fruit abscission
were discussed in section 15.3.4.
Fig 15.30 Effect of auxin (IAA) on abscission of a leaf

petiole. Removal of leaf blade leads to abscission of the 15.4 Phytochrome and effects of light on
petiole. IAA substitutes for the presence of the leaf blade plant development
The importance of environmental stimuli to

the growth and orientation of plant organs has already been
the ‘June drop’ when self-thinning of the plants occurs:
discussed with plant movements. The stimulus which has
only the aborted immature fruitlets have high ABA levels
the widest influence on plant growth is light. Not only does
at this stage. The second corresponds with seed ripening.
it provide the energy for photosynthesis and influence plant
There is some doubt as to whether ABA also affects leaf
movements, but it directly affects development. The effect
abscission. Applications of high concentrations are effec-
of light upon development is called photomorphogenesis.
tive, but this could be a result of stimulating ethene
production. Ethene is produced by senescing leaves and 15.4.1 Etiolation
ripening fruits and always stimulates abscission when
applied to mature organs. Some deciduous shrubs and trees Perhaps the best way to demonstrate the
produce ABA in their leaves just before winter (section importance of light is to grow a plant in the dark! Such a
15.5.1) but this may be purely to induce bud dormancy. plant lacks chlorophyll (is chlorotic) and therefore appears
Abscission is of immense horticultural significance white or pale yellow rather than green. The shoot
because of its involvement in fruit drop. Commercial internodes become elongated and thin and it is described as
applications of auxins and ABA reflect this and have been etiolated. In dicotyledonous plants, the epicotyl or hypo-
discussed earlier in this chapter. cotyl (section 21.6.2) elongates in hypogeal or epigeal
If flowers are not fertilised they are generally abscised germination respectively and the plumule tip is hooked.
(see ‘fruit set’ below). Dicotyledonous leaves remain small and unexpanded. In
monocotyledonous plants, the mesocotyl elongates during
15.3.5 Pollen tube growth, fruit set and fruit germination and the leaves may remain rolled up. In all
development leaves, chloroplasts fail to develop normal membrane
systems and are called etioplasts. Plants make less
Germinating pollen grains are a rich source of
supporting tissue and are fragile and collapse easily.
auxins as well as commonly stimulating the tissues of the Eventually they use up their food reserves and die unless
style and ovary to produce more auxin. This auxin is light is reached for photosynthesis. Yet as soon as the plant
necessary for ‘fruit set’, that is retention of the ovary, which is exposed to light, normal growth ensues. The significance
becomes the fruit after fertilisation. Without it abscission of etiolation is that it allows maximum growth in length
of the flower normally occurs. After fertilisation, the ovary with minimum use of carbon reserves which, in the absence
and the ripe seeds continue to produce auxins which of light, the plant cannot obtain by photosynthesis.
stimulate fruit development.
A few natural examples are known where fruit develop- 15.24 How does the morphology of an
ment proceeds without fertilisation, and therefore without etiolated plant suit it for growing through soil?
seed development, for example banana, pineapple and
some seedless varieties of oranges and grapes. Such 15.4.2 Discovery of phytochrome
development is called parthenocarpy. Unusually high
The first stage in any process affected by light
auxin levels occur in these ovaries. Parthenocarpy can
must be the absorption of light by a pigment, the so-called
sometimes be artificially induced by adding auxins, as in
photoreceptor. The characteristic set of wavelengths of
tomato, squash and peppers. Seedless pea pods can just as
559
660nm 730 nm Normal sunlight contains more red than far-red light, so
the Pp form predominates during the day. This is the
inhibition of lettuce physiologically active form, but reverts slowly to the more
seed germination,
leaf inhibition, stable, but inactive, Pg form at night. Phytochrome was
promotion of lettuce seed hooking response shown to consist of a pigment portion attached to a protein.
germination,
leaf enlargement,
in etiolated bean
seedling, etc.
It is present in minute amounts throughout plants (hence it
removal of hypocotyl hook is not visible, despite its colour (but is particularly
concentrated in the growing tips. Its absorption spectrum is
from etiolated bean seedling,
etc.
shown in fig 15.32.
660 nm
Effectiveness
Pp or Pyow
Absorbance
400 500 600 700 800

400 500 600 700 : 800
Wavelength/nm Wavelength/nm
Fig 15.31 Typical action spectra of a phytochrome-controlled Fig 15.32 Absorption spectra of the two forms of
response phytochrome
light it absorbs form its absorption spectrum (section

15.25 What is the difference between an
9.3.2); remaining wavelengths are reflected and give the
absorption spectrum and an action spectrum?
substance a characteristic colour (chlorophyll, for ex-
ample, absorbs red and blue light and reflects green light).
The seeds of many plants germinate only if exposed to
light. In 1937 it was shown that, for lettuce seeds, red light
promoted germination but far-red light (longer wave-
A number of developmental processes are mediated by low
length) inhibited germination. Borthwick and Hendrick,
intensities of red light and reversed by far-red light or
working at the US Department of Agriculture in the 1950s,
darkness, showing the involvement of phytochrome (fig
plotted an action spectrum for the germination response (a 15.31 and table 15.5). The most significant of these is
spectrum of wavelengths showing their relative effective- flowering, which is discussed later. The involvement of
ness at stimulating the process). Fig 15.31 shows that the phytochrome ina process is shown by matching the action
wavelength most effective for germination was about
spectrum of the response with the absorption spectrum of
660 nm (red light) and for inhibition of germination about
phytochrome.
730 nm (far red light).
They also showed that only brief exposures of light were 15.4.3 Photoperiodism and flowering
necessary and that the effects of red light were reversed by
far-red light and vice versa. Thus the last treatment in an One of the important ways in which light
alternating sequence of red/far-red exposures would exerts its influence on living organisms is through variations
always be the effective one. The US team eventually in daylength (photoperiod). The further from the equator,
isolated the pigment responsible in 1960 and called it where days are almost a constant 12 hours, the greater the
phytochrome. Phytochrome, as they predicted, is a seasonal variation in daylength. Thus daylength is an
blue-green pigment existing in two interconvertible forms. important environmental signal in temperate latitudes
One form, Pz Or P73) absorbs far-red light and the other, Pz where it varies between about 9 and 15 hours during the
Or Pee, absorbs red light. Absorption of light by one form year. The effects of photoperiod on animals are discussed
converts it rapidly and reversibly to the other form (within in section 16.8.5. In plants it is a matter of common
seconds or minutes depending on light intensity): observation that phenomena such as flowering, fruit and
seed production, bud and seed dormancy, leaf fall and
red light (660 nm) germination are closely attuned to seasonal influences like
ee ee aN daylength and temperature, and that survival of the plant
i Pix ———® biochemical process
ES rae year e ae ae
\ _ far-red light (730 nm) i depends on this.
\
NX a
/ The process involving the most profound change is
= _
a
flowering, when shoot meristems switch from producing
Slowly in darknes® leaves and lateral buds to producing flowers. The impor-
560
Table 15.5 Some phytochrome-controlled responses in Table 15.6 Classification of plants according to
plants. photoperiodic requirements for flowering.
General process affected Red light promotes Short-day plants (SDPs) Long-day plants (LDPs)
Germination* Germination of some seeds, e.g. some e.g. cocklebur (Xanthium e.g. henbane (Hyoscyamus
lettuce varieties pennsylvanicum), chrysanthe- niger), snapdragon, cabbage,
Germination of fern spores mum, soybean, tobacco, spring wheat, spring barley
Photomorphogenesis Leaf expansion in dicotyledons. Leaf strawberry
(light-controlled unrolling in grasses (monocotyle- Flowering induced by dark Flowering induced by dark
development of form dons). Chloroplast development periods longer than acritical periods shorter than a critical
and structure) (etioplasts converted to chloroplasts: length, e.g. cocklebur 8.5 h; length, e.g. henbane 13 h
see etiolation). Greening (proto- tobacco 10-11 h
chlorophyll converted to chlorophyll). (Under natural conditions (Under natural conditions
Inhibition of internode growth equivalent to days shorter than equivalent to days longer than
(including epicotyl, hypocotyl, a critical length, e.g. cocklebur a critical length, e.g. henbane
mesocotyl), i.e. preventing of 15.5 h; tobacco 13-14 h) 11 h)
etiolation. Unhooking of plumule in
dicotyledons Day-neutral plants
Photoperiodism Stimulates flowering in long-day e.g. cucumber, tomato, garden pea, maize, cotton
plants. Inhibits flowering in short- Flowering independent of photoperiod
day plants. See flowering
NB Tobacco (SDP) and henbane (LDP) both flower in 12-13 h
*Experiments designed to investigate the effects of light on seed daylength.
germination are described by J. W. Hannay inJ. Biol. Ed. (1967) 1, 65-73.
The variety of lettuce suggested, ‘Grand Rapids’, is no longer available
but some modern varieties could be screened for suitability. Such varieties
currently available are ‘Dandie’, ‘Kloek’ and ‘Kweik’. ‘Dandie’ is
probably the most reliable but is fairly expensive because it is a
winter-forcing variety. It is available from Suttons in small packets or from
E. W. King & Co., Coggeshall, Essex in 10 g packets (enough for about
250 dishes of 50 seeds each). Phacelia seeds make an interesting contrast
to lettuce.
NB In this experiment a green leaf can be used as afar-red filter.
tance of photoperiod in flowering was discovered as early do not cancel the effect of long days. Table 15.6
as 1910 but was first clearly described by Garner and Allard summarises the three main categories of plant.
in 1920. They showed that tobacco plants would flower only
after exposure to a series of short days. This occurred
naturally in autumn, but could be induced by artificially 15.4.4 Quality and quantity of light
short days of seven hours in a greenhouse in summer. As
they examined other plants it became obvious that some The next step is to find the quality (colour) and
required long days for flowering (long-day plants) and quantity of light required. Remembering that the cockle-
some would flower whatever the photoperiod once mature bur (SDP) will not flower if its long night is interrupted,
(day-neutral plants). experiments revealed that red light was effective in
Additional complications have since been found. For preventing flowering, but that far-red light reversed the
example, some plants are day-neutral at one temperature, effect of red light. Therefore phytochrome is the photo-
but not at another; some require one daylength followed by receptor. These experiments were, in fact, part of the
programme which led to the discovery of phytochrome. As
another; in some, the appropriate daylength only acceler-
would be expected, a LDP held in short days is stimulated
ates flowering and is not an absolute requirement.
to flower by a short exposure to red light during the long
An important advance in our understanding came when
it was shown that it is really the length of the dark period
night. Again this is reversed by far-red light. The last light
which is critical. Thus short-day plants are really long-night
treatment always determines response.
In some, though not all, cases low light intensities for a
plants. If they are grown in short days, but the long night is
few minutes are effective, again typical of a phytochrome-
interrupted by a short light period, flowering is prevented.
controlled response. The higher the intensity used, the
Long-day plants will flower in short days ifthe long night
shorter the exposure time required.
period is interrupted. Short dark interruptions, however,
561
15.4.6 Mode of action of phytochrome
15.26 Xanthium (cocklebur) is a short-
day plant. Flowering is therefore normally induced by _ How, then, does phytochrome exert its con-
long nights. In experiments the effects of red and trol? At the end of a light period it exists in the active Prep
red/far-red light interruptions of three consecutive form. At the end of a short night its slow transition back to
12h (‘long’) nights on subsequent flowering were the inactive P, form, which takes place in darkness, may
investigated. Flowering intensity was measured by not be complete. It can be postulated, therefore, that in
an index of ‘floral stage’ from 1 to 8, and the results LDPs Py, promotes flowering and in SDPs it inhibits
are shown in table 15.7. flowering. Only long nights remove sufficient Pr from the
Draw graphs of these results, plotting floral stage on latter to allow flowering to occur. Unfortunately, short
the vertical axis and duration of light on the horizontal exposures to far-red light, which would have the same
axis. (The two graphs can be drawn between the effect as a long night, cannot completely substitute for long
same axes.) Comment on the results. : : nights, so the full explanation is more complex. Some time
factor is also important.
We do know that gibberellins can mimic the effect of red
light in some cases. Gibberellic acid (GA3) promotes
Table 15.7 Effect of red/far-red light interruptions of long flowering in some LDPs, mainly rosette plants like
nights on flowering of cocklebur (after Downs, R. J. henbane which bolt before flowering. (Bolting is a rapid
(1956) Plant Physiol. 31, 279-84).
increase in stem length.) GA also inhibits flowering in
some SDPs. Antigibberellins (growth retardants) nullify
Red light Two minutes red light followed these effects.
by FR light
So, does Pep stimulate gibberellin production, and is this
Floral Duration of red Floral Duration of FR the flowering hormone? There are too many exceptions for
stage _light/s stage _ light/s this to be the case. Abscisic acid inhibits flowering in some
LDPs, such as Lolium, but induces it in some SDPs, such as
6.0 0 0.0 0
5.0 5 4.0 12 strawberry. In short, our understanding of the flowering
4.0 10 4.5 15 process is still incomplete.
2.6 20 5 25
0.0 30 6.0 50
15.5 Vernalisation and flowering
Some plants, especially biennials and peren-
nials, are stimulated to flower by exposure to low
temperatures. This is called vernalisation. Here the
stimulus is perceived by the mature stem apex, or by the
embryo of the seed, but not by the leaves as in
photoperiodism. As with photoperiod, vernalisation may
15.4.5 Perception and transmission of the be an absolute requirement (such as in henbane) or may
stimulus
simply hasten flowering (as in winter cereals).
It was shown in the mid-1930s that the light Long-day plants (for example cabbage), short-day plants
stimulus is perceived by the leaves and not the apex where (such as chrysanthemum) and day-neutral plants (such as
the flowers are produced. ragwort) can all require vernalisation. The length of
chilling required varies from four days to three months,
15.27 How could you demonstrate this temperatures around 4 °C generally being most effective.
with a short-day plant? Like the photoperiodic stimulus, the vernalisation stimulus
can be transmitted between plants by grafting. In this case
In addition to this, a cocklebur plant with just one induced the hypothetical hormone involved was called vernalin. It
leaf will flower even if the rest of the plant is under has subsequently been discovered that during vernalisation
non-inductive conditions. This implies that some agent, gibberellin levels increase, and application of gibberellins
that is a hormone, must pass from the leaf to the apex to to unvernalised plants can substitute for vernalisation (fig
bring about flowering. This concept is supported by the 15.33). It is now believed that ‘vernalin’ is a gibberellin.
observation that the flowering stimulus can be passed from It is clear now that photoperiodism and vernalisation serve
an induced plant to a non-induced plant by grafting, and to synchronise the reproductive behaviour of plants with
that the stimulus is apparently the same for SDP, LDP and their environments, ensuring reproduction at favourable
day-neutral plants because grafting between these is times of the year. They also help to ensure that members of
successful. The hypothetical flowering hormone has been the same species flower at the same time and thus
called ‘florigen’ but has never been isolated. Indeed, its encourage cross-pollination and cross-fertilisation, with
existence is doubted by some plant physiologists. the attendant advantages of genetic variability.
562
15.571 Photoperiodism and dormancy
The formation of winter buds in temperate
trees and shrubs is usually a photoperiodic response to
shortening days in the autumn, for example in birch, beech
and sycamore. The stimulus is perceived by the leaves and,
as mentioned before, abscisic acid (ABA) levels build up.
ABA moves to the meristems and inhibits growth. Short
days also induce leaf fall from deciduous trees (abscission).
Often buds must be chilled before dormancy can be broken
(‘bud-break’). Similarly some seeds require a cold stimulus
(‘stratification’) after imbibing water before they will
germinate, thus preventing them from germinating prema-
turely once ripe. This is discussed further in section 21.10.
Gibberellins can substitute for the cold stimulus, and
natural bud-break is accompanied bya rise in gibberellins
as well as, in many cases, a fall in ABA. Breaking of bud
dormancy in birch and poplar has been shown to coincide
with a rise in cytokinins.
Apart from buds and seeds, storage organs are involved
in dormancy and again photoperiod is important. For
example, short days induce tuber formation in potatoes,
whereas long days induce onion bulb formation.
15.28 Some buds remain dormant Ae

throughout the summer. What causes dormancy —
here?
Fig 15.33 Carrot plants (var. Early french forcing). J/eft:

control; centre: maintained at 17°C but supplied 10 mg of
gibberellin daily for 4 weeks; right: plant given vernalizing cold
treatment (6 weeks). All photographed 8 weeks after completion
of cold treatment
563
Chapter Sixteen
Coordination and control in animals
Irritability or sensitivity is a characteristic feature of all 16.1.1 The nature of the nerve impulse
living organisms and involves their ability to respond to a
stimulus. In all organisms some degree of internal The transmission of information along
coordination and control is necessary in order to ensure neurones as electrical impulses and its effects on muscle
that the events of the stimulus and response bear some contraction and glandular secretion have been known for
mutual relationship associated with the maintenance of the over 200 years. Details of the mechanisms, however, were
steady-state (chapter 18) and survival of the organism. established only in the last 40 years following the discovery
Animals, unlike plants, have two different but related of certain axons in squid which have a diameter of
systems of coordination: the nervous system and the approximately 1 mm (1000 um). These giant axons (sup-
endocrine system. The former is fast acting, its effects are plying the muscles of the mantle involved in escape
localised and it involves electrical and chemical transmis- responses) had large enough diameters to permit the
sion, whereas the latter is slower acting, its effects are earliest electrophysiological investigations to be carried
diffuse and it relies on chemical transmission through the out on them.
circulatory system. It is thought that the two systems have The apparatus currently used to investigate electrical
developed in parallel in the majority of multicellular activity in neurones is shown in fig 16.1. The microelec-
animals. trode, composed of a small glass tube drawn out to a fine
point of 0.5 um diameter, is filled with a solution capable of
conducting an electric current, such as 3M potassium
chloride. This is inserted into an axon and a second
16.1 The nervous system electrode, in the form of a small metal plate, is passed into
the saline solution bathing the neurone being investigated.
The nervous system is composed of highly Both electrodes are connected by leads to a preamplifier to
differentiated cells whose function is to detect sensory complete the circuit. The preamplifier increases the signal
information, code this in the form of electrical impulses and strength in the circuit approximately 1 000 times and
transmit them, often over considerable distances, to other provides the input to a dual-beam cathode ray oscilloscope.
differentiated cells capable of producing a response. All movements of the microelectrode are controlled by a
All sensory information (stimuli) is detected in multi- micromanipulator, a device with adjusting knobs similar to
cellular animals by modified nerve cells called sensory those of the microscope, which enables delicate control
receptors and the structure and function of these is over the position of the tip of the microelectrode.
described in section 16.5. This sensory information is When the tip of the microelectrode penetrates the axon
passed on to effector cells which produce a response which plasma membrane, the beams of the oscilloscope separate.
is associated in some way with the stimulus. The structure The distance between the beams indicates the potential
and function of effectors is described briefly in section 16.6 difference between the two electrodes of the circuit and can
and chapter 17. be measured. This value is called the resting potential (RP)
Interposed between the receptors and effectors are the of the axon and is approximately —65 mV in all species
conductile cells of the nervous system, the neurones. These investigated. The negative sign for the resting potential
are the basic structural and functional units of the nervous indicates that the membrane of the axon is polarised, that is
system and ramify throughout the organism forming an the inside of the axon is negative with respect to the outside
elaborate communication network. Types of neurone, and of the axon. In sensory cells, neurones and muscle cells this
their structures, are shown and described in section 8.6. value changes with the activity of the cells and hence they
The structural complexity and organisation of the neurones are known as excitable cells. All other living cells show a
within the network is related to the phylogenetic position of similar potential difference across the membrane, known
the organism and ranges from the primitive nerve net of the as the membrane potential, but in these cells this is constant
coelenterates (section 16.3) to the complex central nervous and so they are known as non-excitable cells.
system of mammals (section 16.2). Whereas the former is
Resting potential
purely a communication system, the latter provides, in
addition, facilities for the storage, retrieval and integration The resting potential of most mammalian neurones is
of information. constant as long as the cell remains inactive due to lack of
565
axon
Fig 16.1 The basic design used to record electrical activity in the
current which
of an isolated neurone. The stimulator produces a
and recorded
generates an action potential in the axon and is detected
e ray
using an intracellular microelectrode and dual-beam cathod
oscilloscope stimulator
preamplifier
cathode ray oscilloscope
3M KCI solution
external ‘earth’ electrode
microelectrode
bathing saline
stimulating solution
electrodes
axon membrane
axon
stimulation. Curtis and Cole in the USA, and Hodgkin and couple the removal of Na* ions from the axon with the
Huxley in England, in the late 1930s revealed the resting uptake of K* ions as shown in fig 16.2a.
potential to be a physico-chemical phenomenon set up and The active movement of these ions is opposed by the
maintained by the differential concentration of ions across passive diffusion of the ions which constantly pass down
the axon membrane and the selective permeability of the their electrochemical gradients as shown in fig 16.26 at a
membrane to the ions. Analyses of the intracellular fluid of rate determined by the permeability of the axon membrane
the axon and the extracellular sea water bathing the axon to the ion. K* ions have an ionic mobility and membrane
showed that ionic electrochemical gradients exist as shown permeability which is 20 times greater than that of Na*
in table 16.1. ions, therefore K* loss from the axon is greater than Na*
gain. This leads to a net loss of K* ions from the axon, and
Table 16.1. Ionic concentrations of extracellular and the production of a negative charge within the axon. The
intracellular fluids in squid axon. (Values given are value of the resting potential is largely determined by the
approximations in mmol kg' H,O, data from Hodgkin, K* electrochemical gradient.
1958) Changes in the permeability of the membrane of
excitable cells to K* and Na” ions lead to changes in the
Ion Extracellular Intracellular
concentration concentration
potential difference across the membrane and the forma-
tion of action potentials in, and the propagation of nerve
K* 20 400 impulses along, the axon.
Na’ 460 50
(Ol 560 100 OUTSIDE
AL 0 370 low [Kt] high [Na*]
(Organic anions)
The axoplasm inside the axon has a high concentration of

potassium (K*) ions and a low concentration of sodium 7.5 nm axon driven
membrane by ATP
(Na”) ions, in contrast to the fluid outside the axon which Se
has a low concentration of K” ions and a high concentration
of Na’ ions. (The distribution of chloride (Cl) ions is
ignored in the following descriptions since it does not play a high [K+] low [Nat]
Nat Kt
vital role in the activities under consideration.) INSIDE (b)
(a)
These electrochemical gradients are maintained by the
active transport of ions against their electro-chemical Fig 16.2 Active and passive ionic movements associated
with the production of a negative (—) potential within the
gradients by specific regions of the membrane known as
axon. (a) The Na*/K* coupled cation pump actively
cation or sodium pumps. These constantly active carrier exchanges ions which pass across the membrane by passive
mechanisms are driven by energy supplied by ATP and diffusion down their electrochemical gradients as shown in (b)
566
Action potential increases the number of positive ions inside the axon and
The experimental stimulation of an axon by an electrical reduces the membrane potential from its resting value of
impulse, as shown in fig 16.3, results in a change in the —70 mV. The change in membrane potential is called
potential across the axon membrane from a negative inside depolarisation. Sodium conductance and depolarisation
value of about —70 mV to a positive inside value of about influence each other by positive feedback, that is an
+40 mV. This polarity change is called an action potential increase in one factor reinforces an increase in the other,
or spike and appears on the cathode ray oscilloscope as and this produces the steep rising phase of the action
shown in fig 16.3. potential. Calculations have revealed that relatively few
Na* ions (about 10°°% of the internal Na* present,
& +60 depending upon axon diameter) enter the axon and
o> +40
ys
ES +20
produce the depolarisation of about 110 mV associated
with the action potential. At the peak of the action
ee
=
Sf
ae
= E =20 potential, sodium conductance declines (Na inactivation)
See
and about 0.5 ms after the initial depolarisation, potassium
6g
og
conductance increases and K” ions diffuse out of the axon.
Ls. ed ! eee = J As K* ions diffuse outwards the internal positive charge is
0 1 2, 3 4 S replaced by a negative charge. This repolarisation of the
Time/ms
membrane is shown by the falling phase of the action
Fig 16.3 A typical action potential in squid axon potential ‘spike’ and results in the membrane potential
assuming its original level.
An action potential is generated by a sudden momentary From the above account it can be seen that whilst the
increase in the permeability of the axon membrane to Na* resting potential is determined largely by K* ions, the
ions which enter the axon. This increase in sodium action potential is determined largely by Na” ions (fig
conductance (the electrical equivalent of permeability) 16.4).
(a) +50 Fig 16.4 Diagrams to show the

+40 relationships between (a) the membrane
+30 potential, (b) ionic conductance and (c) the
+20
+10 potential distribution across the axon
depolarisation repolarisation membrane during production of an action
-10 potential
-20
-30
-40
-50
Membrane
potential/mV
-60
-70
hyperpolarisation
+__—_!_—_—direction of impulse
(b)
Ionic
conductance
change
ROTC ULA eae ee eS

heer] anne ete ga = 16.1. Give two reasons why there is a
difference axon membrane sudden influx of Na* ions into the axon following an
across axon
membrane ===== increase in Na* permeability of the axon membrane.
PH FER EFF FHT EH tet ttt tet tt 16.2 If the permeability of the axon mem-
(©)
brane to Na* ions and K* ions increased simul-
ee Seen eens
5
taneously what effect would this have on the action
2 3 4
0 1 _ potential?
Time/ms
567
transmitted over an infinite distance, that is, they are
non-decremental. The reason for this is that the production
of an action potential at each point along the axon is a
self-generating event resulting from a change in the local
concentration of ions. So long as the extra- and intra-
cellular environments of the axon have the necessary
differential ionic concentrations an action potential at one
potential/mV
Membrane
point will generate another action potential in its adjacent
region.
Time/ms Refractory period. Nerve impulses only pass
Fig 16.5 Membrane potentials recorded from squid axons
along the axon in one direction from active region to resting
bathed in isotonic sea water containing different relative region. This is because the previously active region
concentrations of ions undergoes a recovery phase during which the axon
membrane cannot respond to a depolarisation by a change
16.3 Hodgkin and Katz, in 1949, investi- in sodium conductance, even if the stimulus intensity is
gated the effect of Na* ions on the production of increased. The phase is called the absolute refractory
action potentials in squid axons. Intracellular micro- period and lasts for about 1 ms. Following this is a period
electrodes recorded action‘ potentials from axons lasting for 5-10 ms called the relative refractory period
bathed in different concentrations of isotonic sea during which a high-intensity stimulus may produce a
water. The results are shown in fig 16.5. Which action depolarisation. The refractory period is also a limiting
potentials correspond with axons placed in normal factor in the speed of conduction of the nerve impulse.
sea water, one-half sea water and one-third sea
water? Explain the effect of these solutions on the 16.4 Describe the ionic changes occur-
action potentials. ring across the axon membrane during the refractory
period. ©
Features of action potentials
Speed of conduction. In non-myelinated
Initiation of an action potential. Stimulation of axons (section 16.2), typical of those found in inverte-
sensory cells leads to depolarisation of their membranes brates, the velocity of the propagated action potential
and if this reaches a certain threshold value in the sensory depends on the longitudinal resistance of the axoplasm.
neurone, the threshold stimulus intensity, it will set up an The resistance, in turn, is related to the diameter of the
action potential. For any given neurone the amplitude axon such that the smaller the diameter the greater the
(fig 16.3) of the action potential is always constant, and resistance. In the case of fine axons (<0.1 mm) the high
increasing the strength or number of stimuli have no effect resistance of the axoplasm has an effect on the spread of
on this. For this reason action potentials are described as current and reduces the length of the local circuits so that
all-or-nothing events. only the region of the membrane immediately in front of
the action potential is involved in the local circuit. These
Transmission of nerve impulses. Infor-
axons conduct impulses at about 0.5 m s |. Giant axons,
mation is transmitted through the nervous system as a
typical of many annelids, arthropods and molluscs, have a
series of nerve impulses, which travel as action potentials.
diameter of approximately 1 mm and conduct impulses at
A nerve impulse passes along an axon as a wave of
velocities up to 100 m s-’, which are ideal for conducting
depolarisation accompanied by a wave of negativity along
information vital for survival.
the surface of the axon marking the position, at any instant,
of the action potential. Action potentials are propagated,
that is self-generated, along the axon by the effects of Na* 16.5 Explain, in terms of the resistance of
ions entering the axon. This creates an area of positive the axoplasm and local circuits, why giant axons
charge and a flow of current is set up ina local circuit conduct impulses at greater velocities than fine
between this active area and the negatively charged resting axons. . :
region immediately ahead. The current flow in the local
circuit reduces the membrane potential in the resting In vertebrates, the majority of neurones, particularly
region and this depolarisation produces an increase in those of the spinal and cranial nerves, have an outer
sodium permeability and the development of an all-or- covering of myelin derived from the spirally wound
nothing action potential in this region. Repeated depolar- Schwann cell (section 8.6.3). Myelin is a fatty material with
isations of immediately adjacent regions of the membrane a high electrical resistance and acts as an electrical insulator
result in the action potential being moved or propagated in the same way as the rubber and plastic covering of
along the axon. Action potentials are propagated along the electrical wiring. The combined resistance of the axon
axon without change in amplitude and are capable of being membrane and myelin sheath is very high but where breaks
568
in the myelin sheath occur, as at the nodes of Ranvier, the (a) ———
direction of
resistance to current flow between the axoplasm and the > propagation
extra-cellular fluid is lower. It is only at these points that £
local circuits are set up and current flows across the axon ‘s
a
membrane generating the next action potential. This 2
°
or
means, in effect, that the action potential ‘jumps’ from o
fon)
node to node and passes along the myelinated axon faster

S
S
me
&
than the series of smaller local currents in a non-myelinated vo
=
axon. This type of conduction is called saltatory (saltare, to
jump) and can lead to conduction velocities of up to
120 m s"' (fig 16.6). cation pump (active
Temperature has an effect on the rate of conduction of diffusion of ions at all times)
nerve impulses and as temperature rises to about 40 °C the

rate of conduction increases.
p> direction of impulse

node of (c) local circuits <==) |
local circuit fea + + + + + ee, ea ae eee See oe eae
state of axon resting active recovering

type of resting potential action potential resting potential
potential
membrane polarised depolarised repolarising
ionic stable, differential | increased almost stable,
movement ionic permeability to differential ionic
concentrations Na‘ ions and K* ions, concentrations
maintained being restored
internal negative positive negative
potential
external positive negative positive
Fig 16.6 Diagrams showing the difference in lengths of the potential
local circuits produced (a) in a myelinated axon and (b) a non-
current flow local currents exist between none
myelinated axon. In (a) conduction is described as saltatory resting and active regions
since the action potential effectively ‘jumps’ from node to
node Fig 16.7 Summary diagrams showing the events
accompanying the production of local circuits within the axon
and the propagation of an action potential in a non-myelinated
axon, (a) action potential and direction of propagation, (b) ionic
movements across the axon membrane, (c) current flow in
16.6 Why do myelinated axons of frog
local circuits
having a diameter of 3.5 um conduct impulses at
30 m s-' whereas axons of the same diameter in cat 16.1.2 The synapse
conduct impulses at 90 m s-1?
A synapse is an area of functional, but not
Coding of nervous information. Nerve im- physical, contact between one neurone and another for the
pulses pass through the nervous system as propagated, purpose of transferring information. Synapses are usually
all-or-nothing action potentials with a fixed amplitude for a found between the fine terminal branches of the axon of
given species, for example 110 mV in squid axon. Informa- one neurone and the dendrites (axodendritic synapses) or
tion cannot therefore be passed as an amplitude code and is cell body (axosomatic synapses) of another neurone. The
passed instead as a frequency code. This code was first number of synapses is usually very large, providing a large
described by Adrian and Zotterman in 1926. They surface area for the transfer of information. For example,
demonstrated that the frequency of nerve impulses is over 1 000 synapses may be found on the dendrites and cell
directly related to the intensity of the stimulus giving rise to body of a motor neurone in the spinal cord. Some cells in
the impulses. the brain may receive up to 10 000 synapses (fig 16.8).
569
produced either in the cell body of the neurone and passes
down the axon to the synaptic knob by axoplasmic
streaming through microtubules, or directly in the synaptic
knob. In both cases synthesis of transmitter substances
requires enzymes produced by ribosomes in the cell body.
In the synaptic knob the transmitter substance is ‘pack-
aged’ into vesicles and stored pending release. The two
main transmitter substances in vertebrate nervous systems
are acetylcholine (ACh) and noradrenaline, although other
substances exist and are described at the end of this section.
Acetylcholine is an ammonium compound with the
formula as shown in fig 16.9. It was the first transmitter
substance to be isolated and was obtained by Otto Loewi,
in 1920, from the endings of parasympathetic neurones of
the vagus nerve (section 16.2) in frog heart. The detailed
structure of noradrenaline is described in section 16.6.6.
Neurones releasing acetylcholine are described as cho-
linergic neurones and those releasing noradrenaline are
described as adrenergic neurones.
Fig 16.8 Scanning electron micrograph of a motor neurone -
transmission at a neuronal synapse,
CH, | CH,
There are two types of synapses, electrical and chemical,

depending upon the nature of transfer of information
CH.—_- NCH,
CH,
__Gh, 0G
A
across the synapse. A structurally dissimilar, but func-
tionally similar, form of synapse exists between the Fig 16.9 Structural formula of acetylcholine
terminals of a motor neurone and the surface of a muscle
fibre and this is called a neuromuscular junction. Details of
Mechanisms of synaptic transmission
the structure and physiological differences between the
synapse and the neuromuscular junction are described The arrival of nerve impulses at the synaptic knob is
later in this section. thought to depolarise the presynaptic membrane and
increase the permeability of the membrane to Ca™* ions. As
Structure of the chemical synapse the Ca’* ions enter the synaptic knob they cause the
Chemical synapses are the commonest type of synapse synaptic vesicles to fuse with the presynaptic membrane
found in vertebrates and they consist of a bulbous and rupture (exocytosis), discharging their contents into
expansion of a nerve terminal called a synaptic knob or the synaptic cleft. This is known as excitation-secretion
bouton terminale lying in close proximity to the membrane coupling. The vesicles then return to the cytoplasm where
of a dendrite. The cytoplasm of the synaptic knob contains they are refilled with transmitter substance. Each vesicle
mitochondria, smooth endoplasmic reticulum, microfila- contains about 3 000 molecules of acetylcholine.
ments and numerous synaptic vesicles. Each vesicle has a The transmitter substance diffuses across the synaptic
diameter of approximately 50 nm and contains a chemical cleft, imposing a delay of about 0.5 ms, and attaches to a
neurotransmitter substance responsible for the transmis- specific receptor site on the postsynaptic membrane which
sion of the nerve impulse across the synapse. The recognises the molecular structure of the acetylcholine
membrane of the synaptic knob nearest the synapse is molecule. The arrival of the transmitter substance causes a
thickened as a result of cytoplasmic condensation and change in the configuration of the receptor site leading to
forms the presynaptic membrane. The membrane of the channels opening up in the postsynaptic membrane and the
dendrite is also thickened and termed the postsynaptic entry of ions which either depolarise or hyperpolarise (fig
membrane. These membranes are separated by a gap, the 16.4a) the membrane according to the nature of the
synaptic cleft, of 20 nm. The presynaptic membrane is transmitter substance released and the molecular proper-
modified for the attachment of synaptic vesicles and the ties of the postsynaptic receptor sites. Having produced a
release of transmitter substance into the synaptic cleft. The change in the permeability of the postsynaptic membrane
postsynaptic membrane contains large protein molecules the transmitter substance is immediately lost from the
which act as receptor sites for the transmitter substances synaptic area by reabsorption by the presynaptic mem-
and numerous channels and pores, normally closed, for the brane, or diffusion out of the cleft or by hydrolysis by
movement of ions into the postsynaptic neurone (fig enzymes. In the case of cholinergic synapses acetylcholine
16.10a). is hydrolysed to choline by the enzyme acetylcholinesterase
Synaptic vesicles contain a transmitter substance which is (AChE), situated on the postsynaptic membrane, and
570
reabsorbed into the synaptic knob to be recycled into
synaptic knob as aresult of an intense stimulus produces
acetylcholine by synthetic pathways in the vesicles (fig individual e.p.s.p.s which are so close together that they
16.10). summate and give rise to an action potential in the
At excitatory synapses ion-specific channels open up postsynaptic neurone. This is temporal (related in time)
allowing Na” ions to enter and K* ions to leave down their summation. Therefore impulses can be set up ina single
respective concentration gradients. This leads to a depolar- postsynaptic neurone asa result of either weak stimulation
isation in the postsynaptic membrane. The depolarising by several of its presynaptic neurones or repeated
response is known as an excitatory postsynaptic potential stimulation by one of its presynaptic neurones.
(e.p.s.p.) and the amplitude of this potential is usually At inhibitory synapses the release of transmitter sub-
small but longer lasting than that of an action potential. stance increases the permeability of the postsynaptic
The amplitude of €.p.s.p.s fluctuates in steps, suggesting membrane by opening up ion-specific channels to Cl” and
that transmitter substance is released in ‘packets’ rather K* ions. As the ions move down their concentration
than individual molecules. Each ‘step’ is thought to gradients they produce a hyperpolarisation of the mem-
correspond to the release of transmitter substance from brane known as an inhibitory postsynaptic potential
one synaptic vesicle. A single e.p.s.p. is normally unable to (i.p.s.p.).
produce sufficient depolarisation to reach the threshold Transmitter substances are neither inherently excitatory
required to propagate an action potential in the post- nor inhibitory. For example, acetylcholine has an excita-
synaptic neurone. The depolarising effect of the e.p.s.p.s tory effect at most neuromuscular junctions and synapses,
is additive, a phenomenon known as summation. Two or but has an inhibitory effect on neuromuscular junctions in
more €.p.s.p.s arising simultaneously at different regions cardiac muscle and visceral muscle. These opposing effects
on the same neurone may produce collectively sufficient are determined by events occurring at the postsynaptic
depolarisation to initiate an action potential in the membrane. The molecular properties of the receptor sites
postsynaptic neurone. This is spatial (related in space) determine which ions enter the postsynaptic cell, which in
summation. The rapid repeated release of transmitter turn determines the nature of the change in postsynaptic
substance from several synaptic vesicles by the same potentials as described above.
(a) | (b) Fig 16.10 Summary diagrams showing the

mechanisms involved in chemical
axon of
presynaptic transmission at a neuronal synapse,
neurone (a) to (e) time sequence
mitochondrion
ae synaptic
knob (Cari Cacia Caer
permeability of presynaptic membrane to
Ca** increases
jalan
membrane of ob 9%
postsynaptic Oatn }
neurone A
ees) Ss SS z; synaptic
presynaptic membrane abe
area of receptor sites postsynaptic
membrane
| (d)
(i) diffusion of transmitter molecules
across cleft
synaptic vesicles fuse transmitter molecules attach to

with presynaptic receptor sites
membrane and rupture
molecules of transmitter
substance channels open up and allow ions to
(e) enter from synaptic cleft
propagated action (i.e. permeability of postsynaptic
potential membrane increases)
local
depolarisation
hydrolytic enzymes break down transmitter

molecules, e.g. acetylcholinesterase
Dil
Electrical synaptic transmission along the sarcolemma and down into the fibre via the
transverse tubule system (T-system) (section 17.4.7). This
Transmission across some synapses in many animals,
action potential results in the initiation of muscular
including those of the coelenterate nerve net and verte-
contraction.
brate nervous systems, occurs by the flow of an electrical
current between the pre- and postsynaptic membranes. Functions of synapses and neuro-
The gap between these membranes is only 2 nm and the muscular junctions
combined resistance to the flow of current by membranes
The primary function of neuronal synapses and neuro-
and fluid in the cleft is very low. There is no delay in
muscular junctions within the vertebrate nervous system is
transmission time across the synapses and they are not
the transmission of information between receptor and
susceptible to the action of drugs or other chemicals.
effector. Several other significant functional features arise
Neuromuscular junction out of the structure and organisation of these sites of
The neuromuscular junction is a specialised form of chemical secretion. They are summarised as follows.
synapse found between the nerve terminals of a motor (1) Unidirectionality. The release of transmitter substance
neurone and the endomysium of muscle fibres (section at the presynaptic membrane, and the location of
17.4.2). Each muscle fibre has a specialised region, the receptor sites on the postsynaptic membrane, ensure
motor end-plate, where the axon of the motor neurone that nerve impulses pass in one direction along a given
divides and forms non-myelinated branches 100 nm wide pathway. This gives precision to the nervous system.
running in shallow troughs on the membrane cell surface. (2) Amplification. Sufficient acetylcholine is released at
The muscle cell membrane, the sarcolemma, has many the neuromuscular junction by each nerve impulse to
deep folds called junctional folds as shown in fig 16.11. The excite the postsynaptic membrane to produce a
cytoplasm of the motor neurone axon terminal has the propagated response in the muscle fibre. Thus nerve
same contents as the synaptic knob, and on stimulation impulses arriving at the neuromuscular junction,
releases acetylcholine by the same mechanisms as pre- however weak, are adequate to produce a response
viously described. Changes in the structure of receptor sites from the effector, thereby increasing the sensitivity of
on the sarcolemma increase the permeability of the the system.
sarcolemma to Na” and K" ions and a local depolarisation (3) Adaptation or accommodation. The amount of trans-
known as end-plate potential (EPP) is produced which is mitter substance released by a synapse steadily falls off
sufficient to lead to a propagated action potential passing in response to constant stimulation until the supply of
axon of motor neurone

covered by Schwann cell motor end-plate
secondary synaptic
Schwann cell clefts
mitochondrion S
synaptic vesicles Sebe
muscle surface
presynaptic membrane
of motor neurone
synaptic cleft
junctional fold of sarcolemma

(postsynaptic membrane)
tubule ofT system
GED
mitochondrion
myofibril
Z disc
Fig 16.11 Diagram showing the structure of a motor end plate and neuromuscular junction. The upper part of the diagram
above X — X shows the course of the axon of a motor neurone on the surface of th e muscle — the mot
the diagram below X -— X shows the ultrastructure of the neuromuscular junction Ete Oreiat an
572
transmitter substance is exhausted and the synapse is Table 16.2. Summary table of chemical substances
described as fatigued. Further information passing affecting the synapse and neuromuscular junction in
along this pathway is inhibited and the adaptive mammals
significance of fatigue is the prevention of damage to an
effector due to overstimulation. Adaptation also Substance Site of action Function
occurs at the level of the receptor and this is described
in section 16.4.2. acetylcholine vertebrate excitation or
nervous system inhibition
(4) Integration. A postsynaptic neurone may receive
gamma aminobutyric mammalian brain inhibition
impulses from a large number of excitatory and acid(GABA)
inhibitory presynaptic neurones. This is known as dopamine
synaptic convergence and the postsynaptic neurone is serotonin (5-
hydroxytryptamine) mammalian brain excitation
able to summate the stimuli from all the presynaptic noradrenaline
neurones. This spatial summation enables the synapse lysergic acid mammalian brain produce hallucinations
to act as one region for the integration of stimuli froma diethylamide(LSD) \ by mimicking the
variety of sources and the production of a coordinated mescaline actions, or acting as
antagonists, of other
response. Facilitation occurs at some synapses and this transmitter substances
involves each stimulus leaving the synapse more tetanus toxin presynaptic prevents release of
responsive to the next stimulus. In this way subsequent membrane inhibiting transmitter
reduced stimuli may evoke a response and this is used substance
to increase the sensitivity of certain synapses. Facilita- botulinum toxin presynaptic prevents release of
membrane acetylcholine
tion is not temporal summation in that it is a chemically
nicotine postsynaptic mimics action of
mediated response of the postsynaptic membrane and membrane acetylcholine
not an electrical summation of postsynaptic membrane eserine postsynaptic inactivates acetylcho-
potentials. strychnine membrane linesterase and
organophosphorus prevents breakdown of
(5) Discrimination. Temporal summation at synapses weedkillers and acetylcholine
enables weak background stimuli to be filtered out insecticides
before it reaches the brain. For example, information curare postsynaptic blocks action of
from exteroceptors in the skin, the eyes and ears membrane of acetylcholine
neuromuscular
receive constant stimuli from the environment which junction
has little immediate importance for the nervous atropine parasympathetic blocks action of
system. Only changes in intensity of stimuli are postganglionic acetylcholine
significant to the nervous system and these increase the endings
frequency of stimuli and pass across the synapse and muscarine parasympathetic mimics action of
postganglionic acetylcholine
evoke a response. endings
(6) Inhibition. The transmission of information across
synapses and neuromuscular junctions may be
prevented postsynaptically by the activity of certain disorders such as anxiety and depression are believed to be
chemical blocking agents, described in the next effective due to their ability to interact with chemical
section, or presynaptically. Presynaptic inhibition transmission at synapses. Many tranquillisers and seda-
occurs at synaptic knobs that are in close contact with tives, such as the tricyclic anti-depressant imipramine and
synaptic knobs from inhibitory synapses. Stimulation reserpine and monoamine oxidase inhibitors, exert their
of these inhibitory synapses reduces the number of effects by interacting with transmitter substances or their
synaptic vesicles released by the inhibited synaptic receptor sites. For example, monoamine oxidase inhibitors
knob. This arrangement enables a given nerve terminal prevent the activity of an enzyme involved in the
to produce a variable response depending upon the breakdown of adrenaline and noradrenaline and, presu-
activity of its excitatory and inhibitory synapses. mably, are effective in treating depression by prolonging
the effects of these transmitter substances. Hallucinogenic
Chemical influences on the synapse and drugs, such as lysergic acid diethylamide (LSD) and
neuromuscular junction mescaline, are believed to produce their effects by either
Chemical substances carry out a variety of different mimicking the actions of naturally occurring brain trans-
functions in the nervous system. The effects of some mitter substances or having antagonistic effects on other
chemical substances are widespread and well known, such transmitter substances.
as the excitatory effects of acetylcholine and adrenaline, Recent research into the activity of the pain-suppressing
whereas the effects of other substances are localised and, as
opiate drugs, heroin and morphine in the mammalian brain
yet, require further clarification. Some of these substances have revealed the presence of naturally occurring (endo-
and their functions are described in table 16.2. genous) substances having similar effects. These substances
Several drugs used in the alleviation of psychiatric which react with the opiate receptors are collectively called
Dh
endorphins. Many so far have been identified and the best motor neurones and are described as mixed nerves. Further
known are a group of low relative molecular mass peptides details of spinal nerves and the spinal cord are given below.
known as enkephalins, for example metenkephalin and Cranial nerves arise from the ventral surface of the brain
8-endorphin. They are thought to reduce pain, influence and, with one exception, supply receptors and effectors of
emotion and are involved with certain types of mental the head. There are 10 pairs of cranial nerves in lower
illness. vertebrates and 12 in mammals, numbered I—-XII in Roman
This research has opened up new ideas on brain numerals. Not all cranial nerves are mixed (table 16.3).
functioning and offers a biochemical basis for the control of The tenth cranial nerve, the vagus, includes an important
pain and healing by such diverse activities as hypnosis, motor nerve of the autonomic nervous system supplying
acupuncture and faith healing. Many more chemical the heart, bronchi and alimentary canal.
substances of this type have yet to be isolated, identified
and have their function determined. As techniques of 16.2.2 Reflex action and reflex arcs
extracting and analysing substances found in such minute The simplest form of irritability associated
quantities continue to improve, it is only a matter of time with the nervous system is reflex action. This is a rapid,
before they will help to provide a more complete automatic stereotyped response to a stimulus and because
understanding of brain activity. it is not under the conscious control of the brain it is
described as an involuntary action. The neurones forming
16.2 The vertebrate nervous system the pathway taken by the nerve impulses in reflex action
make up a reflex arc. The simplest reflex arc -found in
The nervous system of vertebrates is animals involves a single neurone and the following
characterised by the structural and functional diversity of pathway:
its neurones and their complex organisation within the neurone
stimulus o_>),€- receptor ——————» effector mmm response
body.
There are several systems of classification of the This level of organisation characterises the nervous system
vertebrate nervous system and all have advantages and of coelenterates. Reflex arcs in all animal groups showing a
limitations. The system given in fig 16.12 divides the greater level of structural and functional complexity than
peripheral nervous system according to the position of coelenterates include at least two neurones, an afferent or
innervation in the body. Internuncial neurones, modified sensory neurone (a, towards) carrying impulses from a
nerve cells of the brain and sense organs are not included in receptor towards an aggregation of nervous tissue which
this classification. may be a ganglion, nerve cord or the central nervous
system, and an efferent or motor neurone (e, away from)
16.2.1 The peripheral nervous system
carrying impulses away from this aggregation to an effector
Spinal nerves arise from the spinal cord and (fig 16.13). There is a wide range of reflexes showing
emerge between adjacent vertebrae along most of the varying structural and functional complexity broadly
length of the spinal cord. They all carry both sensory and involving four courses of action.
vertebrate nervous system
central nervous system (CNS) peripheral nervous system
visceral nervous somatic nervous
ie
system (autonomic) system (voluntary)
brain spinal cord
sympathetic parasympathetic
a
nervous system nervous system
Fig 16.12 Outline classification of
the vertebrate nervous system. (All
spinal nerves and some cranial
nerves have both sensory and
motor components) spinal nerves only spinal nerves cranial nerves spinal nerves cranial nerves
574
Table 16.3. Summary of mammalian cranial nerves, their innervations
and functions
<<< — titi
EEE——————————————
eee
Cranial nerve Name Type Innervation Function

I olfactory sensory olfactory organ smell
II optic sensory retina sight
Ill oculomotor motor four eye muscles eye movements
IV trochlear motor one eye muscle eye movements
Vv trigeminal mixed jaw muscles, teeth, skin of face jaw movements, touch and pain receptors
VI abducens motor one eye muscle eye movements
VII facial mixed cheek, face muscles, tongue salivation, facial expression, sweet, sour, salt
taste
VIII auditory sensory cochlea, semicircular canals hearing, balance
Ix glossopharyngeal mixed tongue, pharyngeal muscle bitter taste, swallowing
x vagus mixed larynx, pharynx, heart, gut speech, swallowing, decrease heart rate,
stimulus for peristalsis
XI accessory motor head and neck head movement
XII hypoglossal motor tongue tongue movement
(1) Monosynaptic reflex. This is the simplest reflex arc seen arc: an internuncial (intermediate or relay) neurone.
in vertebrates. The sensory neurone synapses directly The synapses are found between the sensory neurone
on to the motor neurone cell body. Only one synapse in and internuncial neurone and between the internuncial
the central nervous system is involved in this arc. These neurone and the motor neurone as shown in fig 16.13b.
reflexes are common in vertebrates and are involved in This type of reflex arc provides a simple illustration of
the control of muscle tone and posture, such as the localised reflex action within the spinal cord. Fig 16.14
knee jerk or patellar reflex. In these reflex arcs no shows a much simplified example of the spinal reflex
neurones pass to the brain and reflex actions are associated with the reflex action following pricking a
carried out without the involvement of the brain finger on a pin.
because they are routine and no conscious thought or Simple reflex arcs such as (1) and (2) allow the body to
decision is required for their operation. They are make automatic involuntary homeostatic adjustments to
economical on the number of neurones in the central changes in the external environment, such as the iris—pupil
nervous system and do not make trivial demands on the reflex and balance during locomotion, and also in the
brain which can ‘concentrate’ on more important internal environment, such as breathing rate and blood
matters. pressure, and to prevent damage to the body as in cuts and
(2) Polysynaptic spinal reflex. This has at least two burns.
synapses situated within the central nervous system as (3) Polysynaptic spinal/brain reflexes. Here the sensory
a result of the inclusion of a third type of neurone in the neurone synapses in the spinal cord with a second
(1) stimulus SPINAL CORD stimulus

white matter | grey matter
|
|
|
|
(3) sensory
neurone
(2) receptor
internuncial neurone
(5) effector
(4) motor
neurone
(6) response (a) (b) response
Fig 16.13 Diagrammatic representation of two simple forms of reflex arc, (a) monosynaptic reflex arc, including the main
features of a reflex arc (numbered 1 to 6) (b) simple polysynaptic reflex arc. (R represents receptor, E represents effector)
S75
cell body dorsal root synapse
(3) sensory neurone
ganglion
direction of white matter (axons

impulse running to and from
aaa brain)
sensory nerve
(1) pin (stimulus) ending in skin internuncial neurone
(2) (receptor) sets
up nerve impulse
4 grey matter (cell
bodies and relay
(4) motor neurones)
neurone /
ventral root
synapse (transmission
from neurone to neurone) TS spinal cord
finger pulled away from biceps muscle — contracts

pin (response) (6) (effector) (5)
triceps muscle — relaxes
(effector) (5)
any
Fig 16.14 A simplified example of reflex action and a reflex arc. (The numbers in brackets refer to the basic structures in
reflex arc shown in Fig 16.13)
sensory neurone which passes to the brain. The latter Fig 16.15 Simplified diagram of sections through the brain
sensory neurones are part of the ascending nerve fibre and spinal cord showing (a) the pathway of sensory impulses
from receptor, via the spinal cord, to the cerebral cortex, and
tract and have their origin in the pre-internuncial
(b) the pathway of motor impulses, initiated within the cortex
neurone synapse (fig 16.15a). The brain identifies this to effector, via the spinal cord
sensory information and stores it for further use.
Alternatively motor activity may be initiated at any
time by the brain and impulses transmitted down
motor neurones in descending nerve fibre tracts to cerebral cortex
synapse directly with spinal motor neurones in the
post-internuncial synaptic region (fig 16.15b).
thalamus
(4) Conditioned reflexes. These are forms of reflex actions basal ganglion
where the type of response is modified by past
experience. These reflexes are coordinated by the
brain. Learning forms the basis of all conditioned
reflexes, such as in toilet training, salivation on the
sight and smell of food and awareness of danger (a) ascending (b) descending
sensory neurone motor neurone
(section 16.9).
dorsal horn white matter
Many simple reflex situations arise where there are
two immediate responses involving the activity of a lateral horn grey matter
given set of muscles which can either contract or relax

and produce opposite responses. The normal spinal
reflex response in such situations would pass through
the reflex arc shown in fig 16.14, but ‘conditions’
associated with the stimulus may modify the response.
In these situations more complex reflex pathways exist root ventral spinal
ventral horn centralcanal ganglion root nerve
involving excitatory and inhibitory neurones.
For example, if any empty metal baking tin is picked In this situation reflex pathways exist as shown in fig
up and found to be extremely hot, burning the fingers, 16.16.
it will probably be dropped immediately whereas a The stimulus in both cases produces impulses
boiling hot, cooked casserole in an expensive dish, passing to the brain in an ascending sensory neurone.
equally hot and painful, will probably be put down, When the information reaches the brain it is inter-
quickly but gently. The reason for the difference in the preted and associated with related sensory information
response reveals the involvement of conditioning and coming from other sense organs, for example the eyes,
memory, followed by a conscious decision by the brain. concerning the cause of the stimulus. The incoming
576
inhibitory motor neurone
of body functions is much more complex. For example,
excitatory motor neurone neurones connect different levels of the spinal cord
together, controlling say the arms and legs, so that activity
ascending sensory
neurone in one can be related to the other whilst at the same time
|
other neurones from the brain achieve overall control.
afferent Whilst combined activity of the brain and endocrine
sensory
@) neurone system is important in the coordination of many nervous
activities described later in the chapter, another reflex
) ©) system, based solely on neuronal activity, exists for the
control of visceral activities. This is the autonomic nervous
| . system.
ee
16.2.3 The autonomic nervous system

+
SO The autonomic nervous system (autos, self;

as internuncial efferent nomos, governing) is that part of the peripheral nervous
neurone motor system controlling activities of the internal environment
(excitatory) neurone
that are normally involuntary, such as heart rate, peristalsis
Fig 16.16 Simplified diagram showing the relationships and sweating. It consists of motor neurones passing to the
within the spinal cord between an internuncial neurone and smooth muscles of internal organs. Most of the activity of
the modifying effects of excitatory and inhibitory motor the autonomic nervous system is integrated locally within
neurones from the brain
the spinal cord or brain by visceral reflexes and does not
involve the conscious control of higher centres of the brain.
information is compared with stored information However, some activities, such as the control of anal and
concerning the nature and cause of the present bladder sphincter muscles, are also under the conscious
stimulus and the likely outcome of allowing the spinal control of the brain and control of these has to be learned.
reflex to proceed. In the case of the metal tin, the brain It is thought that many other autonomic activities may be
computes that no further damage to either the body or able to be controlled by conscious effort and learning:
the tin will occur if it is dropped and so initiates many forms of meditation and relaxation have their
impulses in an excitatory motor neurone. This passes physiological roots in the control of autonomic activities,
down the spinal cord to the level where the stimulus and considerable success has already been achieved in
entered and synapses with the cell body of the motor regulating heart rate and reducing blood pressure by
neurone of the spinal reflex. Such is the speed of conscious control or ‘will power’. The overall control of the
conduction through the pathway described above that autonomic nervous system is maintained, however, by
the impulses from the excitatory motor neurone reach centres in the medulla and hypothalamus (see section
the spinal motor neurone at the same time as impulses 16.2.4). These receive and integrate sensory information
from the internuncial neurone. The combined effect of and coordinate this with information from other parts of
these sends excitatory impulses to the muscle effector the nervous system to produce the appropriate response.
along the spinal motor neurone and the tin is dropped. The autonomic nervous system is composed of two types
In the case of the casserole dish, the brain computes of neurones, an unmyelinated preganglionic neurone,
that dropping the casserole would probably scald the which leaves the central nervous system in the ventral root
legs and feet, ruin the meal and break an expensive of the segmental nerve before synapsing with several
dish, whereas holding it until it could be put down unmyelinated postganglionic neurones leading to effectors.
safely would not cause much more damage to the There are two divisions of the autonomic nervous
fingers. If this decision is reached, impulses are system: the sympathetic and the parasympathetic nervous
initiated which pass down the spinal cord in an systems. The two systems differ primarily in the structural
inhibitory motor neurone. These impulses arrive at the organisation of their neurones and these differences are
synapse with the spinal motor neurone at the same time shown in fig 16.17.
as stimulatory impulses from the internuncial neurone In the sympathetic nervous system the synapses and cell
and the latter are cancelled out. No impulses pass along bodies of the postganglionic neurones in the trunk region
the motor neurone to the muscle effector and the dish are situated in ganglia close to the spinal cord. Each
is held. Simultaneous brain activity would initiate an sympathetic ganglion is connected to the spinal cord by a
alternative muscle response which would result in the white ramus communicans and to the spinal nerve by a grey
dish being put down quickly and safely. ramus communicans as shown in fig 16.18. Adjacent
The accounts of reflex arcs and reflex activity given segmental sympathetic ganglia on each side of the spinal
cord are linked together by the sympathetic nerve tract to
above are, of necessity, simplified generalisations. The
form a chain of sympathetic ganglia running alongside the
whole process of the coordination, integration and control
OTT
visceral sensory neurones
effector
ffect ay
errec aeons Die ° —)
——————— Ad
QoS
ACh ACh
parasympathetic ganglion ¢ Pk es (sympathetic ganglion
S situated centrally)
(situated peripherally)
preganglionic neurones
(a)
Fig 16.17 Simplified diagram showing the basic features of preganglionic neurone
(a) the parasympathetic nervous system and (b) the postganglionic neurone
sympathetic nervous system (visceral sensory neurones are ACh acetylcholine
not part of the autonomic nervous system) Ad _ noradrenaline
Fig 16.18 Simplified diagram showing the position of a normal level when secretion from both systems balances
sympathetic ganglion and its relationship to the spinal cord out. A summary of the antagonistic effects of these systems
and spinal nerve is shown in table 16.5.
grey
matter
white 16.2.4 The central nervous system
wae postganglionic
sympathetic The central nervous system (CNS) develops
neurone
from an infolding of the ectoderm immediately above the
embryonic notochord and forms a dorsal, hollow neural
tube running the length of the animal. The neural tube
differentiates during development to form an expanded
anterior region, the brain, and a long cylindrical spinal
cord.
preganglionic
ventral root
The central nervous system is covered by three mem-
sympathetic
neurone grey ramus branes called meninges and is completely encased within
communicans the protective bones of the skull and vertebral column. The
outer membrane forms the tough dura mater attached to
sympathetic the periosteum of the skull and vertebrae, and the inner pia
white ramus ganglion mater which directly overlies the nervous tissues. Between
communicans
the two is the arachnoid ‘membrane’, composed of pillars
spinal cord. The ganglia of the parasympathetic nervous of connective tissue supporting a space beneath it, the
system are situated close to, or within, the effector organ. subarachnoid space, which contains the cerebrospinal fluid
Other differences between the two systems include the (CSF). Most of this fluid is contained in the central canal of
nature of the chemical transmitter substance released at the the spinal cord and continues forward to occupy four
postganglionic effector synapse, their general effects on the expanded regions, the ventricles, of the brain. The fluid
body and the conditions under which they are active. These bathes the outside and inside of the brain and blood vessels
differences are summarised in table 16.4. lie within it for the supply of nutrients and oxygen to the
The sympathetic and parasympathetic nervous systems nervous tissues and the removal of wastes (see fig 16.19).
generally have opposing effects on organs they supply and The cells of the vascular anterior and posterior choroid
this enables the body to make rapid and precise adjust- plexuses in the roof of the brain secrete CSF and provide a
ments of visceral activities in order to maintain a steady link between the fluid outside and inside the brain. About
state. For example, an increase in heart rate due to the 100 cm’ of fluid is present in the CNS and, apart from its
release of noradrenaline by sympathetic neurones is nutritive and excretory functions, it supports the nervous
compensated for by the release of acetylcholine by tissues and protects them against mechanical shock. A
parasympathetic neurones. This action prevents heart rate continual circulation of fluid is maintained by ciliated cells
becoming excessive and will eventually restore it to its lining the ventricles and central canal.
578
Table 16.4. Summary of the differences between the sympathetic and parasympathetic nervous systems
eee
Feature Sympathetic Parasympathetic

Origin of neurones Emerges from cranial, thoracic and lumbar regions Emerges from cranial and sacral regions of CNS
of CNS
Position of ganglion Close to spinal cord Close to effector
Length of fibres Short preganglionic fibres Long preganglionic fibres
Long postganglionic fibres Short postganglionic fibres
Number of fibres Numerous postganglionic fibres Few postganglionic fibres
Distribution of fibres Preganglionic fibres innervate a wide area Preganglionic fibres innervate a restricted
region
Area of influence Effect diffuse Effect localised
Transmitter substance Noradrenaline released at effector Acetylcholine released at effector
General effects Increases metabolite levels Decreases metabolite levels
Increases metabolic rate None
Increases rhythmic activities Decreases rhythmic activities
Lowers sensory threshold Restores sensory threshold to normal levels
Overall effect Excitatory homeostatic effect Inhibitory homeostatic effect
Conditions when active Dominant during danger, stress and activity; controls Dominant during rest
reactions to stress Controls routine body activities
Table 16.5 Summary of the effects of the sympathetic and parasympathetic nervous systems on the body
Region Sympathetic Parasympathetic
Head Dilates pupils Constricts pupils

None Stimulates secretion of tears
Inhibits secretion of saliva Stimulates secretion of saliva
Heart Increases amplitude and rate of heart beat Decreases amplitude and rate of heart beat
Lungs Dilates bronchi and bronchioles Constricts bronchi and bronchioles
Increases ventilation rate Decreases ventilation rate
Gut Inhibits peristalsis Stimulates peristalsis
Inhibits secretion of alimentary juices Stimulates secretion of alimentary juices
Contracts anal sphincter muscle Inhibits contraction of anal sphincter muscle
Blood Constricts arterioles to gut and smooth muscle Maintains steady muscle tone in arterioles to gut, smooth
Dilates arterioles to brain and skeletal muscle muscle, brain and skeletal muscle
Increases blood pressure Reduces blood pressure
Increases blood volume by contraction of spleen None
Skin Contracts erector pili muscles of hair None ;
Constricts arterioles in skin of limbs Dilates arterioles in skin of face
Increases secretion of sweat None
Kidney Decreases output of urine None
Bladder Contracts bladder sphincter muscle Inhibits contraction of bladder sphincter muscles
Penis Induces ejaculation Stimulates erection
Glands Releases adrenaline from adrenal medulla None
dura mater
arachnoid layer
subarachnoid space
blood vessel
arachnoid trabecula
pia mater
perivascular space Fig 16.19 Diagram showing the structure of the meninges
and associated blood vessels. Notice that the channels
eweito ots through the nerve tissue are lined by the pia mater
a79
The functions of the central nervous system involve the and the complexity of their interconnections have in-
coordination, integration and control of most nervous creased. The ‘nuclei’ vary in size from isolated groups of
activities and it works in conjunction with the peripheral several hundred cells to large regions such as the cerebral
nervous system. In higher organisms, advanced forms of cortex and the cerebellar cortex in Man consisting of
nervous activity, such as memory and intelligence, are several hundred million cells.
possible due to the increased size of certain regions of the The
The structure of the vertebrate brain.
brain.
vertebrate brain differentiates initially into three regions
The spinal cord during its embryological development and these are the
The spinal cord is a dorso-ventrally flattened cylinder of forebrain, midbrain and hindbrain. Primitively, these
nervous tissue running from the base of the brain to the three regions are associated with the coordination of the
lumbar region and protected by vertebrae. It consists of an senses of smell, sight and balance respectively. Subsequent
H-shaped central core of grey matter, composed of nerve development of the brain varies between and within each
cell bodies, dendrites and synapses surrounding a central vertebrate class, and the original tripartite structure
canal, and an outer layer, the white matter, containing becomes obscured as the forebrain and hindbrain each
nerve fibres whose fatty myelin sheaths give it its subdivide. The adult vertebrate brain has five regions, and
characteristic colour. There are 31 pairs of segmental spinal they assume a particular significance within each class
nerves present and these divide close to the spinal cord to associated with the mode of life and level of structural and
form two branches called the dorsal root and ventral root. functional complexity attained by the class. Some regions
Sensory neurones enter the dorsal root and have their cell of the brain have increased in size reflecting their
bodies in a swelling, the dorsal root ganglion, close to the importance whereas others have diminished. The overall
spinal cord. The sensory neurones then enter the dorsal structure of the generalised vertebrate brain is shown in fig
horn of the grey matter where they synapse with internun- 16.20 and a summary showing the development of the
cial (intermediate or relay) neurones. These, in turn, regions of the brain in non-mammalian and mammalian
synapse with motor neurones in the ventral horn and leave vertebrates is given in table 16.6.
the spinal cord via the ventral root (fig 16.15). Since there Fish are very dependent upon smell in order to find food
are many more internuncial neurones than motor and have large olfactory lobes to accommodate the sensory
neurones, some integration must occur within the grey input. A large cerebellum coordinates movements, and the
matter. Some sensory neurones synapse directly with large optic lobes act partly in visual responses but mainly as
motor neurones in the ventral horn, as in the familiar the main coordination centre of the brain. The optic lobes
knee-jerk reflex (fig 16.13a). In the thoracic, upper lumbar form the dominant feature of the amphibian brain where
and sacral regions a lateral horn is present (fig 16.15) they have the same functions as in fish. Reptiles show a
containing the cell bodies of the preganglionic autonomic reduction in the size of the midbrain and an increase in the
neurones. The white matter is composed of groups of nerve size of the forebrain. This trend continues in birds with an
fibres, forming tracts, running between the grey matter and increase in the sizes of the thalamus and corpora striata
the brain and providing a means of communication where most of the complex, instinctive behavioural
between spinal nerves and the brain. Ascending tracts carry activities of birds are coordinated. Finally, mammals are
sensory information to the brain and descending tracts characterised by a large pair of cerebral hemispheres
relay motor information to the spinal cord. covering a well-developed thalamus. These two structures,
The functions of the spinal cord include acting as a containing nuclei and tracts respectively, reflect the
coordinating centre for simple spinal reflexes, such as the dependence placed on the storage of sensory information
knee-jerk response, and autonomic reflexes, such as and the integration of all voluntary activities. The medulla
contraction of the bladder, and providing a means of shows little change in relative size throughout the verte-
communication between spinal nerves and the brain. brates and this stresses the importance placed on the reflex
control of all essential functions such as heart rate.
The brain
The brain is the swollen anterior end of the vertebrate Methods of studying brain function
neural tube and it coordinates and controls the activities of Little knowledge of the functions of the various regions of
the whole nervous system. The brain is composed entirely the brain can be obtained by simply studying their anatomy
of groups of cell bodies, nerve tracts and blood vessels. The and histology. Whilst this provides valuable information
nerve fibre tracts form the white matter of the brain and about structural interrelationships, it does not explain the
carry bundles of neurones to and from various regions physiological activities occurring in the brain. Brain
called ‘nuclei’ or centres composed of groups of cell bodies activity has been studied using electrophysiological techni-
and synapses collectively forming the grey matter of the ques, including the use of electroencephalograms. Good
brain. The tracts connect the various ‘nuclei’ together and electrical contact is made between electrodes and the scalp
link the brain and spinal cord. During the phylogenetic using electrode jelly and electrical changes produced by the
development of the vertebrate brain the number of tracts activity of many cells of the cerebral cortex are detected,
580
forebrain midbrain hindbrain
telence phalon i
diencephalon | mesencephalon | metencephalon | myelencephalon
. 1 |
anterior |
choroid
cerebrum | plexus |
| |
thalamus optic lobe cerebellum | posterior
ry ; \ (tectum) } choroid plexus
ee — pineal ! |
2 \ body
us lateral si |
ag ventricles x
ep D Se BS \
Seay ‘
eo |
Se [
a \ |
=~ I ! is
| ! | '
spinal cord
olfactory aoa nsire<i i
lobe
!
medulla
| hypothalamus , pons oblongata |
Fig 16.20 Diagram showing a median section of a generalised vertebrate brain. The numbers refer to the positions of the \lIrd
and |Vth ventricles
Table 16.6. The origins and regions of the vertebrate brain
Structural/functional regions
Embryonic Adult Non-mammal Mammal
division region
olfactory lobes (fish .

telencephalon recente nana Cee price
corpora striata (birds) basal ganglia
forebrain
thalamus thalamus
diencephalon
hypothalamus hypothalamus
optic lobes (amphibia)

midbrain mesencephalon
tectum corpora quadrigemina
cerebellum cerebellum
metencephalon
hindbrain pons pons
[
myelencephalon medulla oblongata medulla oblongata
ee
amplified and recorded on a pen trace. Three major wave CNS and occurs in two forms. The first, grand mal produces
frequencies have been detected. a-waves are recorded powerful convulsions lasting from several seconds to
from relaxed subjects with their eyes closed and they minutes whereas the second, petit mal produces mild
usually mask the higher frequency 6-waves which are convulsions lasting for much shorter periods of time.
present at all times and best seen in patients under Investigations of brain function are carried out on the
anaesthesia. The absence of $-waves or any electrical brains of patients under local anaesthesia. In one type of
activity from the brain of a patient indicates ‘brain-stem- investigation patients describe sensations produced whilst
death’ and, along with cessation of heart beat and various regions of the brain are electrically stimulated. This
ventilation is taken as a Clinical definition of death. 5-waves technique enables motor activities of regions of the
have the lowest frequency and greatest amplitude and are cerebral cortex to be mapped (fig 16.23). Likewise
recorded during sleep. Whilst electroencephalograms yield recordings of electrical activity obtained using microelec-
little immediate information regarding brain function, they trodes inserted into cell bodies or axons in specific regions
are valuable in diagnosing the position of localised brain during stimulation of sensory cells and organs have enabled
abnormalities, and, for example, the various forms of maps to be drawn showing regions of brain sensory
epilepsy. Epilepsy is a result of excessive activity of the function.
581
The structure and functions of the human Telencephalon
brain This is the anterior region of the forebrain and consists of
The following account of the brain is brief and represents a the cerebrum and basal ganglia. The cerebrum forms the
summary of current knowledge of the structures and roof and walls of the telencephalon and is greatly enlarged
functions of the human brain. Elaboration of function and to form the left and right cerebral hemispheres covering
evidence for this is omitted except where it is considered most of the brain. The basal ganglia occupy the floor of the
necessary for clarification. In this chapter the brain is telencephalon.
considered from astructural standpoint and the activities of
Cerebrum. The cerebral hemispheres are
each region shown in table 16.6 are described in turn. By
composed of a thin outer layer (3 mm) of densely packed
necessity this approach and account is a gross oversimp-
nerve cells (10°) forming a region of grey matter called the
lification since many brain activities span several regions,
cerebral cortex. Beneath this is a central mass of white
and it is useful to bear in mind that functionally there are
matter composed of nerve fibre tracts. Left and right
three main interrelated regions, the cerebrum (cerebral
hemispheres), the cerebellum and the brain stem, the latter cerebral hemispheres are linked by a broad nerve fibre tract
being composed of the medulla, pons, mesencephalon and called the corpus callosum and the surface area of the
thalamus. The brain stem is an extension of the spinal cord cortex is increased by numerous infoldings called convolu-
and contains complex neuronal pathways controlling tions. Each cerebral hemisphere is divided, for conve-
cardiovascular function, ventilation, gastrointestinal func- nience, into four lobes as shown in fig 16.23, and
tion, eye movement, equilibrium and most of the electrophysiological techniques have enabled three dis-
stereotyped activities of the body. crete areas to be recognised according to the functions their
cells perform. These areas are:
(1) sensory — receiving impulses indirectly from receptors
X
cerebral
(input),
—
hemisphere (2) association — interpreting the input, storing the input
corpus callosum cerebral and initiating a response in the light of similar past
;
lateral ventricles
cortex
‘
experience, and
subarachnoid (3) motor — transmitting impulses to effectors (output).
space
thalamus
anterior The interrelationships between the above areas enable
hypothalamus choroid plexus the cerebral cortex to dominate and coordinate all
pineal body voluntary and some involuntary activities of the body,
pituitary body
corpora including highly developed functions such as memory,
quadrigemina
learning, reasoning, conscience and personality. If the
cerebellum
posterior
cerebral cortex is entirely destroyed it would not cause
choroid death but the patient would show no spontaneous activity.
medulla oblongata plexus The patient would be able to respond to certain stimuli but
spinal cord would be unable to learn or reason and all visible signs of
central canal
intelligence and personality would disappear. Only those
reflexes controlled predominantly by the medulla and
cerebellum, such as feeding and sleeping, would remain.
Fig 16.21 (above) Simplified diagram showing a vertical Sensory areas are the input areas of the cortex and
section through the human brain (the numbers indicate the receive sensory impulses via ascending tracts from recep-
ventricles of the brain)
tors originating from most parts of the body. The sensory
areas form localised regions of the cortex associated with
certain senses as shown in fig 16.23. The size of the region is
Fig 16.22 (below) Simplified diagram showing a cross-
section of the human brain through X — X on Fig. 16.21
related to the number of receptors in the sensory structure.
Association areas are so named for several reasons. First,
they associate incoming sensory information with previous-
cerebral cortex
ly perceived information stored in memory units, so that
corpus callosum
the information is ‘recognised’. Secondly, the information
lateral ventricle
is associated with incoming sensory information from other
subarachnoid space
receptors. Thirdly, the information is ‘interpreted’ and
basal ganglion
given meaning within its present context and, if necessary,
IIIrd ventricle the interpreted information is associated with the ‘com-
puted’ most appropriate response which the association
area initiates and passes to its associated motor area.
hypothalamus thalamus Association areas, therefore, are involved in memory,
582
16.23 Diagram showing the positions somatic motor area
of the four main lobes of the brain:
frontal, parietal, occipital and Pern orerarsa somatic senso ry area
temporal. Superimposed on these are
the major: sensory, association and
motor areas and centres. The positions
of certain regions of the brain
associated with specific activities and
regions of the body are also shown speech association area
visual sensory area
speech motor memory

centre
tem
auditory association area
auditory sensory area
learning and reasoning, and the degree of success of the which relays motor neurones from the reticular formation
outcome may be loosely termed intelligence. situated in the brain stem between the thalamus and
Several major association areas are adjacent to their medulla. Motor impulses from various regions of the brain
related sensory area; for example, the visual association controlling muscular activity pass to specific areas of the
area is situated immediately anterior to the visual cortex in reticular formation where this activity is modified by
the occipital lobe where, in a visual context, it carries out impulses from the cortex and becomes either inhibitory or
the association functions described above. Some associa- excitatory. For example, impulses from the cerebellum and
tion areas may have arestricted specialised function and premotor area of the cortex, a region involved in the
these are linked to other association centres that can control of coordinated movements, pass to an area of the
further develop the activity. For example, the auditory reticular formation in the medulla where the combined
association area only interprets sounds into broad categor- effect stimulates inhibitory motor neurones. These have a
ies which are relayed to more specialised association areas suppressive regulatory effect on muscle activity enabling
such as the speech association area where ‘sense’ is made of complex coordinated movements of the body to occur,
the words. Speech is initiated in the speech motor centre such as the varied control involved in angling, directing and
which is an example of the third type of functional area powering a tennis racquet, cricket ball or violin bow,
found in the cerebrum. according to circumstances. Other combinations of motor
Motor areas are the output areas of the cortex where impulses stimulate excitatory motor neurones; in fact, the
motor impulses are initiated to pass to voluntary muscles overall motor output of the reticular formation is excita-
via descending tracts originating in the white matter of the tory.
cerebrum. Most sensory neurones run to the reticular formation
Many motor impulses pass directly to the spinal cord before passing to the cortex via the thalamus. Some of
through two large pyramidal tracts (corticospinal tracts) these sensory neurones form the reticular activating system
via the brain stem. All other motor impulses pass through and are responsible for activating the cortex and arousing
extrapyramidal tracts which contain motor impulses from the body from its natural state of sleep. Underactivity or
other regions of the brain such as the basal ganglia and destruction of the reticular activating system induces deep
cerebellum. In the medulla all tracts cross over so that sleep or coma respectively. Many general anaesthetics are
impulses from the left cerebral cortex innervate the right thought to act by temporarily blocking synaptic transmis-
side of the body and vice versa. sion in this system. The reticular activating system is also
Neurones passing through the pyramidal tracts have believed to be responsible for producing and sustaining
their cell bodies in the motor cortex and their axons pass motivation and concentration hence the inverse relation-
directly to synapse with the motor neurones that they ship between tiredness and concentration.
activate in the spinal segment at the point where the latter Finally, the functions of certain regions of the cortex,
emerge. There are no intervening synapses in the brain, particularly the large anterior regions, the prefrontal lobes,
therefore impulses and subsequent responses are neither are still uncertain. These regions, along with others in the
delayed nor modified en route. Localised regions of the brain, are called silent areas because they fail to produce
motor cortex have been mapped, and examples of these are either sensation or response when stimulated electrically.
shown in fig 16.23. The size of each specific motor area is They are believed to be responsible for our individual
related to the complexity of the motor activity. characteristics or personality. The removal of the lobes or
The main extrapyramidal tract is the reticulospinal tract cutting the tracts leading from them to the rest of the brain
583
(prefrontal lobotomy) was used to relieve acute anxiety Mesencephalon
states in patients but it has been discontinued. The This connects the anterior two regions of the brain to the
side-effects included a reduction in mental awareness, posterior two regions and, therefore, all nerve fibre tracts
intelligence, judgement and creativity and gave an indica- within the brain pass through this region which is part of the
tion of the functions of the prefrontal lobes. brain stem. The roof of the mesencephalon is composed of
Basal ganglia. These are regions of the the tectum consisting of the four corpora quadrigemina and
forebrain containing cell bodies receiving motor neurones these function as visual and auditory reflex centres. The
from parts of the cortex and passing on impulses to the superior pair of corpora quadrigemina receive sensory
reticular formation. The functions of the basal ganglia are neurones from the eyes and muscles of the head and control
varied, but one ganglion, for example, provides inhibitory visual reflexes. For example, they control the movement of
stimuli for the antagonistic control of muscle tone during the head and eyes to fix and focus on an object. The inferior
slow movements. Damage to this basal ganglion produces pair of corpora quadrigemina receive sensory neurones
the tremor of the hands associated with Parkinson’s from the ears and muscles of the head and control auditory
disease, a form of muscle paralysis. reflexes such as the movement of the head to locate and
detect the source of a sound.
Diencephalon Situated in the floor of the mesencephalon are many
This is the posterior region of the forebrain and its dorsal centres or nuclei controlling specific subconscious
and lateral regions form the thalamus and the ventral stereotyped muscular movements, such as bending for-
region forms the hypothalamus. The pineal body arises in wards and backwards and rotation of the head and trunk.
this region and its function is described in section 16.6.3. For example, stimulation of part of one of these nuclei, the
Thalamus. The majority of sensory neur- red nucleus, causes the head and upper trunk to extend
ones carrying impulses to the cortex terminate in the backwards.
thalamus. Here the origin and nature of the impulses are Metencephalon
‘analysed’ and relayed to the appropriate sensory areas of
The dorsal region of the metencephalon forms the
the cortex by neurones originating in the thalamus. It
cerebellum and the ventral region forms the pons.
therefore acts as a processing, integrating and relay centre
for all sensory information. In this function it shows Cerebellum. This is made up of the two
similarities to a switchboard in a telephone exchange. cerebellar hemispheres and, like the cerebrum, has its grey
Information from certain regions of the cortex is modified matter on the outside. The grey matter contains character-
by the thalamus, which is also thought to be involved in the istically large flask-shaped Purkinje cells bearing many
perception of pain and pleasure. Part of the reticular dendrites. These cells receive impulses concerning muscu-
formation, whose function was described above for the lar movement from a number of different sources,
cerebral motor areas, originates in the thalamus. The including sensory receptors in the balance organs (the
dorsal region immediately anterior to the thalamus, the vestibular apparatus) of the ear concerned with balance,
anterior choroid plexus, facilitates transfer of substances proprioceptors in joints, tendons and muscles and motor
between the CSF in the third ventricle and the sub- centres of the cortex. The cerebellum is thought to
arachnoid space (fig 16.21). integrate this information and produce coordinated muscu-
Hypothalamus. This is the main coordinat- lar activity in all the muscles involved in a given movement
ing and control centre for the autonomic nervous system. It including the reflex control of body posture. Damage to the
receives sensory neurones from all the visceral receptors cerebellum results in jerky, poorly controlled movements.
and taste and smell receptors. Information is relayed from The cerebellum is vital to the control of rapid muscular
here to effectors via the medulla and spinal cord, and is activities such as running, typing and even talking. All the
used in the regulation and control of heart rate, blood activities of the cerebellum are involuntary but may involve
pressure, ventilation rate and peristalsis. Other regions of learning in their early stages. During these training periods
the hypothalamus contain specific centres for the initiation the cortex directs the control of the cerebellum and
of feeding, drinking and sleeping, and behavioural activi- concentration is required, such as when learning to walk,
ties associated with aggression and reproduction. The swim or ride a bicycle. Once the skill is acquired reflex
hypothalamus is the most vascular region of the brain and control by the cerebellum takes over.
monitors the metabolite and hormone levels of the blood as Pons. This forms the part of the brain stem
well as blood temperature. Using this information the in the floor of the metencephalon and, apart from acting as
hypothalamus, in association with the pituitary gland a bridge (pons, bridge) carrying ascending and descending
situated immediately beneath it, directs and controls the tracts, it contains several ‘nuclei’ relaying impulses to the
release of most of the hormones from the body, and cerebellum.
maintains the steady-state composition of the blood and
tissues. The detailed neuroendocrine role of the hypothala- Myelencephalon
mus is described in section 16.6.2. This is the posterior region of the brain and is continuous
584
with the spinal cord. It is composed of the dorsally situated
little value to the whole organism unless the stimulus is
posterior choroid plexus and the ventro-lateral medulla
intense or prolonged. In most advanced coelenterates, such
oblongata. as jellyfish and sea anemones, in addition to the nerve net
In the medulla the ascending and descending nerve fibre there is a system of elongate bipolar neurones arranged in
tracts cross over from left to right and vice versa. The tracts, called through conduction tracts, and able to
eighth to twelfth cranial nerves originate from the medulla transmit impulses rapidly over considerable distance and
and it contains important reflex centres for the regulation without apparent loss. This system enables the organism to
of autonomic activities including the control of heart rate make fairly rapid responses of the whole body to harmful
(chapters 14 and 18), blood pressure (chapters 14 and 18), stimuli, such as the withdrawal of tentacles, and this
ventilation rate (chapters 11 and 18), swallowing, saliva- foreshadows the aggregation of neurones into nerves seen
tion, sneezing, vomiting and coughing. in higher organisms.
In the annelids the association of neurones into nerves
has resulted in a nervous system consisting of a single
16.3 The phylogenetic development of longitudinal tract, the ventral nerve cord, running the
the nervous system entire length of the organism. This consists of paired
segmentally arranged ganglia joined by connecting
A study of animal phylogeny shows a progres- neurones and supplying segmental nerves to the tissues of
sive increase in structural and functional complexity from each segment as shown in fig 17.37.
protozoa to mammals. Several trends and patterns in the As a result of the unidirectional method of locomotion,
organisation of organ systems have become established and annelids possess a head. This structure is specialised to
this is clearly reflected in the development of the nervous assist with feeding and, since it is the first part of the body to
system as shown bya study of irritability in protozoa, come into contact with new environmental situations, it
coelenterates, annelids, arthropods and mammals. contains all the sensory structures necessary to detect
In protozoa the ability to respond to a stimulus resides stimuli associated with these situations. The increased
within a single cell of the organism. There is no spatial input of sensory information from these receptors to the
separation between the stimulus and response and the cell nervous system is dealt with by the enlarged anterior end of
functions as both receptor and effector. Investigations the nerve cord. This concentration of feeding apparatus,
carried out on irritability in Amoeba suggested that the sense organs and nervous tissue into one region is called
mechanism of transducing a stimulus, for example prod- cephalisation. It should be emphasised though that the
ding it with a blunt needle, involves the plasma membrane term applies to the development of all the features
and the granular endoplasm. The mechanism probably associated with the head and not just the nervous tissue.
involves the release of energy by an ATP/ATPase system The degree of cephalisation shown by an organism varies
which provides energy for amoeboid movement (section according to the level of structural complexity attained by
17.6.1) thus enabling Amoeba to make an avoidance the organism and its mode of life.
response to the stimulus. The annelid nervous system shows all the basic features
The development of multicellular organisation in the found in all other invertebrate groups. The enlarged
coelenterates has led to an increase in spatial separation of anterior region of the nerve cord forms apair of cerebral
stimulus and response, receptor and effector. Fortunately, ganglia situated above the pharynx and linked to the
the attainment of the multicellular state was accompanied ventral nerve cord by a pair of circumpharyngeal connec-
by tissue differentiation and the appearance of nerve cells tives.
linking receptor to effector, thus overcoming difficulties of In arthropods the basic organisation of the nervous
spatial separation. The nervous system of primitive system is almost identical to that of annelids except that the
coelenterates, for example Hydra, is a nerve net or plexus cerebral ganglia overlie the oesophagus and consequently
composed ofa single layer of short multipolar neurones in are linked to the ventral nerve cord by circumoesophageal
synaptic contact throughout the organism. Impulses spread connectives. Cerebral ganglia are analogous to the verte-
out in all directions from the point of stimulation and at brate brain but do not possess the same degree of
each synapse an impulse is lost. This impulse is used, autonomy over the entire nervous system as seen in
effectively, to ‘charge’ the synapse so that subsequent vertebrates. For example, removal of the head of an
impulses can cross the synapse. This process is called invertebrate has very little effect on movement, whereas in
facilitation and, since an impulse is lost at each synapse in vertebrates the brain initiates and controls all movement of
facilitating (making easier) the passage of the next impulse, the body. Invertebrate cerebral ganglia, in fact, appear to
the mechanism of conduction is called decremental conduc- act simply as relay centres between receptors and effectors
tion. Nervous conduction in these organisms is therefore and their role in integration and coordination is limited to a
slow, due to the number of synapses to cross, and spatially few neuroendocrine responses such as the timing of
restricted because impulses die out as they progress reproductive activities in annelids and the control of
outwards from the stimulus. This system is useful in ecdysis and moulting in arthropods (section 21.7.3).
producing localised responses, say within a tentacle, but of
585
Animals only detect stimuli existing in one of the forms
16.4 Sensory receptors of energy shown in table 16.7. Structures transforming
stimulus energy into electrical responses in axons are
The coordinated activity of an organism relies
known as transducers and, in this respect, receptors act as
upon a continuous input of information from the internal biological transducers.
and external environments. If this information leads to a All receptors transform the energy of the stimulus into a
change in activity or behaviour of the animals, it is a
localised non-propagated electrical response which initi-
stimulus. The specialised region of the body detecting the
ates nerve impulses in the neurone leaving the receptor.
stimulus is known as a sensory receptor.
Thus receptors encode a variety of stimuli into nerve
The simplest and most primitive type of receptor consists
impulses which pass into the central nervous system where
of a single unspecialised primary sense cell composed of a
they are decoded and utilised to produce the required
single sensory neurone whose terminal end is capable of
detecting the stimulus and giving rise to a nerve impulse responses as shown in fig 16.13b. The type of response and
passing to the central nervous system, for example, skin its extent and duration are all directly related to the nature
mechanoreceptors such as the Pacinian corpuscle (section
of the stimulus.
16.5.1). More complex receptors are known as secondary
sense cells, and they consist of modified epithelial cells able
to detect stimuli. These form synaptic connections with 16.4.1 The mechanism of transduction
their sensory neurones which transmit impulses to the
All sensory cells are excitable cells, and they
CNS, for example mammalian taste buds (fig 16.31). The
share with nerve cells and muscle cells the ability to
most complex receptors are sensory organs composed of a
respond to an appropriate stimulus by producing a rapid
large number of sense cells, sensory neurones and
change in their electrical properties. When not stimulated,
associated accessory structures. The mammalian eye and
ear show the level of complexity which is attained by sense sensory cells are able to maintain a resting potential as
described in section 16.1.1, but respond to a stimulus by
organs. In the eye there are two types of secondary sense
cells, rods and cones, many connecting neurones and many producing a change in membrane potential. Bernard Katz,
accessory structures such as the lens and iris. in 1950, using a specialised stretch receptor known as a
The accessory structures often serve the double function muscle spindle was able to demonstrate the presence of a
of eliminating the effects of unwanted stimuli and amplify- depolarisation in the immediate region of the sensory nerve
ing the effects of desired stimuli. ending in the muscle spindle. This localised depolarisation
On the basis of the position of the receptor and the is found only in the sensory cell and is known as the
stimulus three types of receptor are identified: generator or receptor potential. Subsequent investigations
(1) exteroceptors — these respond to stimuli originating involving intracellular recordings, made by penetrating the
outside the body, as with the ear and sound; membranes of receptor cells in muscle spindles and the
(2) interoceptors — these respond to stimuli originating mechanoreceptors of the skin, the Pacinian corpuscles,
inside the body, such as blood pressure and carbon have revealed the following information about transduc-
dioxide receptors in the carotid arteries; tion:
(3) proprioceptors — these respond to stimuli concerned (1) the generator potential results from the stimulus
with the relative positions and movements of muscles producing a non-selective increase in the permeability
and the skeleton. of the sensory cell membrane to sodium and potassium
An alternative, and somewhat preferable, system of ions which flow down their electro-chemical gradients;
classification is based on the type of stimulus detected by (2) the magnitude of the generator potential varies with
the receptor, and this is shown in table 16.7. the intensity of the stimulus;
(3) when the generator potential reaches a predetermined
Table 16.7. Types of receptors and the stimuli detected by threshold it gives rise to a propagated action potential
them in the sensory axon leading from the sensory cell (fig
16.24);
(4) the frequency of the nerve impulses in the sensory axon
Type of receptor Type of stimulus Nature of stimulus
energy is directly related to the intensity of the stimulus.
This latter point was, in fact, established in Cambridge in
photoreceptor electromagnetic light 1926 by Lord Adrian, who demonstrated that sensory
electroreceptor electromagnetic electricity information is carried by all-or-nothing action potentials as
mechanoreceptor mechanical sound, touch, a frequency code. We now know that only the generator
pressure, gravity potential exhibits an amplitude code but, following the
thermoreceptor thermal temperature change attainment of a certain threshold value in the sensory
chemoreceptor chemical humidity, smell, taste neurone, this gives rise to a propagated all-or-nothing
SSS
————— response in the sensory neurone.
586
pressure preamplifier
(stimulus)
Y OM et oe Ly cathode ray oscilloscope

microelectrode I
)
fine glass rod microelectrode II
a axon
Pacinian corpuscle myelin sheath

unmyelinated nerve terminal
concentric rings of connective tissue
Recordings from Recordings from

microelectrode I microelectrode II
0 0|
=() -20
Light Potential difference
pressure across membrane/mV _40 _40
-60 —60
0 0|
=)() -20
Medium mv
pressure _40 _40
60 -60
+ 40
+ 20
0 0
=9() -20
Heavy mV
pressure 40 _40
-60 -60
Fig 16.24 The electrical activity recorded by two microelectrodes (\) and (II) inserted into (\) the axon terminal within a Pacinian
corpuscle and (Il) the axon of the sensory neurone leaving the corpuscle. As the pressure on the fine glass rod, acting as the
stimulus is increased, the amplitude of the localised, non-propagated generator potential increases and at a certain threshold
produces a propagated action potential in the sensory neurone
587
4p
300P
— 3F
Za 3 200K
=
= =
a.
ee 100+
(le
02 0.4 0.6
of 100 é
Frequency/s' Generator potential/mV
(a)
(b)
Zotterman, 1926 and (b) after Katz,
16.25 Graphs obtained during investigations on frog muscle spindles (a) after Adrian and
1950
sensory neurone. The frequency of these impulses gradu-

16.7 Study the graphs shown in fig 16.25, ally declines and this reduction in response, with time, is
_ obtained from investigations on frog muscle spindle, called adaptation. For example, on entering a room you
and describe the relationship between stimulus, may immediately notice a clock ticking but after a while
generator potential, and frequency of nerve im- become unaware of its presence. The rate and extent of
pulses. adaptation ina receptor cell is related to its function and
there are two types, rapidly and slowly adapting receptors.
16.4.2 Properties of receptors Rapidly adapting receptors (phasic receptors) respond to
changes in stimulus level by producing a high frequency of
The input to the central nervous system from
impulses at the moments when the stimulus is switched ‘on’
receptors provides the organism with all the essential or ‘off’. For example, the Pacinian corpuscle and other
information to enable it to survive. As a rule a single receptors concerned with touch and the detection of
receptor cell cannot monitor the whole range of a given sudden changes act in this way and register the dynamic
type of stimulus, however the organism needs to have aspects of a stimulus.
information concerning the strength or intensity of each Slowly adapting receptors (tonic receptors) register a
stimulus in order to produce the correct response. constant stimulus with a slowly decreasing frequency of
Receptors have two major properties which increase their impulses. For example, crayfish stretch receptors register
effectiveness and efficiency. These are sensitivity and static aspects of a stimulus associated with more or less
discrimination and they are obtained by the following steady conditions.
structural and functional adaptations. Adaptation is thought to be related to a decrease in the
Parallel sensory cells with various permeability of the receptor membrane to ions due to
thresholds sustained stimulation. This progressively reduces the
Some sense organs, such as stretch receptors in muscle, are amplitude and duration of the generator potential and
composed of many sense cells having a range of thresholds. when this falls below the threshold level the sensory
Those having a low threshold are stimulated by weak neurone ceases to fire.
stimuli, and as the strength of the stimuli increases they The advantage of adaptation of sense cells is that it
respond by producing an increasing number of impulses in provides the animal with precise information about
the sensory neurone leaving the sense cell. At a given point changes in the environment. At other times the cells
saturation occurs and the frequency of impulses in the remain quiescent, thus preventing overloading of the
neurone cannot be increased. A further increase in central nervous system with irrelevant and unmanageable
intensity of stimulus will excite sense cells with higher information. This contributes to the overall efficiency and
thresholds, and these too will produce a frequency of economy of the nervous system and enables it to ignore
responses which is proportional to the intensity of the static background information and to concentrate on
applied stimulus. In this way the range of receptors is monitoring aspects of the environment having most
extended (fig 16.26). survival value.
Adaptation Convergence and summation

Most receptors initially respond to a strong constant A high degree of sensitivity is achieved in many sense
stimulus by producing a high frequency of impulses in the organs by an anatomical arrangement of sense cells and
588
increased visual sensitivity produced by this arrangement
of rods is highly adapted to dim-light vision and is well
developed in nocturnal species such as owls, badgers and
foxes. This high degree of sensitivity, however, is linked
with a decrease in visual precision, as may be observed
when attempting to read in poor light. In the human eye
and that of many other diurnal species (active during
daylight) this problem is counteracted by the presence of
cones which, with few exceptions, do not show con-
vergence or summation. What cones lose in sensitivity they
gain in discrimination as described in section 16.5.4.
Spontaneous activity
Some receptors produce nerve impulses in sensory
units)
(arbitrary
stimulus
of
Intensity
neurones in the absence of stimulation. This system is not
as meaningless as it might appear as it has two important
advantages. Firstly it increases the sensitivity of the
stimulus ON receptor by enabling it to make an immediate response to a
Time/s x 1073 stimulus that would normally be too small to produce a
response in the sensory neurone. Any slight change in the
intensity of the stimulus will now produce a change in the
frequency of impulses along the sensory neurone. Secondly
the direction of the change in stimulus can be registered by
this system as an increase or decrease in the frequency of
the response in the sensory neurone. For example,
infra-red receptors in pits in the face of the rattlesnake
which act as direction finders in locating prey and predators
show spontaneous activity and are sensitive and able to
discriminate increases or decreases in temperature of
activity
Electrical
potentials)
(action
neurones
sensory
In
Once
16.26 The frequency of action potentials produced in Feedback control of receptors
sensory neurones leaving three sense cells, A, B and C, each
The threshold of some sense organs can be raised or
having different threshold levels. In the case of B and C the
point at which the receptors become active coincides with the lowered by efferent impulses from the central nervous
saturation point of the sense cell with the lower threshold system. This ‘resets’ the sensitivity of the receptor to
respond to different ranges of stimulus intensities with
equal sensitivity. In many cases the mechanism of control
sensory neurones known as convergence. In such cases, involves feedback from the receptors, which produces
several sense cells are connected to, or converge on, a changes in accessory structures enabling the receptor cells
single sensory neurone. These cells are characteristically to function over a new range. This occurs, for example, in
small, are found in large numbers and are extremely the muscle spindle and the iris of the eye.
sensitive to stimuli. Whilst the effect of a stimulus on a
Lateral inhibition
single one of these cells would not produce a response in
the sensory neurone, the combined effect of the simul- This phenomenon is important in increasing both sensitiv-
taneous stimulation of several cells is cumulative. This ity and discrimination in a receptor. The basic principle
cumulative stimulatory effect produced in the sensory involves the mutual inhibition of response from adjacent
neurone is known as summation and is similar in function to sense cells when stimulated. Investigations carried out on
the summative effect described for synapses in section the individual sense organs (ommatidia) of the compound
16.1.2 and effectors in sections 16.6 and 17.4.5. A good eye of the horseshoe crab, Limulus, revealed that adjacent
example of convergence and summation is provided by rod ommatidia produce a lower response if stimulated simul-
cells of the mammalian retina. Some of these cells are taneously than they do if stimulated independently. This is
capable of detecting a single quantum of light, but the significant in accentuating the differences between two
generator potential produced is inadequate to produce a adjacent areas stimulated by different intensities. Fig 16.27
propagated action potential in a neurone of the optic nerve. shows a model system based on visual perception and
However, several rods (ranging from two or three to demonstrates how light falling on two adjacent receptors is
several hundred) are connected to a single optic nerve fibre enhanced at the bright/dim boundary thereby ‘highlighting’
by a bipolar neurone. Stimulation of at least six rods is the edge. This effect forms the basis of the optical illusion
required to produce an impulse in an optic nerve fibre. The shown in fig 16.28.
589
Lateral inhibition in the human eye increases the
resolving power or visual acuity of the eye. The resolving
power of a system is its ability to distinguish two or more
stimuli of equal intensity as separate stimuli. For example,
the distance at which two black lines of equal density can be
seen as separate lines and not as a single line is a measure of
the eyes’ resolving power. The visual acuity of the human
eye is very high and in part is due to lateral inhibition but
mainly to the anatomical arrangement of cones in the
retina. Light from two sources falling on the same or
immediately adjacent cones is not resolved as coming from
two sources. However, light from two sources falling on
cones separated by only one cone can be perceived as
separate sources. Approximately 95% of the seven million
photoreceptor action potential spikes Fig 16.28 The above diagram illustrates an optical illusion
cells sensory
neurones | | | |
which may be caused by lateral inhibition. The grey spots at
the intersections of the white lines are thought to be produced
4
/
by the image of the white lines inhibiting adjacent receptors
/ which fail to be stimulated. The lack of stimulation from the
/
2S x SS dark areas and the stimulation from the white lines combine to
produce the sensation of grey at the intersections of the white
LIGHT lines
—+( WUE)
\
\ cones in the eye are situated within a 1 mm diameter fovea
\
in the centre of the retina. Here they each have a bipolar
neurone connecting them to their own sensory neurone of
crt: light the optic nerve. It is the absence of convergence and the
(a) no lateral inhibition Pine
close packing of cones in the fovea that is responsible for
the high visual acuity.
= WwW
if
/
4
16.5 Structure and function of receptors
/
eae EZ X ee ET IGE
There is enormous variation in the structure
LIGHT inhibitory and function of receptors in the animal kingdom. In this
neurones
—> z Y eS EL section the structure and function of some mammalian
\
receptors is considered briefly and detailed consideration is
.
given only to the mammalian eye and ear, and the
4
Z
arthropod eye.
(b) with lateral inhibition ON OFF 16.5.1 Mechanoreceptors

light Mechanoreceptors are considered the most
stimulus
primitive type of receptors and may respond to a range of
Fig 16.27 The diagram shows action potentials recorded mechanical stimuli such as pressure, gravity, displacement
from four adjacent ommatidia W, X, Y and Z. X and Y are and vibration.
stimulated directly by a narrow beam of light whilst W and Z
are stimulated weakly by light scattered from the beam Touch and pressure
stimulating X and Y. In (a) with no lateral inhibition W and Z
are excited and action potentials are recorded in their sensory The distinction between touch and pressure is one of
neurones. In (0) with lateral inhibition direct stimulation excites degree, and the detection of these stimuli depends on the
X and Y and inhibitory neurones linking these with W and Z position of the receptors within the skin. Touch receptors
prevent any propagated action potentials being set up in the sensitive to small pressures are found close to the surface of
latter. This mechanism sharpens the contrast, as perceived by the skin and are composed of many fine, free endings of
the cerebral ganglia, as a result of the differential illumination
of adjacent photoreceptors. The same principle applies to the sensory neurones (primary sense cells). These may be
rods and cones of the vertebrate eye (after Lamb, Ingram, situated in the epidermis or attached to hairs in the hair
Johnson and Pitman, 1980) follicle and respond to deformation of the skin and hairs.
590
Touch receptors are localised in certain regions of the body 16.5.2 Thermoreceptors
and account for the increased sensitivity in these regions.
For example, two stimuli may be resolved by the tip of the Two types of sense organs found in the dermis
tongue when 1 mm apart whereas this is only possible at a are claimed to be responsible for detecting the temperature
distance of 60 mm in the middle of the back. of the skin. Organs of Ruffini are thought to respond to
Specialised sense organs known as Meissner’s corpuscles warm temperatures and bulbs of Krause are thought to
are situated immediately beneath the epidermis and respond to low temperatures. However, it is almost certain
respond to touch (fig 16.29). They consist of the single that the many free nerve endings in the skin are the major
convoluted ending of a neurone enclosed within a structures detecting temperature (fig 16.30).
fluid-filled capsule. Pacinian corpuscles are situated in the
dermis, joints, tendons, muscles and mesenteries of the free cornified
nerve
gut, and consist of the ending of a single neurone endings
layer
surrounded by connective tissue lamellae. They respond to

epidermis
pressure (fig 16.24).
All touch and pressure receptors are thought to produce
a generator potential as a result of deformation of the
receptor membrane leading to an increase in the per- dermis
meability of the receptor cell to ions.

Fig 16.30 Free nerve endings in the epidermis and dermis
which detect temperature
Se ___ lead squamous
a epithelial
; cells
stratified
16.5.3 Chemoreceptors
squamous
epithelial The sensations of taste and smell in Man are
cells due to chemical substances exciting specific chemorecep-
tors and play important roles in feeding, avoiding un-
Malpighian favourable environments and other behavioural activities
layer
involving courtship, mating and social organisation.
Meissner’s
corpuscle
Taste
Many of the sensations registered as taste are really
flavours which are detected both by smell and taste
receptors. If a person is blindfolded and pinches his
Fig 16.29 Simplified diagram of a touch receptor, Meissner’s nostrils, preventing chemical stimuli to reach the smell
corpuscle, showing its relationship to the skin receptors, whilst chewing onion and apple, it is unlikely
that the difference between them will be distinguished.
This illustrates the lack of sensitivity of taste receptors.
Muscle spindle Taste or gustation is detected by receptor molecules
Specialised proprioceptors called muscle spindles have situated in microvilli projecting from sensory cells sunk
been found in the muscles of mammals, amphibia, into goblet-shaped organs known as taste-buds found on
crustacea and insects. They respond to changes in tension the top and sides of the tongue (fig 16.31).
in the muscles and act as stretch receptors in all activities There are four ‘taste sensations’ and the uneven
associated with the control of muscular contraction. The distribution of their receptors may be demonstrated by
muscle spindle has three main functions, one static and two applying the following stimulating substances to various
dynamic: areas of the tongue: sweet — saccharin; sour — weak acid;
(1) to provide information to the central nervous system salt — sea salt; bitter — quinine. The distribution of these
on the state and position of muscles and structures receptor areas is shown in fig 16.31b. No correlation has yet
attached to them, a static function; been found between the chemical structure of a substance
(2) to initiate reflex contraction of the muscle and return it and the taste produced. It is assumed, however, that the
to its previous length when stimulated by a load, a production of a generator potential within the receptor cell
dynamic function; depends upon substances either penetrating the receptor
membrane or attaching to specific receptor sites on the
(3) to alter the state of tension in the muscle and reset it to
maintain a new length, a dynamic function. surface of the receptor cell membrane. -Taste receptors
require higher concentrations of stimulating substances
The structure and function of the muscle spindle is
than smell receptors.
described in detail in section 17.5.4.
591
facial nerve
(a) fluid-filled pore
epithelial
cell
supporting
microvilli
cell
glossopharyngeal
sensory cell nerve
basal cell
sweet salt sour bitter
Fig 16.31 (a) Simplified diagram showing the structure of a tastebud. Substances to be tasted dissolve in the fluid Fs
surrounding the microvilli and diffuse to receptor cells embedded in the microvilli. (Redrawn and reproduced with permission,
from Schmidt, 1978). (bo) Diagram showing the symmetrical distribution of taste receptors of the human tongue and their
innervation
Smell Amoore in 1963 proposed a stereochemical theory of

The sensation of smell is more acute than that of taste and is olfaction that related the properties of receptor sites on the
due to airborne odoriferous substances dissolving in the cilia-like projections of the olfactory cells to the shape and
layer of mucus and stimulating an area of olfactory dimension of odour-producing molecules. He proposed
epithelium situated high up in the nasal passages. Each that there are seven primary odours such as putrid,
nostril has an area of 2.5 cm’ containing three types of cells pungent, peppery etc. and that they each have specific
as shown in fig 16.32. receptor sites. About 10 000 odours can be differentiated,
and this is thought to be due to varying degrees of
mucus layer stimulation of these seven types of receptor sites. Water-
vi olfactory knob and lipid-soluble substances produce the strongest sensa-
20 a AA | olfactory tions of smell.
H at — boi ] cilium
EL ED _ Hered microtubule Taste and smell in insects
UY &) JIOGOY
ky ET
mitochondrion The sense of taste in insects is extremely sensitive and has
been studied in detail. Taste organs in insects are known as
supporting taste hairs and they are situated on the antennae, maxillary
cell
and labial palps and feet of insects. They consist of a hollow
structure containing endings of sensory neurones and are
sensory cell sensitive to both chemical substances and displacement.
In the blowfly, Calliphora, there is a taste reflex which
extends the proboscis when the legs are in contact with
acceptable food substances. The response threshold in
nucleus of blowfly to sucrose is as low as 0.8 x 10°* M compared with
sensory cell
0.2 x 107’ Mm in Man.
Despite these remarkably low thresholds for taste
basal cell
receptors the sensitivity of insects to smell is even greater.
The olfactory receptors are usually situated on the
___ basement antennae and have been extensively studied in moths.
WH be} { membrane Female silk moths, Bombyx mori, are capable of attracting
axon of males of the same species at distances of several kilometres
sensory
neurone as a result of releasing a specific chemical substance.
Substances, such as this sex attractant, that act as a form of
Fig. 16.32 Simplified diagram showing the structure and
communication within a species are known as pheromones.
relationships between sensory cell, supporting cell and basal
cell in the olfactory epithelium. Receptor molecules to In one experiment a single silk moth female in a muslin
odoriferous substances are located in the membranes of the cage attracted 40% of a large population of males released
olfactory cilia 4 km away. Several species that utilise pheromones over
ey
great distances have enlarged antennae carrying, in the muscles which control eye movements. Each human
case of the male polyphemus moth, Telea polyphemus, 15 eyeball is about 24 mm in diameter and weighs 6-8 g. Most
x 10* sensory cells. Molecules of sex attractant are of the eye is composed of accessory structures concerned
absorbed on to the antennae as air filters through them. with bringing the visual field to the photoreceptor cells
Contrary to popular belief male moths do not fly up situated in the innermost layer of the eye, the retina. The
concentration gradients to find female moths. Instead they eye is composed of three concentric layers: the sclera
take off into the wind carrying the pheromone and zig-zag (sclerotic coat) and cornea; the choroid, ciliary body, lens
across the aerial trail reversing their direction whenever and iris; and the retina, and is supported by the hydrostatic
they lose the scent. Once in the immediate vicinity of the pressure (25 mmHg) of the aqueous and vitreous humours.
female, her exact location is determined by following the The gross structure of the human eye is shown in fig 16.33
concentration gradient. and brief notes on the function of the various parts are
given below.
16.5.4 Mammalian eye Sclera — external covering of eye; very tough, containing
collagen fibres, protects and maintains shape of eyeball.
The mammalian eye is a sense organ com- Cornea — transparent front part of the sclera; the curved
posed of many sense cells, the rods and cones of the retina, surface acts as a main structure refracting (bending) light
sensory neurones of the optic nerve and a complex array of towards the retina.
accessory structures. This arrangement enables the eye to Conjunctiva — thin transparent layer of cells protecting the
convert light of various wavelengths reflected from objects cornea and continuous with the epithelium of eyelids; the
at varying distances, the visual field, into electrical impulses conjunctiva does not cover the cornea overlying the iris.
which nerves transmit to the brain where an image of Eyelid — protec‘s the cornea from mechanical and chemical
remarkable precision is perceived. damage and the retina from bright light by reflex action.
Light travels as waves of electromagnetic radiation and Choroid — rich in blood vessels supplying the retina and
the wavelengths perceived by the human eye occupy a covered with pigment cells preventing reflection of light
narrow band, the visible spectrum, from 380-760 nm within the eye.
(appendix A 1.7). Light is a form of energy and is emitted Ciliary body — junction of sclera and cornea; contains
and absorbed in discrete packets called quanta or photons. tissue, blood vessels and ciliary muscles.
The wavelengths of the visible spectrum carry sufficient Ciliary muscles — circular sheet of smooth muscle fibres
energy in each quantum of radiation to produce a forming bundles of circular and radial muscles which alter
photochemical response in the sense cells of the eye. the shape of the lens during accommodation.
The camera and the eye work on the same basic Suspensory ligament — attaches the ciliary body to the lens.
principles and these are: Lens — transparent, elastic biconvex structure; provides
(1) controlling the amount of light entering the structure, fine adjustment for focusing light on to the retina and
(2) focusing images of the external world by means of a separates the aqueous and vitreous humours.
lens system; Aqueous humour - clear solution of salts secreted by the
(3) registering the image on asensitive surface; ciliary body, finally draining into the blood through the
(4) processing the ‘captured’ images to produce a pattern canal of Schlemm.
which can be ‘seen’. Iris — circular, muscular diaphragm containing the pigment
giving eye its colour; it separates the aqueous humour
Structure and function of the human eye region into anterior and posterior chambers and controls
The eyes are held in protective bony sockets of the skull the amount of light entering eye.
called orbits by four rectus muscles and two oblique Pupil — opening in iris; all light enters eye through this.
upper eyelid sclera

choroid
conjunctiva
retina
cornea
aqueous humour
pupil vitreous humour
fovea
lens
blind spot
iris
suspensory ligament
ciliary body /WSF-QR— ciliary muscle
Fig 16.33 Structure of the mammalian eye
593
Light from distant object Light from near object
Vitreous humour — clear semi-solid substance supporting
the eyeball.
Retina — contains the photoreceptor cells, rods and cones, focus on focus on
retina
and cell bodies and axons of neurones supplying the optic retina
nerve.
Fovea — most sensitive part of retina, contains cones only; 1 Parallel light rays reach eye 1 Diverging light rays reach eye
2 Cornea refracts (bends) light rays
most light rays are focused here. 2 Cornea refracts (bends) light rays
3 Circular ciliary muscle contracted
3 Circular ciliary muscle relaxed
Optic nerve — bundle of nerve fibres carrying impulses from 4 Suspensory ligament taut 4 Suspensory ligament slack
the retina to the brain. 5 Lens pulled out thin 5 Elastic lens more convex
6 Light focused on retina 6 Light focused on retina
Blind spot — point where the optic nerve leaves eye; there
are no rods or cones here, therefore it is not light-sensitive.
(b)
16.8 List, in order, the structures through
which light passes before striking the retina. _
Accommodation Fig 16.35 Events occurring during accommodation of light

rays from objects at various distances, (a) side views of eye,
Accommodation is the reflex mechanism by which light (b) front views of eye
rays from an object are brought to focus on the retina. It
involves two processes and these will be considered
Table 16.8. Relationship between structures changing the
separately.
shape of the lens and the degree of refraction —
Reflex adjustment of pupil size. In bright
light the circular muscle of the iris diaphragm contracts, the Ciliary muscle — Tension in Radius of Refraction
suspensory curvature
radial muscle relaxes, the pupil becomes smaller and less ligament
light enters the eye, preventing damage to the retina (fig
16.34). In poor light the opposite muscular contractions contracted no tension decreased increased
and relaxations occur. The added advantage of reducing (lens thick)
the pupil size is that it increases the depth of focus of the eye relaxed taut increased decreased
so that any displacement of the sense cells in the retina will (lens thin)
not impair the focus.
Refraction of light rays. Light rays from contraction of the ciliary muscles changes the tension on
distant objects (> 6 m) are parallel when they strike the the suspensory ligaments. This acts on the natural elasticity
eye. Light rays from near objects (< 6 m) are diverging of the lens which causes it to change its shape (radius of
when they strike the eye. In both cases the light rays must curvature) and thus the degree of refraction. As the radius
be refracted or bent to focus on the retina and refraction of curvature of the lens decreases it becomes thicker and
must be greater for light from near objects. The normal
the amount of refraction increases. The complete rela-
eye is able to accommodate light from objects from about tionship between these three structures and refraction is
25 cm to infinity. Refraction occurs when light passes from shown in table 16.8. Fig 16.35 shows the changes occurring
one medium into another with a different refractive index,
in the eye during accommodation to light from distant and
and this occurs at the air—corneal surface and at the lens. near objects.
The degree of refraction at the corneal surface cannot be
The image produced by the eye on the retina is inverted
varied and depends on the angle at which light strikes the
and reversed but the mental image is perceived in the
cornea (which, in turn, depends upon the distance of the
correct position because the brain learns to accept an
object from the cornea). Most refraction occurs here, and
inverted reversed image as normal.
consequently the function of the lens is to produce the final
refraction that brings light to a sharp focus on the retina; Structure of the retina
this is regulated by the ciliary muscles. The state of
The retina develops as an outgrowth of the forebrain called
the optic vesicle. During the embryonic formation of the
Bright light Dim light
circular radial
eye the photoreceptor cells of the retina turn inwards
muscle muscle (invaginate) and lie against the choroid layer. The cells are
< contracts contracts
covered by the cell bodies and axons linking the photo-
pupil pupil
smaller a dilates
receptor cells to the brain as shown in fig 16.36.
radial circular The retina is composed of three layers of cells each
muscle muscle containing a characteristic type of cell. First there is the
relaxes relaxes
photoreceptor layer (outermost layer) containing the
Fig 16.34 The iris/pupil response to variations in light photosensitive cells, the rods and cones, partially embed-
intensity ded in the microvilli of pigment epithelium cells of the
594
sclera outer membrane. The two regions remain in contact by
choroid epithelium cytoplasm andapair of cilia which pass between the two.
showing pigment
——- layer These cilia consist of nine peripheral fibres only, the usual
rod y
vesicles central two being absent.
outer
infoldings of segment Inner segment. This is an actively metabolic
plasma region. It is packed with mitochondria producing energy
membrane ae
pair of cilia constriction for visual processes, and polysomes for the synthesis of
photoreceptor proteins involved in the production of the membranous
layer vesicles and visual pigment. The nucleus is situated in this
inner region.
segment
Synaptic region. Here the cells form syn-
—— apses with bipolar cells. Diffuse bipolar cells may have
synapses with several rods. This is called synaptic con-
synaptic
region vergence, and whilst it lowers visual acuity it increases
ee
ae visual sensitivity. Monosynaptic bipolar cells link one cone
synaptic to one ganglion cell and this gives the cones greater visual
intermediate
terminals layer acuity than rods. Horizontal cells and amacrine cells link
bipolar
neurone
certain numbers of rods together and cones together. This
amacrine cell
allows a certain amount of processing of visual information
internal to occur before it leaves the retina, for instance these cells
Direction
surface layer are involved in lateral inhibition.
of light neurones of
rays Be
woe optic nerve Differences between rods and cones
Rods are more numerous than cones, 120 x 10° as opposed
Fig 16.36 Diagrammatic section through the retina of the
eye showing the ultrastructure of a rod and a cone. to 6 x 10°, and have a different distribution. The thin
Connections between the sensory cells and the neurones of elongate rods (50 xX 3 um) are distributed uniformly
the optic nerve are shown in the inner segment. Light rays throughout the retina except at the fovea, where the
must pass through the ganglion cells and the intermediate conical elongate cones (60 x 1.5 um) have their greatest
layers before reaching the rods and cones concentration (5 x 10* mm~’). Since the cones are tightly
packed together at the fovea this gives them higher visual
choroid. Next is the intermediate layer containing bipolar acuity (section 16.4.2). Rods are much more sensitive to
neurones with synapses connecting the photoreceptor layer light than cones and respond to lower light intensities.
to the cells of the third layer. Horizontal and amacrine cells Rods only contain one visual pigment, and being unable to
found in this layer enable lateral inhibition to occur. The discriminate colour are used principally for night vision.
third layer is the internal surface layer containing ganglion Cones contain three visual pigments and these enable the
cells with dendrites in contact with bipolar neurones and cones to differentiate colours. They are used principally in
axons of the optic nerve. daylight. The rods have a lower visual acuity because they
are less tightly packed together and they undergo synaptic
Structure and function of rods and cones convergence, but this latter point gives them increased
Rods and cones have an essentially similar structure as collective sensitivity required for night vision.
shown in fig 16.36 and their photosensitive pigments are
attached to the outer surfaces of the membranes in the 16.9 Explain why synaptic convergence
outer segment. They have four similar regions whose should increase visual sensitivity.
structure and function are summarised below. 16.10 Explain why objects are seen more
Outer segment. This is the photosensitive clearly at night by not looking directly at them.
region where light energy is converted into a generator
potential. The entire outer segment is composed of The mechanism of photoreception
flattened membranous vesicles containing the photosensi-
Rods contain the photosensitive pigment rhodopsin
tive pigments. Rods contain 600-1 000 of these vesicles
attached to the outer surface of the membranous vesicles.
stacked up like a pile of coins and they are enclosed by an Rhodopsin, or visual purple, is a complex molecule formed
outer membrane. Cones have fewer membranous vesicles by the reversible combination of a lipoprotein, scotopsin,
and they are formed by repeated infoldings of the outer and a small light-absorbing carotenoid molecule retinene
membrane. (retinol). The latter is an aldehyde derivative of vitamin A
Constriction. The outer segment is almost and exists in two isomeric forms according to the light
separated from the inner segment by an infolding of the conditions as shown in fig 16.37.
ooo
no light
Glan light CHa CHe CH, CH,

(lak, (RR outer
segment
O
'
Nh a Nat
,
TN
+
I
'
|
! |
CH, CH,
7\ !
\
| |
|
|
11 cis retinene O all trans retinene
Nat
Nat
Fig 16.37 The action of light changes the structure of inner
retinene from the 11 cis isomer to the all trans isomer segment
When rhodopsin is exposed to light it is known that one

4 ACHVE
photon of light will produce the above isomeric change. transport
Retinene acts as a prosthetic group and is believed to
occupy a certain site on the surface of the scotopsin
molecule where it inhibits reactive groups in this molecule
involved in initiating electrical activity in the rods. The
exact mechanisms of photoreception are not yet known but
it is thought that it involves two processes: first the isomeric Fig 16.38 Diagram of a rod showing the proposed changes
conversion of 11 cis retinene to all trans retinene by the in sodium permeability of the outer segment produced in the
presence of light. The negative charges © on the right side of
action of light, and second the decomposition of rhodopsin the rod indicate the normal resting potential whereas those on
via a series of intermediate molecules into retinene and the left indicate the hyperpolarisation
scotopsin, a process known as bleaching.
bleaching
Table 16.9. Colours of the visible spectrum and
rhodopsin ————> retinene t scotopsin approximate ranges of their wavelength
Rhodopsin is reformed immediately in the absence of Colour

ae ee
Wavelength/nm
Eee
ON
further light stimulation. All trans retinene is first con- above 620
red
verted into 11 cis retinene in the presence of the enzyme orange 590-620
retinene isomerase, and is then recombined with scotopsin. yellow 570-590
This process is called ‘dark adaptation’ and in total green 500-570
darkness it takes 30 min for all rods to adapt and the eyes to blue 440-500
violet below 440
achieve maximum sensitivity. During the second process,
however, the permeability of the outer segment membrane
to sodium ions decreases whilst the inner segment
continues to pump out sodium ions, thus creating increased have shown absorb light of different wavelengths. There
negativity within the rod (fig 16.38). This situation gives are red, green and blue cones.
rise to hyperpolarisation of the rod. This situation is exactly Colour vision is explained in terms of the trichromacy
opposite to the effect normally found in sensory receptors theory, which states that different colours and shades are
where the stimulus produces a depolarisation and not a produced by the degree of stimulation of each type of cone
hyperpolarisation. The hyperpolarisation reduces the rate produced by the light reflected from an object. For
of release of excitatory transmitter substance from the rod example, equal stimulation of all cones produces the colour
which is released maximally during darkness. The bipolar sensation of white. The initial discrimination of colour
neurone linked by synapses to the rod cell also responds by occurs in the retina but the final colour perceived involves
producing a hyperpolarisation, but the ganglion cells of the the integrative properties of the brain. The mixing effect of
optic nerve supplied by the bipolar neurone respond to this pigments forms the basis of colour television, photography
by producing a propagated action potential. and painting.
Colour vision Colour-blindness. The complete absence of
The human eye absorbs light from all wavelengths of the a particular cone or a shortage of one type can lead to
visible spectrum and perceives these as six colours broadly various forms of colour-blindness or degrees of ‘colour-
associated with particular wavelengths as shown in table weakness’. For example, a person lacking red or green
16.9. There are three types of cones each possessing a cones is ‘red-green colour-blind’, whereas a person with a
different pigment which electrophysiological investigations reduced number of either cones will have difficulty in
596
distinguishing a range of red-green shades. Colour- right eye left eye
blindness, or its extent, is determined using test charts
,
such as the Ishihara test charts, composed of aseries of dots
of several colours. Some charts bear a number which
a optic nerve
person with normal colour vision can perceive, whilst the
colour-blind sufferer sees a different number or no number
at all. optic chiasma
Colour-blindness is a sex-linked recessive characteristic
resulting in the absence of appropriate colour genes in the
X chromosome. About 2% of men are red colour-blind, lateral
geniculate
6% are green colour-blind, but only 0.4% of women show nucleus
any sign of colour-blindness.
visual cortex
16.11 A person places a green filter over
one eye and ared filter over the other eye and looks Fig 16.39 Diagram of the human visual pathway as seen
at an object. Using the data given in table 16.9, from the underside of the brain
describe and explain the apparent colour of the
- object. involving perhaps only a few rods and cones, is represented
and it is here that the visual input is interpreted and we
‘see’. However, what we see has meaning only after
Binocular vision and stereoscopic reference to other regions of the cortex and the temporal
vision lobes, where previous visual information is stored and used
Binocular vision occurs when the visual fields of both eyes in the analysis and identification of the present visual input
overlap so that the fovea of both eyes are focused on the (section 16.2.4). In Man, axons from the left side of the
same object. It has several advantages over monocular retina of both eyes pass to the left visual cortex and axons
vision and these include alarger visual field, damage to one from the right side of each retina pass to the right visual
eye being compensated for by the other, for example it cortex. The point where the axons from those regions of the
cancels the effect of the blind spot, and it provides the basis retina closest to the nose cross is called the optic chiasma
of stereoscopic vision. Stereoscopic vision depends upon and is included in the visual pathway shown in fig 16.39.
the eyes simultaneously producing slightly different retinal About 20% of the optic neurones do not pass to the visual
images which the brain resolves as one image. The more cortex but enter the midbrain, where they are involved in
frontally the eyes are situated the greater the stereoscopic reflex control of pupil size and eye movements.
visual field. For example, Man hasa total visual field of
180° and a stereoscopic visual field of 140°. The horse has 16.5.5 Arthropod eye
laterally placed eyes with a limited forward stereoscopic Arthropods have two types of eye: the simple
visual field which it uses for viewing distant objects. For eye (ocellus), which is found in some larval insects, and the
nearer objects the horse turns its head and uses monocular more characteristic compound eye. The simple eye consists
vision to examine detail. Frontally placed eyes and of a single lens covering a few light-sensitive cells and is
centrally situated foveas, producing good visual acuity, are able to discriminate light and dark but unable to produce
essential for good stereoscopic vision which provides an images. These eyes are often found in conjunction with the
increased appreciation of size and perception of the depth larger, and functionally more important, compound eyes.
and distance of objects. Stereoscopic vision is found mainly Each compound eye is composed of thousands of
in predatory animals where it is vital when capturing prey separate structures called ommatidia; for example there
by pouncing or swooping, as shown by members of the cat are 30 000 in the dragonfly. Each ommatidium is composed
family, hawks and eagles. Animals which are hunted have of two refractive bodies, a biconvex lens formed from
laterally placed eyes giving wide visual fields but restricted cuticle and a crystalline cone secreted by vitrellar cells.
stereoscopic vision, for instance the rabbit has a total These focus light on to a group of photosensitive retinulae
binocular visual field of 360° and frontal stereoscopic vision cells, whose inner membranes bear microvilli which fuse
of 20°. The resolution of the two retinal images produced in together to form a structure called a rhabdome. The visual
stereoscopic vision occurs in two areas of the brain called pigment is located within the microvilli of the rhabdome
the visual cortex. where the nerve impulses are set up and pass via the optic
nerve to the cerebral ganglia. Each ommatidium is
Visual pathways and the visual cortex surrounded and isolated from the next by elongate pigment
Nerve impulses generated in the retina are carried by the cells (fig 16.40).
million or so neurones of the optic nerve to the visual Since each ommatidium receives light from a very small
cortex, which is situated in the occipital lobe at the rear area, the image formed by the eye as a whole is made up of
base of the brain. Here each minute region of the retina, a series of separate overlapping points and is known as a
397
= aes EMS semicircular
(a) Cee
canal
___crystalline core stapes
ampulla
mee Sy itrellarcell incus
____ proximal pigment cell +e a utr iculus

malleus
___distal pigment cell ear-drum oval

ee sacculus
(b) window
' /median
igment cell _ retinula cell canal
ae
retinula cell
aAR
_____rhabdome direction of
sound waves middle ear ;
tympanic
vestibular\ canal
window canal
external auditory
meatus (ear tube)
perilymph
eustachian tube (peri, around)
| re) ____neurones of optic nerve
rhabdome
m cochlea (normally
Fig 16.40 Diagrams of (a) an LS of a single ommatidiu (all structures are endolymph
coiled)
encased in bone) (endo, inside)
and (b) TS of an ommatidium
Fig 16.41 Diagram showing the major structures in the
mammal ian ear involved in hearing and balance, (not to
scale)
mosaic image. This type of eye lacks the resolution of the connection between the air-filled middle ear and the
mammalian eye but its overlapping images make it very pharynx, the eustachian tube. Finally there is the inner ear
sensitive to movements of objects. which consists of a complex system of canals and cavities
within the skull bone containing a fluid called perilymph.
Within these canals are membranous sacs filled with
16.5.6 Mammalian ear endolymph and sensory receptors. Auditory receptors are
The mammalian ear is a sense organ contain- found in the cochlea and balance receptors are found in the
ing mechanoreceptors sensitive to gravity, displacement utricle and saccule and the ampullae of the semicircular
and sound. Movements and positions of the head relative canals. The perilymph is enclosed by the membranes of the
to gravity are detected by the vestibular apparatus oval window and round window.
composed of the semicircular canals, the utricle and the
The nature of sound
saccule. All other structures of the ear are involved in
receiving, amplifying and transducing sound energy into Sound is produced by the vibration of particles within a
electrical impulses and producing the sensation of hearing medium. It travels as waves consisting of alternating
in the auditory regions of the brain. regions of high and low pressure and will pass through
liquids, solids and gases. The distance between two
Structure and function of the ear identical points on adjacent waves is the wavelength and.
The ear consists of three sections specialising in different this determines the frequency of vibrations or pitch
functions (fig 16.41). The outer ear consists of the pinna, (whether it sounds high or low). The human ear is sensitive
strengthened by elastic cartilage, which focuses and to wavelengths between 40 and 20 000 Hz (cycles per
collects sound waves into the ear tube (external auditory second). The audible range of dogs reaches 40 000 Hz, and
meatus); the sound waves cause the tympanic membrane that of bats 100 000 Hz. Human speech frequencies vary
(eardrum) to vibrate. In the middle ear, vibrations of the between 500 and 3000 Hz, and sensitivity to high
tympanic membrane are transmitted across to the membra- frequencies decreases with age.
nous oval window by movement of the three ear ossicles, Tone depends upon the number of different frequencies
the malleus, incus and stapes (hammer, anvil and stirrup). making up the sound. For example, a violin and trumpet
A lever system between these three bones and the relative playing the same note, say middle C, produce the same
areas of contact of the malleus with the tympanic fundamental frequency of 256 Hz but sound different. This
membrane (60 mm’) and the stapes with the oval window is due to overtones or harmonics produced by the
(3.2 mm’) amplifies the movement of the tympanic instrument which give it its distinctive quality or timbre.
membrane 22 times. Damage to the tympanic membrane, The same principle applies to the human voice and gives it
due to atmospheric pressure changes, is prevented by a its characteristic sound.
598
Pitch and intensity discrimination
vestibular canal The ability to distinguish the pitch of a sound depends upon
the frequency of the vibration producing movement of the
Reissner’s membrane
basilar membrane and stimulating sensory cells in a specific
region of the organ of Corti. These cells supply a particular
median canal region of the auditory cortex of the brain where the
sensation of pitch is perceived. The basilar membrane
becomes broader and more flexible as it passes from the
tectorial
membrane base of the cochlea to its apex and its sensitivity to vibration
See Dasilan organ of changes along its length so that only low-frequency sounds
membrane Corti
can pass to the apex. High-frequency (pitch) sounds
sensory
hair cell stimulate the basilar membrane at the base of the cochlea
auditory and low frequency sounds stimulate it at the apical end. A
neurone pure sound, consisting of a single frequency, will only
tympanic canal stimulate one small area of the basilar membrane whereas
most sounds, containing several frequencies, simul-
Fig 16.42 Diagram of a TS cochlea showing the organ of taneously stimulate many regions of the basilar membrane.
Corti The auditory cortex integrates the stimuli from these
various regions of the basilar membrane and a ‘single’
blended sound is perceived.
The intensity (loudness) of a sound depends upon the The intensity or loudness of the sound depends upon
amplitude of the sound waves produced at the source and is each region of the basilar membrane containing a range of
a measure of the energy they contain. sensory cells responding to different thresholds of vibra-
Cochlea and hearing tion. For example, a quiet sound at a given frequency may
only stimulate a few sensory cells, whereas a louder sound
The cochlea is a spiral canal 35 mm long and subdivided
at the same frequency would stimulate several other
longitudinally by a membranous triangle into three regions
sensory cells having higher thresholds of vibration. This is
as shown in fig 16.42. an example of spatial summation.
Both the vestibular and tympanic canals contain
perilymph, and the two canals are connected at the extreme Balance
end of the cochlea via a small hole, the helicotrema. The Maintaining balance at rest and during movement of the
median canal contains endolymph. The basilar membrane body relies upon the brain receiving a continual input of
separates the median and tympanic canals and supports sensory information concerning the position of various
sensory hair cells that can be brought into contact with the parts of the body. Information from proprioceptors in
tectorial membrane above. This unit, consisting of basilar joints and muscles indicates the positions and state of the
membrane, sensory cells and tectorial membrane, is called limbs, but vital information related to position and
the organ of Corti and is the region where transduction of movement of the head is provided by the vestibular
sound waves into electrical impulses occurs. apparatus of the ear, the utricle, saccule and semicircular
Sound waves transmitted from the ear tube to the oval canals.
window produce vibrations in the perilymph of the The basic sensory receptors within these structures
vestibular canal and these are transmitted via Reissner’s consist of cells which have hair-like extensions, hair cells,
membrane to the endolymph in the median canal. From attached to dense structures supported in the endolymph.
here they are transferred to the basilar membrane and the Movement of the head results in deflection of the hairs and
perilymph in the tympanic canal, and are finally dissipated the production of a generator potential in the hair cells.
into the air of the middle ear as vibration of the round Regions of the walls of the utricle and saccule called
window. maculae contain receptor cells which have their hair-like
The precise mechanism of transduction of pressure processes embedded in a gelatinous mass containing gran-
waves into nerve impulses is not known but is believed to ules of calcium carbonate called an otoconium. Otoconia
involve relative movement of the basilar and tectorial respond to the pull of gravity acting at right-angles to the
membranes. Vibrations of the basilar membrane, induced Earth’s surface and are mainly responsible for detecting the
by pressure waves, push the sensory hairs against the direction of movement of the head with respect to gravity.
tectorial membrane and force the two membranes to slide The utricle responds to vertical movements of the head
past each other. The distortion produced in the sensory and the otoconia produce maximum stimulation when
hairs due to the shearing forces causes a depolarisation of pulling the receptor hairs downwards, such as when the
the sensory cells, the production of generator potentials body is upside down.
and the initiation of action potentials in the axons of the The saccule responds to lateral (sideways) movement of
auditory nerve. the head. The hair cells of the saccule are horizontal when
599
Penile.)
otoconia and (b) the otoconial layer

Fig 16.43 Scanning electron micrographs of the internal structure of the ear. (a) the
with a portion removed to show the sensory cells of the macula beneath it
ampulla
cupula
n
oO
&
endolymph
a2
Aao og
° ec So
sensory hair
e238
38
S56
sensory cell
Les
Position of
head (as vestibular neurone
shown below) utiricle
——_—_——
sensory cell Fig 16.45 Diagram showing a TS through the ampulla of a
utricle hair processes semicircular canal
otolith
gradual
change in
position from
left left to right 8, The three semicircular canals are arranged in three
saccule o planes at right-angles to each other and detect the direction
&
s
& and rate of change of position of the head. At the base of
each canal is a swelling, the ampulla, containing a conical
gelatinous structure, the cupula. This encloses the hair-like
Fig 16.44 Graph showing the effect of head position on the projections of the receptor cells. The cupula fully extends
activity of receptor cells of the utricle and saccule across the ampulla (fig 16.45). Rotational movement of the
head, semicircular canals and cupula is resisted by the
inertia of the endolymph which remains stationary. This
the head is upright. Tilting of the head to the left produces a produces a relative displacement of the cupula which is
differential response from left and right saccules. The left bent in the opposite direction to the head movement. The
receives increased stimulation as the otoconia pull down- receptor cells respond by producing generator potentials
wards on the hairs whereas decreased stimulation occurs on leading to propagated action potentials in the vestibular
the right. These displacements produce impulses passing to neurones. The direction and rate of displacement are both
the cerebellum where the orientation of the head is detected by the receptor cells. Linear acceleration is
perceived. detected by both the maculae and cupulae.
600
16.6 —Effectors - the endocrine system substances produced by one part of the body; these then
enter the bloodstream and pass toa distant organ, tissue or
An effector is a differentiated structure, such
group of cells where they exert a specific regulatory effect.
as a cell, tissue, organ or organ system, performing a
The major endocrine glands of the body and their
specific reaction relative to the environment in response to
hormones and effects are summarised in table 16.10.
a stimulus from the nervous system. The most widespread
and important effectors are those involved in movement Methods of studying glands and
and secretion, hormones
Organisms responding to external stimuli by some form Most of the original information concerning the function of
of movement or locomotion may utilise amoeboid, ciliary, glands and their secretions involved studying the change in
flagellate or muscular movements, according to their body activity associated with overactivity or underactivity
structure and mode of life, in order to make the response. of the gland during disease. In some cases the experimental
Details of these mechanisms can be found in chapter 17. In removal of the gland was also used to determine its
the case of multicellular organisms all types of movement function. As techniques for the isolation, purification,
may be involved in response to internal stimuli, and analysis and synthesis of hormones developed, biologists
examples include the amoeboid movement of white blood were able to counteract the symptoms of removing the
cells through capillary walls to a site of infection in the gland by injecting extracts of the gland or synthetic
tissues, the increased movement of cilia in the trachea in preparations of the hormone. These techniques clearly
response to infection in the lungs, the activation of sperm established the role of many glands and their secretions but
flagella in response to secretions of the vagina, and reflex they could not be applied to all glands, for example the
contractions of the peristaltic muscles of the oesophagus in liver and kidney. The removal of these glands would
response to a food bolus. produce undesirable conditions other than those connected
Other stimuli, which may be external or internal, bring with their endocrine role.
about a response in the organism involving secretory Radioimmunoassay (RIA) is a technique which is widely
activity of either an exocrine gland or an endocrine gland used to measure the concentration of hormones, drugs,
(section 8.3.3). For example, exocrine responses may be enzymes, viruses, bacterial and tumour antigens and other
stimulated by particles of dirt in the eye causing secretion of organic substances of biological interest in blood, urine or
tears from the tear glands, acidic food in the duodenum other media. The principle of RIA is simple. The
causing the release of pancreatic juice and damage to a components of the assay consist of a specific anti-serum to
joint causing an increased production of synovial fluid. the substance (hormone) under investigation, a radioac-
Endocrine glands too respond to a variety of stimuli, for tively labelled form of the substance and the unlabelled
example the release of insulin by the cells of the islets of substance. The unlabelled substance competes for a limited
Langerhans in response to rising blood glucose levels, the number of antibody-binding sites with the labelled sub-
release of human chorionic gonadotrophin at the beginning stance. The amount of inhibition of binding of the labelled
of pregnancy by the uterus wall following implantation by a substance is proportional to the amount of unlabelled
blastocyst and the increased release of thyroxine following substance present. Measurement of this inhibition is
prolonged exposure to low temperatures. possible by separating (usually precipitating) the antibody-
bound material from the unbound, and the technique is
16.6.1 The endocrine system made quantitative by comparing the results with a set of
The endocrine and nervous systems function tubes containing a known concentration of the substance
in a coordinated fashion to maintain a homeostatic state under investigation. This is shown diagrammatically as
within the body. The nervous system transmits nerve follows.
impulses whereas the endocrine system utilises the blood as Ab + XO A ae eA aeNCO ea
specific unlabelled labelled unlabelled labelled
a transport medium. Despite obvious differences in the antibody antigen antigen (antibody—antigen complexes)
mechanism of transfer of information, both systems share a
common feature in the release of chemical substances as a Current trends in this field are to replace radioactive
means of communication between cells. It is believed that labels with enzyme-fluorescent or chemiluminescent labels
these two systems originated and developed side by side as and thus remove the hazards associated with handling
the needs of intercellular communication became more radioactive sources.
complex due to the increase in the size and complexity of
Mechanisms of hormone action
organisms. In both cases the principal role of the systems is
the integration, coordination and control of many of the All vertebrate hormones belong to one of four chemical
major physiological activities of organisms. _ groups: derivatives of amines, such as tyrosine; peptides
A gland is a structure secreting a specific chemical and proteins; steroids; and fatty acids, as summarised in
substance, and there are two types of glands in the body; table 16.11. The mechanisms controlling the release of
exocrine and endocrine glands. Endocrine glands secrete hormones by glands are as follows.
hormones (hormon, to urge on) which are specific chemical (a) The presence of a specific metabolite in the blood. For
601
%
Table 16.10. Summary of the major human endocrines, their functions and the control of their secretion
eee OOD
Gland Hormone Functions Secretion control mechanism

Oe ie ee ee ee
Hypothalamus Releasing and inhibiting hormones Control of specific anterior pituitary Feedback mechanisms
and factors, seven identified, hormones involving metabolite and
possible number unknown (see hormone levels
table 16.2)
Posterior pituitary hormones
produced here
Posterior No hormones synthesised here, Feedback mechanisms
pituitary gland stores and secretes the following: involving hormones and
Oxytocin Ejection of milk from mammary gland, nervous system
contraction of uterus during birth
Antidiuretic hormone (ADH) Reduction of urine secretion by kidney Blood osmotic pressure
(vasopressin)
Anterior Follicle stimulating hormone (FSH) In male, stimulates spermatogenesis Plasma oestrogen and testos-
pituitary gland In female, growth of ovarian follicles terone via hypothalamus
Luteinising hormone (LH) In male, testosterone secretion Plasma testosterone via
hypothalamus
In female, secretion of oestrogen and Plasma oestrogen level via
progesterone, ovulation and maintenance hypothalamus
of corpus luteum
Prolactin Stimulates milk production and secretion Hypothalamic hormones
Thyroid stimulating hormone Synthesis and secretion of thyroid Plasma T, and T, levels via
(TSH) hormones, growth of thyroid glands hypothalamus
Adrenocorticotrophic hormone Synthesis and secretion of adrenal cortex Plasma ACTH via
(ACTH or corticotrophin) hormones, growth of gland hypothalamus
Growth hormone (GH) Protein synthesis, growth, especially of Hypothalamic hormones
bones of limbs
Parathyroid Parathormone Increases blood calcium level lasma Ca’ level, and
gland Decreases blood phosphate level plasma PO,’ level
Thyroid gland Triiodothyronine (T;) and Regulation of basal metabolic rate, TSH
thyroxine (T,) growth and development
Calcitonin Decreases blood calcium level Plasma Ca’ level
Adrenal cortex Glucocorticoids (cortisol) Protein breakdown, glucose/glycogen ACTH
synthesis, adaptation to stress, anti-
inflammatory/allergy effects
Mineralocorticoids (aldosterone) Na’ retention in kidney, Na’ and K* Plasma Na’ and K levels
ratios in extracellular and intracellular and low blood pressure
fluids, raises blood pressure
Adrenal Adrenaline (epinephrine) Increases rate and force of heartbeat, Sympathetic nervous system
medulla constriction of skin and visceral capillaries
Dilation of arterioles of heart and skeletal
muscles, raises blood glucose level
Noradrenaline (norepinephrine) General constriction of small arteries, Nervous system
elevation of blood pressure
Islets of Insulin (beta cells) — Decreases blood glucose level, increases Plasma glucose and amino
Langerhans glucose and amino acid uptake and acid levels
utilisation by cells
Glucagon (alpha cells) Increases blood glucose level, breakdown Plasma glucose level
of glycogen to glucose in liver
Stomach Gastrin Secretion of gastric juices Food in stomach
Duodenum Secretin Secretion of pancreatic juice Acidic food in duodenum
Inhibits gastric secretion
Cholecystokinin (pancreozymin) Emptying of gall bladder and liberation of Fatty acids and amino acids
pancreatic juice into duodenum in duodenum
Kidney Renin Conversion of angiotensinogen into Plasma Na’ level, decreased
angiotensin blood pressure
Ovarian follicle Oestrogens (17(-oestradiol) Female secondary sex characteristics, FSH and LH
oestrous cycle
Progesterone Gestation, inhibition of ovulation LH
Corpus luteum Progesterone and oestrogen Uterine growth and development LH
Progesterone and oestrogen Foetal development Developing foetus
Placenta Chorionic gonadotrophin Maintenance of corpus luteum Developing foetus
Human placental lactogen Stimulates mammary growth Developing foetus
Testis Testosterone Male secondary sexual characteristics LH and FSH
SSS
602
Table 16.11. Summary table showing chemical nature of the major hormones of the
body
sss
Chemical group Hormone Major source

, catecholamines Adrenaline Sympathetic nervous system
Amines Noradrenaline Adrenal medulla
tyrosine Thyroxine Thyroid gland
Triiodothyronine Thyroid gland
Hypothalamic releasing Hypothalamus
and inhibiting
hormones and factors
Follicle stimulating
hormone
Luteinising hormone Anterior pituitary gland
Prolactin
Thyroid stimulating
hormone
Adrenocorticotrophic
hormone
Peptides and proteins Growth hormone
Oxytocin Posterior pituitary gland
Vasopressin (ADH)
Parathormone Parathyroid gland
Calcitonin Thyroid gland
Insulin Islets of Langerhans (pancreas)
Glucagon
Gastrin Stomach mucosa
Secretin Duodenal mucosa
Steroids Testosterone Testis
Oestrogens Ovary and placenta
Progesterone
Corticosteroids Adrenal cortex
Fatty acids Prostaglandins Many tissues
example, excess glucose in the blood causes the release the cascade effect, is significant because it enables the effect
of insulin from the pancreas which lowers the blood of the release of a small amount of initial hormone to
glucose level. become amplified as it passes along the endocrine pathway.
(b) The presence of another hormone in the blood. For
example, many of the hormones released from the
anterior pituitary gland are ‘stimulating’ hormones ereren
hypothalamus
ays Amount of
sulistares
which cause the release of other hormones from other
glands in the body. adrenocorticotrophic
releasing factor 0.1 pg
(c) Stimulation by neurones from the autonomic nervous (CRF)
system. For example, adrenaline and noradrenaline
are released from the cells of the adrenal medulla by anterior pituitary
the arrival of nerve impulses in situations of anxiety,
adrenocorticotrophic 1.0
stress and danger. hormone (ACTH) Pe
In the first two cases the timing of hormone release and
the amount of hormone released are regulated by feedback adrenal cortex
control. Positive and negative feedback mechanisms both
operate in endocrine control but positive feedback, since it cortisol 40 jig
tends to increase instability, only operates as part of a
larger control system. For example, the release of
luteinising hormone under the stimulus of oestrogen is an LIVER
example of positive feedback but its continued release is

prevented by the later release of progesterone, glucose glycogen 5 600 pg
Hormones which are released by the presence of another
Fig 16.46 An example of the ‘cascade’ effect in the control
circulating hormone are usually under the control of the of the conversion of glucose to glycogen as a result of the
hypothalamus and pituitary glands and the final metabolic release of adrenocorticotrophic releasing factor. The total
or growth effect may involve the secretion of three separate amplification in this example is 56 000 times (data from
hormones as shown in fig 16.46. This mechanism, known as Bradley, 1976)
603
(4) Steroid hormones, and the insect moulting hormone
ecdysone, pass through the cell membrane and bind to
BLOODSTREAM glucose
a receptor in the cytoplasm. The complex formed

cell membrane passes to the cell nucleus where the hormones exert a
direct effect upon the chromosomes by activating
genes and stimulating transcription (messenger RNA
formation).
In many cases of hormone action hormones appear to
exert their effects by influencing enzymes associated with
membranes or genetic systems.
ATP——__———> cyclic AMP
(several
steps) 16.6.2 The hypothalamus and pituitary gland
|
Intercellular communication is achieved, as
|
|
stated earlier, by the activity of nervous and endocrine
inactive nee.) a active
phosphorylase b phosphorylase a systems acting independently or together. The major
|
centres in the body for the coordination and integration of
the two systems of control are the hypothalamus and
glycogen ————————_ glucose pituitary gland. The hypothalamus plays a dominant role in
collecting information from other regions of the brain and
Fig 16.47 Simplified diagram showing how adrenaline from blood vessels passing through it. This information
causes the release of glucose from aliver cell. The release of passes to the pituitary gland which, by its secretions,
membrane bound adenyl cyclase produces cyclic AMP which
activates enzyme systems leading to the breakdown of
directly or indirecily regulates the activity of all other
glycogen to glucose which diffuses out the cells into the blood glands.
stream
The hypothalamus
The hypothalamus is situated at the base of the forebrain
Hormones are specific chemical substances and only immediately beneath the thalamus and above the pituitary
exert their effects on target cells which possess the specific gland. It is composed of several distinct regions called
protein or lipoprotein receptors that interact with the nuclei made up of collections of cell bodies whose axons
hormone. Non-target cells lack these receptors, therefore terminate on blood capillaries in the median eminence and
there is no response to the circulating hormone. Many posterior pituitary as shown in fig 16.48. Many physiologi-
hormones have specific regions of their molecular structure cal activities such as hunger, thirst, sleep and temperature
for attachment to the receptor, and once attached the regulation are regulated by nervous control exercised
hormone may exert its effect at one of the following levels through nerve impulses passing from the hypothalamus
in the cell: (1) the cell membrane, (2) enzymes located in along autonomic neurones. The hypothalamic control of
the cell membrane, (3) cellular organelles, and (4) genes.
An example of each of these levels follows. primary plexus neurosecretory
(releasing and cells
(1) Insulin exerts one of its effects by increasing the uptake inhibiting factors
of glucose into cells by binding with a receptor site and secreted here)
altering the permeability of the membrane to glucose.
(2) Adrenaline and many peptide hormones bind to
receptor sites on the cell membrane and cause the
release of a ‘second messenger’ which initiates a
sequence of enzyme mechanisms which produce the hypothalamo-hypophysial
appropriate hormonal response. In many cases this portal system
‘second messenger’ is the nucleotide 3,5—adenosine hypothalamo-
monophosphate (cyclic AMP) which is formed by the secondary plexus
hypophysial
fibre tract
action of the enzyme adenyl cyclase on ATP following (hormones released
the enzyme’s release from the receptor site. A here)
neurohaemal region
simplified representation of this mechanism is seen in
fig 16.47.
(3) One of the effects of thyroxine is seen at the level of the anterior pituitary posterior pituitary
mitochondrion where it influences the enzymes of the branches of
electron carrier system involved with the formation of pituitary vein
ATP (section 11.5.4). Much of the energy passing Fig 16.48 Diagram showing the relationship between
along the electron transport chain in those circum- neurosecretory cells and blood vessels in the hypothalamus
stances is lost as heat. and pituitary gland
604
endocrine secretion, however, lies in the ability of the trophic hormones which are produced and stored by the
hypothalamus to monitor metabolite and hormone levels in anterior pituitary. These six hormones pass into the blood
the blood. Information generated this way, together with vessels leaving the pituitary and exert their effects on
information from almost all parts of the brain, then passes specific target organs throughout the body, as shown in
to the pituitary gland either by the release of ‘hormones’ table 16.12.
into the blood vessels of the median eminence, which The release of the first two hormones is under the dual
supply the pituitary, or by neurones. The information inhibitory and stimulatory control of the hypothalamus,
relayed by neurones passes through specialised neurones whereas the release of the other four is regulated by
called neurosecretory cells. negative feedback of hormones from the target glands
All nerve cells release a chemical substance, a transmit- acting on receptors in the hypothalamus and anterior
ter substance, at their terminal synapse, but neurosecre- pituitary. Pituitary hormones stimulate the release of
tory cells are nerve cells that have developed the secretory target gland hormones and as the levels of these rise they
capacity to a high level. Chemical substances are produced inhibit the secretion of hypothalamic and pituitary hor-
in the cell bodies of these cells and packaged into granules mones. When the circulatory level of these target hor-
or droplets before being transported down the axon by mones falls below a certain level the hypothalamic and
axoplasmic streaming. At the terminal end of the neurone pituitary inhibition ceases allowing the increased secretion
these cells synapse on to capillaries into which they release from these glands. This control mechanism is described in
their secretion when stimulated by nerve impulses passing sections 18.4.4 and 19.6.
down the axon. Posterior pituitary or neurohypophysis (neuro,
The pituitary gland nerve). ‘This is derived from a downgrowth of the floor of
The pituitary gland or hypophysis (hypo, under; phyein, to the brain. It does not synthesise any hormones but stores
and releases two hormones, antidiuretic hormone (ADH or
grow) is a small red-grey gland weighing about 0.5 g and
vasopressin) and oxytocin. Antidiuretic hormone is re-
connected to the brain by the pituitary stalk (infundibu-
leased in response to a fall in the water content of plasma
lum). It has a dual origin and retains features of those
and leads to an increase in the permeability to water of the
origins in its functions. The two lobes of the pituitary gland
distal and collecting tubules of the nephron so that water is
are the anterior pituitary and the posterior pituitary.
retained in the blood plasma. A reduced volume of
Anterior pituitary or adenohypophysis (aden, hypertonic urine is excreted (section 19.6). Oxytocin
gland). This is derived from an upgrowth of the glandular causes contraction of the uterus during birth and the
roof of the mouth. It is connected to the hypothalamus by ejection of milk from the nipple (section 20.3.8).
blood vessels of the hypothalamo-hypophysial portal ADH and oxytocin are produced by neurosecretory cell
system which has a capillary bed, the primary plexus, in the bodies lying in the nuclei of the hypothalamus and pass
median eminence of the hypothalamus and a secondary down the nerve fibres attached to carrier protein molecules
plexus in the anterior pituitary. called neurophysins. These neurosecretory cells are much
Nerve terminals from specialised neurosecretory cells more specialised than those connected with the secretion of
release two groups of chemical substances, known as releasing factors and they form structures in the posterior
‘releasing factors’ and ‘inhibiting factors’, into the blood pituitary known as neurohaemal organs. These are struc-
capillaries of the primary plexus and these pass to the tures consisting of a swollen synapse attached to a capillary
secondary plexus where they cause the release of six and surrounded by connective tissue (fig 16.49).
Table 16.12. Summary table showing the main hypothalamic hormones, the anterior pituitary hormones influenced
by them and their target organs
SS ——<$——j .—00o’™’—_—ar—|-|.
ee
Hypothalamic hormone Anterior pituitary hormone Site of action

and response
a
eee SS Se eS SS
Growth hormone releasing factor (GHRF)
Growth hormone release-inhibiting hormone (GHRIH)* Growth hormone (GH) Most tissues
(somatostatin)
Prolactin releasing factor (PRF) Prolactin (luteotrophin) (LTH)
Ovary and mammary gland
Prolactin inhibiting factor (PIF) Inhibition of prolactin secretion
Luteinising hormone releasing hormone (LHRH)* {Follicle stimulating hormone (FSH) Ovary and testis
Luteinising hormone (LH)
Thyroid stimulating hormone (TSH) Thyroid gland
Thyrotrophin releasing hormone (TRH)*
Adrenocorticotrophic hormone Adrenal cortex
Adrenocorticotrophin releasing factor (CRF)
(ACTH)
SSS
eS
identi
i d identity
i an establishe i hormone isis a able to stimulate the release
inisi g hormone releasing
are known as A ‘hormones’.: Luteinisin
*Releasing factors with
of both FSH and LH.
605
afferent boys result in precocious sexual development. The pineal
nerve
impulses gland, then, appears to function as a ‘biological clock’ and
acts aS a ‘neuroendocrine transducer’ converting cyclic
nervous activity, generated by light, into endocrine
neurosecretory secretion. Much more research is required to clarify the
cell body
many functions of this gland.
nerve
impulses mm 16.6.4 Parathyroid glands
pass down | |
axon
There are four small parathyroid glands in
Man and they are embedded in the thyroid gland. They
produce only one hormone called parathormone which is a
mitochondrion peptide composed of 84 amino acids. Parathormone and
the thyroid hormone calcitonin work antagonistically to
neurosecretory
vesicle regulate the plasma calcium and phosphate levels. The
neurohaemal
basement region
release of parathormone increases the plasma calcium level
membrane to its normal level of 2.5 mmol dm~’ and decreases the
hormones released
plasma phosphate level. The activity of the parathyroid
from neurosecretory glands is controlled by the simple negative feedback
granules into the mechanism shown in fig 16.50.
bloodstream me ICUSIOn
ee endothelial Overactivity of the gland, hyperparathyroidism, reduces
eG
capillary wall cell the level of calcium in the plasma and tissues due to calcium
Fig 16.49 Diagram showing a neurosecretory cell and
excretion in urine. This can lead to a state of tetany in
which muscles remain contracted. Also the rate of
J
neurohaemal organ, (not drawn to scale)
excretion of phosphate is reduced and the level of
phosphate ions in the plasma rises.
Afferent nerve stimuli are relayed to the cell bodies of
high Ca?+
these neurosecretory cells from other regions of the brain
and transmitted down the axons to the neurohaemal organ PO,*-
high
where secretions stored in vesicles are released into the
\
bloodstream and carried to target organs. The physio-
logical response produced by such a circuit involving the rae
translation of sensory information into hormonal secre- calcitonin parathormone
tions, and eventually a physiological response, is known as
a neuroendocrine response. Many neuroendocrine res- a
ponses result in a type of behavioural pattern known as a
neuroendocrine reflex and many examples of these are low
PO,>
associated with courtship and breeding activity. -
low Ca2+
16.6.3 Pineal gland or epiphysis, (ep/, upon: Fig 16.50 Hole of parathormone and calcitonin in the
phyein, to grow) regulation of blood calcium level
The pineal gland is an extremely small gland
found in the roof of the forebrain (diencephalon) and 16.6.5 Thyroid gland
covered by the corpus callosum and cerebral hemispheres. The thyroid gland in Man is a bow-tie-shaped
It is variable in size and weighs about 150 mg. The Romans structure situated in the neck region with lobes, on each
believed the pineal gland to be the ‘seat of the soul’ but it is side of the trachea and larynx, connected by a thin band
now recognised as an endocrine gland. It has no direct of tissue. The entire gland weighs approximately 25 g and
connection with the central nervous system but is richly produces three active hormones, triiodothyronine (T;),
vascularised and believed to secrete several hormones, thyroxine (T,) and calcitonin. T; and T, have a regulatory
including melatonin. It is innervated by sympathetic effect on metabolic rate, growth and development, whilst
neurones which are stimulated by centres in the brain calcitonin is involved in the regulation of the plasma
controlled by light. Exposure to darkness stimulates calcium level.
melatonin synthesis which acts via the brain and modifies
the function of the thyroid and adrenal glands and gonads. The structure of the thyroid gland
There is also evidence to suggest that melatonin acts on the The thyroid gland is made up of numerous follicles which
brain and influences several physiological processes which have a diameter of 0.1 mm and contain a clear colloid
are dependent upon time, such as the onset of puberty, composed of the glycoprotein thyroglobulin. The wall of
ovulation and sleep. Tumours of the pineal gland in young each follicle is composed ofa single layer of cuboidal cells
606
I I NH, hormones such as insulin, adrenaline and glucocorticoids
(section 18.5.2).
HO OC) hh co
Control of T; and T, release
I The effects of T,; and T, are longer lasting than those of
3, 5, 3-triiodothyronine (T;) most other hormones and hence it is homeostatically vital
that fluctuations in their secretion be prevented. This is one
I I of the reasons for the storage of T; and T, within the gland
NH, :
so that it is readily available for release.
HO 1
ho The levels of T; and T, circulating in the blood control
their release from the thyroid gland by negative feedback
I l mechanisms involving the hypothalamus and anterior
pituitary.
3, 5, 3, 5-tetraiodothyronine (T,)
Thyrotrophic releasing hormone (TRH) and TSH release
thyroxine
is inhibited by the amounts of T,; and T, in the blood in
Fig 16.51 Structural formulae of the main thyroid hormones excess of those required to maintain the metabolic rate at a
T3 and T, steady state. Superimposed on these are environmental
factors, such as temperature, which influence higher
that become columnar and bear microvilli on their inner centres of the brain to stimulate the hypothalamus to
surface when the gland is activated by thyroid stimulating secrete TRH. This ‘sets’ the threshold in the pituitary for
hormone (TSH) from the anterior pituitary gland. the negative feedback mechanisms. These mechanisms of
The formation and release of thyroid control are shown in fig 16.52.
hormones
— — — — p inhibition
Iodine is taken up by active transport from the plasma, in
——_——» stimulation
the numerous capillaries surrounding the follicles, as
iodine ions (I) and secreted into the lumen of the follicle.
Here it is bound to a protein molecule and oxidised to
iodine by peroxidase enzymes which form part of a
cytochrome enzyme system. The iodine formed reacts with cold
|
warmth
the amino acid tyrosine which is bound to thyroglobulin |
formed by the follicle cells. Further iodination occurs and

subsequent interactions give rise to the two main iodinated hypothalamus Fi
\
amino acid hormones shown in fig 16.51.
The function of T, and T,
OR TRH T, and T,
Whilst T,; and T, have a major influence upon many regulates
metabolic processes, including carbohydrate, protein, fat ‘set-point’
for thyroxine
and vitamin metabolism, their main influence is upon the level in =
rate of metabolic processes, an effect known as calori-
\
genesis, (calor, heat, genesis, production). The basal
metabolic rate (BMR) in Man, 160 kJ m~™ body surface
h7!, is maintained at a steady state by the action of TSH
thyroxine which promotes the breakdown of glucose and
fats into forms which can readily yield energy (section
18.4.2). Further calorigenic effects include increases in the erandelly,
thyroid
uptake of oxygen by the body and the rate of enzyme
reactions involved in the electron transport chain in the
mitochondria. This increases the rate of ATP formation
and heat production by the tissues. The exact mechanism of ivanell,
this action is not known.
T; and T, and the growth hormone, somatomedin, have a
joint stimulatory effect on protein synthesis leading to an
increase in growth rate. This effect is clearly seen in frog blood/tissues
metamorphosis (section 21.7.4) and in children suffering
from a lack of thyroid hormones. its Fig 16.52 Summary diagram showing the factors regulating
In many of the metabolic processes with which it is thyroxine secretion and leading to homeostatic control of
metabolic rate
involved thyroxine appears to enhance the effects of other
607
Overactivity of the thyroid gland (hyper- a condition known as myxoedema and the symptoms are a
thyroidism) reduction in metabolic rate accompanied by decreased
oxygen consumption, ventilation, heart rate and body
Both over- and underactivity of the thyroid gland can
temperature. Mental activity and movement become
produce a swelling in the neck known as a goitre.
slower and weight increases due to the formation and
Overactivity may be due to overproduction of thyroxine
storage of a semi-fluid material under the skin. This causes
from an enlarged thyroid gland. The symptoms are
the face and eyelids to become puffy, the tongue swells, the
increases in heart rate (tachycardia) (section 18.1.5),
skin becomes rough and hair is lost from the scalp and
ventilation rate and body temperature. The basal metabo-
eyebrows. All of these symptoms can be eliminated and the
lic rate may increase by 50% with associated increases in
condition treated by taking thyroxine tablets.
oxygen consumption and heat production. Patients
become very nervous and irritable and the hands shake Calcitonin
when held out. Extreme hyperthyroidism is termed In addition to follicular cells the thyroid gland contains
thyrotoxicosis and is associated with increased excitability other cells known as C cells. These secrete a polypeptide
of cardiac muscle which may lead to heart failure unless hormone calcitonin which works antagonistically towards
treated. This may involve the surgical removal of most of parathormone and lowers blood calcium levels.
the gland or the destruction of the same amount by
administering radioactive iodine. 16.6.6 Adrenal glands
Underactivity of the thyroid gland There are a pair of adrenal (ad, to; renes,
(hypothyroidism) kidneys) glands weighing approximately 5 g each which are
A lack of TSH production by the anterior pituitary, iodine situated anterior to the kidneys. Each gland is composed of
deficiency in the diet or failure of enzyme systems involved cells having two different embryological origins, and these
in thyroxine production may result in hypothyroidism. If cells function independently. The outer cortex is derived
there is a deficiency of thyroxine at birth this will lead to from cells of the neural plate and forms 80% of the gland. It
poor growth and mental retardation, a condition known as is a firm structure and covered by a fibrous capsule and is
cretinism. If the condition is diagnosed at an early stage essential for life. The inner medulla is derived from cells of
thyroxine can be given to restore normal growth and the neural crest and has retained its close origin with the
development. Thyroxine deficiency in later life gives rise to nervous system. It is not essential for life.
Table 16.13. Summary table showing the main cytological detail, secretion and functions of the three regions of the
adrenal cortex
Region Cytological detail Secretion Function Notes
Zona Composed of rounded Mineralocorticoids Control water and electrolyte Renin released from juxta-
glomerulosa clusters of small cells e.g. aldosterone metabolism by stimulating cation glomerular apparatus in kidney
containing elongated pumps in membranes to conserve produces angiotensin. This
mitochondria Na’ and Cl and remove K’. stimulates release of aldo-
Prevent excessive Na’ loss in sterone which increases Na"
sweat, saliva and urine and uptake by kidney and leads to
maintain osmotic concentration release of ADH which in-
of body fluids at a steady state creases reabsorption of water
by kidney tubules. Release is
not stimulated by ACTH
(A) Carbohydrate metabolism Overactivity leads to Cushings’
(1) promote gluconeogenesis syndrome and patients show
(2) promote liver glycogen abdominal obesity, wasting of
formation muscles, high blood pressure,
Zona 2-3 rows of parallel Glucocorticoids (3) raise blood glucose level diabetes and increased hair
fasciculata columns of cells e.g. cortisol growth. Overproduction of
(B) Protein metabolism
arranged at right-angles to ACTH by the anterior
the surface of the gland
(1) promote breakdown of
plasma protein pituitary is Cushings’ disease.
and containing spherical Underactivity leads to
mitochondria
(2) increase availability of
amino acids for enzyme Addison’s disease as shown by
synthesis in the liver muscular weakness, low blood
pressure, decreased resistance
Zona Composed of irregularly (C) Other roles to infection, fatigue and
reticularis arranged columns of (1) prevent inflammatory and darkening of the skin
cells containing allergic reactions
elongated mitochondria (2) decrease antibody
production
SS. —~wW~0w ee6$3uqou0uQqoOoee_— a Se
608
Adrenal cortex Control of cortical hormone release
The adrenal cortex has three histologically distinct regions Mineralocorticoid release is stimulated by the activity of
composed of cells grouped around blood capillaries and renin and angiotensin as described in table 16.13, whilst
produces steroid hormones of two types. These regions, glucocorticoids are secreted in response to adrenocortico-
their cytological detail, secretions and functions are shown trophic hormone (ACTH). The mechanisms of the release
in table 16.13. of these hormones is shown in fig 16.53.
All steroids are formed from a common precursor ACTH is a polypeptide molecule containing 39 amino
molecule called cholesterol which the cortex is able both to acids. It attaches to receptors on the surface of the cortical
synthesise and take up from the circulation following cells and activates adenyl cyclase to convert ATP to cyclic
absorption from the diet. The three regions of the cortex AMP (section 16.6.1). This acts as a cofactor in activating
then convert it into specific steroid hormones. enzymes known as protein kinases which stimulate the
Steroids are lipid-soluble substances which diffuse conversion of cholesterol to pregnenolone. In addition to
through cell membranes and attach to cytoplasmic receptor this ACTH is responsible for the uptake of cholesterol into
proteins. The complexes formed then migrate into the the cortex, the maintenance of the size of the adrenal
nucleus where they attach to specific areas of the cortex and the maintenance of the enzymes involved in
chromosome and de-repress or activate certain genes steroid production.
inducing the formation of messenger RNA.
Adrenal medulla
The size of the adrenal gland is closely linked to the
output of ACTH and the ability to withstand stress. During The adrenal medulla forms the centre of the adrenal gland.
long periods of stress the size of thé gland increases. It is a soft tissue composed of strands of cells surrounded by
Investigations into the behaviour of organisms under stress blood capillaries and is richly supplied with nerves. The
have shown that the output of adrenal hormones increases cells are modified postganglionic sympathetic neurones
with the rise in number in the population. In organisms and when stimulated by preganglionic sympathetic
where social hierarchies exist, there is a positive correlation neurones they secrete noradrenaline and adrenaline (sec-
between position in the hierarchy and increased size of the tion 16.6.1). The adrenal medulla is not essential to life
adrenal gland. since its function is to augment that of the sympathetic
nervous system.
Noradrenaline (norepinephrine) and adrenaline
(epinephrine) are formed from the amino acid tyrosine and
low plasma Nat ‘biological pain fever disease anxiety
clock’
belong to a group of biologically active molecules called
catecholamines (fig 16.54). They are both secreted by the
sleep cells of the medulla but only noradrenaline is secreted by
kidney the postganglionic synapses of the sympathetic nervous
(juxtaglomerular hypothalamus a system. The effects of both hormones are basically
apparatus)
identical as shown in table 16.14, but they differ in their
adrenocorticotrophic
releasing factor HO H H O H
(ACRF)
cc | |
renin HO = CNH HC C—C—N—CH,
| |
anterior pituitary HO Be HO H Ee
lela
cyclic AMP
plasma proteins noradrenaline adrenaline
(angiotensinogen)
ACTH-secreting cells
Fig 16.54 Molecular structure of noradrenaline and adrenaline
Table 16.14. Physiological effects of noradrenaline and

iotensin adrenocorticotrophic negative adrenaline
hormone (ACTH) feedback
ats
Dilate pupils of eyes
Cause hair to stand on end
Relax bronchioles thus increasing air flow to lungs
zona fasciculata
and ; Inhibit peristalsis
zona glomerulosa
zona reticularis Inhibit digestion
Prevent bladder contraction
Increase amplitude and rate of heartbeat
Cause almost general vasoconstriction
cortisolp= — — — — — — — Increase blood pressure
aldosterone
(mineralocorticoid) (glucocorticoid) Stimulate conversion of liver glycogen to glucose
Decrease sensory threshold
Fig 16.53 Summary diagram for the control of the main Increase mental awareness
adrenal cortex hormones
609
effects on blood vessels. Noradrenaline causes vasocon- Table 16.15. Some of the effects of insulin deficiency
striction of all blood vessels whereas adrenaline causes and excess
eo Ma\wwmwaowo
vasoconstriction of blood vessels supplying the skin and gut

and vasodilation of blood vessels to muscles and the brain. Deficiency Excess
Both of these hormones activate two types of receptor sites High blood glucose level Low blood glucose level
on the target tissues known as o. and f adrenergic receptors. (hyperglycaemia) (hypoglycaemia)
These release adenyl cyclase (cyclic AMP) into the cell and Breakdown of muscle tissue Hunger
this leads to the specific tissue response as shown in fig Loss of weight Sweating
16.53. Most organs have both a- and £-receptors with Tiredness Irritability
a-receptors appearing to be more receptive to noradrena- Double vision
line than adrenaline and vice versa in the case of
B-receptors.
from the pancreas of animals. In 1950 Sanger, in
The actions of these hormones are widespread through-
Cambridge, succeeded in determining the primary struc-
out the body and prepare the animal for situations of ‘fight
ture of insulin and opened up the way for its synthetic
or flight’ and facilitate the response to sudden demands
imposed by stress such as pain, shock, cold, low blood preparation. The effects of insulin deficiency and excess are
sugar, low blood pressure, anger and passion. summarised in table 16.15.
16.6.7 Pancreas Glucagon

Glucagon is a peptide composed of 29 amino acids and is
The pancreas has both exocrine and endocrine
released, along with several other hormones, in response to
functions and is associated with the alimentary canal
a fall of blood glucose level induced by an increase in
(section 10.4). The bulk of this gland is composed of
exocrine acinar cells which form rounded structures
metabolic demand. It stimulates the breakdown of proteins
enclosing a lumen into which the digestive enzymes of the and fats to carbohydrates (gluconeogenesis) (section
zymogen granules are secreted (section 10.4.9). Inter- 18.5.2). Receptor sites in the liver cell membrane bind
spersed amongst groups of acinar cells are the islets of glucagon which causes the release of adenyl cyclase to form
Langerhans containing a small number of large alpha (a) cyclic AMP. The action of glucagon is similar to that of
cells, numerous small beta ((}) cells and blood capillaries. adrenaline and both activate phosphorylase enzymes which
Alpha cells secrete glucagon and beta cells insulin and stimulate the breakdown of glycogen to glucose-6-
these have antagonistic effects on the blood glucose level. phosphate as shown in fig 16.47. Glucagon has no effect on
muscle glycogen.
Insulin Summary diagrams of the role of insulin and glucagon
Insulin is a protein composed of 51 amino acids and is are shown in figs 18.7 and 18.23.
released in response to a rise in blood glucose level above
100 mg 100 cm™ blood andarise in glucagon level. It is 16.6.8 Differences between nervous and en-
carried in the plasma bound to f-globulin and has an docrine coordination
important anabolic effect on every organ of the body. The nervous and endocrine systems function
Receptor sites on cell membranes bind insulin, and this separately and together in the coordination of many of the
reaction leads to changes both in cell permeability and the body’s activities. This chapter describes both situations but
activity of enzyme systems within the cell with the following it is useful as a summary to examine the differences
effects: between the two systems as this highlights the advantage of
(1) increase in the rate of conversion of glucose to each system in the light of their roles (table 16.16).
glycogen (glycogenesis),
(2) increase in the rate of uptake of glucose by muscle and Table 16.16. Differences between nervous and endocrine
fat by cell membranes, coordination
(3) increase in the rate of protein and fat synthesis,
(4) increase in the formation of ATP, DNA and RNA. Nervous Chemical
aaa cre
The secretion of insulin is vital to life since it is the only
Information passes as Information passes as a
hormone which lowers the blood glucose level. A defici- electrical impulses along chemical substance through the
ency in insulin production leads to the metabolic condition axons (chemical across bloodstream
known as diabetes mellitus in which the blood glucose synapses)
reaches such a level that it exceeds that which the kidney Rapid transmission Slow transmission
can reabsorb, the renal threshold, and is excreted in the Response immediate Response usually slow, e.g.
urine. The involvement of the pancreas in diabetes mellitus growth
has been known for almost a hundred years but it was not Response short-lived Response long-lasting
until 1921 that the Canadian doctors Banting and Best Response very exact Response usually widespread
succeeded in treating the condition with insulin extracted e—aooeoos$“=@s@>s@>"@mmwwO0O9aoawmwWDOOOOOO—SS So
610
16.7 =‘The study of behaviour (ethology) Recent advances in audio-visual technology have
assisted the recording of behavioural activities. Infra-red
Behaviour may be defined as the outwardly photography has enabled animals to be filmed at night and
expressed course of action produced in organisms in time-lapse photography and slow-motion cinematography
response to stimuli from a given situation. The action have enabled respectively slow-moving activities, such as
modifies, in some way, the relationship between the moulting in insects, and fast-moving activities, such as bird
organism and its environment and its adaptive significance flight, to be recorded and subsequently seen at speeds more
suited to analysis by behaviouralists. The use of miniature
is the perpetuation of the species. All living organisms
exhibit a variety of forms of behavioural activity deter- cassette tape recorders for recording sounds and their
subsequent analysis using sound spectrographs and compu-
mined by the extent to which they are able to respond to
ters has helped in the study of auditory communication
stimuli. This varies from the relatively simple action of the
between organisms. The movement of organisms is now
growth of a plant stem towards a light source, to the
studied either using implanted miniaturised signal gener-
complex sexual behaviour patterns of territory defence,
ators, emitting signals that can be followed using direction-
courtship and mating seen in birds and mammals.
finding equipment, or by the use of tracking radar. The two
Plant behaviour is restricted to movements produced by
techniques are employed successfully in following the
growth or turgor changes and is stereotyped and predict-
migrations of mammals, birds and locusts.
able. The two main activities associated with plant
Whatever the approach and techniques used in the
behaviour are tropisms and nasties and details of these are
investigation of behaviour the fundamental explanation of
described in section 15.1.
behavioural activity must begin with a stimulus, end with a
Animal behaviour is far more complex and diverse than
response and include all the stages occurring at various
plant behaviour and therefore it is extremely difficult to
levels of organisation within the body linking cause and
investigate and account for with any degree of scientific
effect.
validity. The three main approaches to behavioural studies
Broadly speaking there are two forms of behaviour,
are the vitalistic, mechanistic and ethological approaches.
innate behaviour and learned behaviour, but the distinction
Vitalistic approach. This seeks to account for between the two is not clear-cut and the majority of
behavioural activities in terms of what animals are seen to behavioural responses in higher organisms undoubtedly
do, and attempts to relate this to changes in the contain components of both. However, for simplification in
environment. It involves the total rejection of any study of this elementary introduction to behaviour, the various
the animal outside its natural environment. The technique aspects of behaviour are considered under these two
has its foundations in natural history and has provided a headings in the next two sections.
wealth of valuable data, but it is essentially non-scientific
since all the observations relate to past events which cannot
be tested experimentally. 16.8 Innate behaviour
Mechanistic approach. This is an _ ex- Innate behaviour does not involve a single
perimental approach and involves the study of particular clear-cut category of behaviour but rather a collection of
aspects of behaviour under controlled conditions in a responses that are predetermined by the inheritance of
laboratory. It may be criticised on the grounds of the specific nerve or cytoplasmic pathways in multicellular or
artificiality of the experimental situation, the nature of the acellular organisms. Asa result of these ‘built-in’ pathways
behavioural activities and the way in which the results are a given stimulus will produce, invariably, the same
interpreted. This technique is, however, used extensively response. These behaviour patterns have developed and
in psychology and was pioneered by Pavlov. been refined over many generations (selected) and their
Ethological approach. This is the contem- primary adaptive significance lies in their survival value to
porary approach to behavioural investigations and the species. Another valuable feature of innate behaviour
attempts to explain responses observed in the field in terms is the economy it places on nerve pathways within
of the stimuli eliciting the behaviour. It involves both of the multicellular organisms, since it does not make enormous
techniques outlined above and was pioneered by Lorenz, demands on the higher centres of the nervous system.
von Frisch and Tinbergen. There is a gradation of complexity associated with
patterns of innate behaviour which is related to the
In all behavioural studies great care has to be taken in complexity of nerve pathways involved in their perfor-
mance. Innate behaviour patterns include orientations
interpreting the results of observations in order to
(taxes and kineses), simple reflexes and instincts. The latter
eliminate subjectivity. For example, care must be taken to
are extremely complex and include biological rhythms,
avoid putting oneself in the place of the animal (anthropo-
territorial behaviour, courtship, mating, aggression, altru-
centrism), or interpreting what is observed in terms of
ism, social hierarchies and social organisation. All plant
human experience (anthropomorphism) or interpreting the
behaviour is innate.
cause of the observation in terms of its outcome (teleology).
611
Taxes Experiment 16.1: To investigate orientation
A taxis or tactic response is a movement of the whole
behaviour in woodlice by the use of a simple
choice chamber.
organism in response to an external directional stimulus.
Tactic movements may be towards the stimulus (positive, Materials
+), away from the stimulus (negative, —), or at a particular old pair of tights
angle to the stimulus, and are classified according to the bases of 2 Petri dishes
nature of the stimulus. Some examples of types of taxes are Araldite
shown in table 15.2. In some cases organisms are able to hot metal rod
move by maiutaining a fixed angle relative to the cotton wool
directional stimulus. For example, certain species of ants anhydrous calcium
can follow a path back to their nest by setting a course chloride
relative to the sun’s direction. Other organisms orientate adhesive tape
themselves so that, for example, their dorsal side is always ten woodlice
uppermost. This is called the dorsal light reaction and is plasticine
found in fish such as plaice which maintain their dorsal
surface at right-angles to the sky. Method
Many organisms detect the direction of the stimulus by (1) Cut a circle out of an old pair of tights 10cm in
moving the head, which bears the major sensory receptors, diameter and stretch over the base of an 8.5 cm Petri
from side to side. This is known as a klinotactic response dish. Attach with Araldite, held in place by an elastic
and enables symmetrically placed receptors on the head, band until it sets.
such as photoreceptors, to detect the stimulus. If both (2) Burn out a 1.0 cm hole in the bottom of this Petri dish
receptors are equally stimulated the organism will move using a hot metal rod.
forwards in approximately a straight line. This type of (3) Divide the base of another Petri dish in half using a
response is shown by Planaria moving towards a food plasticine strip 8.5 cm long, 1.4 cm deep and 0.5 cm
source and by blowfly larvae moving away from alight wide.
source. In all cases of klinotaxis it is thought that successive (4) Place cotton wool soaked in water in one half of this
stimulation of receptors on each side of the body is Petri dish and granules of anhydrous calcium chloride
necessary in order to provide the ‘brain’ with a contin- in the other half.
uous supply of information since there is no long-term (5) Attach the Petri dish base prepared in (1) above to the
‘memory’. Petri dish base prepared in (4) with adhesive tape as
Kineses shown in fig 16.55.
(6) Introduce ten woodlice into the apparatus through the
A kinetic response is a non-directional movement response
hole in the upper Petri dish and record the position and
in which the rate of movement is related to the intensity of
number active at 1 min intervals in a table such as
the stimulus and not the direction of the stimulus. For
table 16.17.
example, the direction of movement of the tentacles of
(7) After 20 min plot a graph of numbers present against
Hydra in search of food is random and slow, but if saliva,
time for each environment.
glutathione or water fleas are placed in the immediate
(8) Calculate the percentage number of woodlice active in
vicinity of the Hydra the rate of movement of the tentacles
the dry environment for each minute interval and plot
increases.
on a graph against time.
Both kinetic and tactic responses are observed through
(9) Explain the nature of the results obtained in terms of
the use of woodlice in a choice chamber as described in
kineses and taxes.
experiment 16.1.
adhesive tape woodlice bases of two Petri dishes

Table 16.17. Specimen arrangement of table of results
stretched ‘tights’ hole for introducing woodlice Humid Dry
Timel Number Number Number Number Percentage

min present active present active active dry side
0 4 1 6 4 40%
1 5 a 5 5 50%
2 etc. etc.
3
moist cotton wool strip of plasticine anhydrous calcium
chloride granules
Fig 16.55 Choice chamber apparatus for investigating 20
orientation behaviour in woodlice.
612
In simple experiments of this type the response of behaviour patterns of immediate adaptive survival value to
organisms to environments having extremes of a given the organism and are produced in response to sudden
variable can be investigated. Tactic responses are observed changes in the environment. They are unique to each
by the preference shown by the organisms for a particular species and differ from simple reflexes in their degree of
environment. For example, woodlice exposed to areas of complexity. Konrad Lorenz, a Nobel prize-winning etho-
high and low humidity in a choice chamber congregate in logist, defined instincts as ‘unlearned species-specific
larger numbers in the area of highest humidity, showing motor patterns’.
them to be positively hydrotactic. More complex experi- Instinctive behaviour is predominant in the lives of
ments can be devised using combinations of variables in invertebrate animals where, in insects in particular, short
order to determine which is strongest in eliciting a final life cycles prevent modifications in behaviour occurring as a
response. result of trial-and-error learning. Instinctive behaviour in
Kinetic responses are observed by recording the activity insects and in vertebrates therefore is a ‘neuronal economy
of woodlice at, say, 30s intervals, in relation to their measure’ and provides the organism with a ready-made set
position in the choice chamber. Results of such investiga- of behavioural responses. These responses are handed
tions show that when first introduced into the choice down from generation to generation and, having under-
chamber at the junction of two environments some gone successfully the rigorous test of natural selection,
woodlice move around whilst others remain stationary. clearly have important survival significance.
After a short time all the woodlice begin moving and the However, before concluding that instinctive behaviour
speed of movement and rate of turning is always greatest in patterns are completely inflexible as a result of their genetic
the drier side of the choice chamber than in the humid side. origin it must be stressed that this is not so. All aspects of
The increased, apparently random, moving and turning of the development of an organism, whether anatomical,
the woodlice on the dry side is believed to indicate an biochemical, physiological, ecological or behavioural, are
attempt to find optimal conditions and, when these are the result of the influence of constantly varying environ-
found, the moving and turning response diminishes. These mental factors acting on a genetic framework. In view of
responses are examples of orthokinesis. The woodlice this no behavioural pattern can be purely instinctive (that is
move more slowly on the humid side and consequently genetic) or purely learned (that is environmental), and any
usually congregate there. The preference shown for the subsequently described behavioural activity, whilst being
humid side of the choice chamber indicates a positively either superficially instinctive or superficially learned, is
tactic response to humidity. influenced by both patterns. Some authorities prefer the
Not all orientation behaviour patterns are rigid, and the terms species-characteristic behaviour, in preference to
instinctive behaviour, and individual-characteristic be-
response shown by an organism may vary depending upon
other factors such as the degree of hunger, thirst, light, haviour, in preference to learned behaviour. But despite
dark, heat, cold and humidity. this terminology the same principle of genetic and
environmental interaction applies. This point was demons-
16.8.1 Simple reflexes in vertebrates trated very clearly by Professor W. H. Thorpe in his
investigations of chaffinch song. He found that chaffinches
A simple reflex is an involuntary stereotyped whether reared by parents, reared in isolation, or deaf from
response of part of an organism to a given stimulus. It is the time of hatching, all produce sounds clearly identifiable
determined by the presence of an inherited pattern of to the human ear as those of a chaffinch. Sound
neurones forming spinal and cranial reflex arcs, and the spectrograms show, however, that these are only
structure and function of these is described in section 16.2. rudimentary songs, and that chaffinches reared by parents,
In terms of behaviour, simple spinal reflexes are either listening to the songs of their parents and other chaffinches
flexion responses, involving withdrawal of a limb from a in the population, develop identical sound patterns to the
painful stimulus, or stretch responses, involving the older birds, characteristic of the local population. It was
balance and posture of the organism. Both of these apparent that bird songs within a species have ‘local
responses are primarily involuntary and most require no dialects’. Songs of deaf birds or those in isolation remained
integration or coordination outside that found in the spinal rudimentary, thus demonstrating that the environment can
cord. However both types of response may be modified by significantly modify an instinctive pattern.
the brain according to circumstances and in the light of
previous experience. When this happens innate and
16.8.3 Motivation
learned behaviour patterns overlap and the reflex action 1s
now described as ‘conditioned’ as described in table 16.18. The extent and nature of any behavioural
for
Many simple cranial reflexes too, may be conditioned, response is modified by a variety of factors that are
example blinking in response to a sudden movement. collectively known as motivation. For example, the same
stimulus does not always evoke the same response in the
16.8.2 Instincts same organism. The difference is always circumstantial and
Instincts are complex, inborn, stereotyped may be controlled by either internal or external factors.
613
Presenting food to a starved animal will produce a different during hibernation results in awakening and food seeking.
response from that shown by an animal that has been fed. Motivational stimuli provide the ‘drive’ or ‘goal’ preparing
In between the two extremes responses of varying strengths the organism for activity which may be triggered off by the
will be produced depending upon the degree of hunger second type of sign stimulus.
experienced by the organism. However, if the act of Releasing stimuli or ‘releasers’. A releaser
feeding would place a hungry animal in danger of being is either a simple stimulus or a sequence of stimuli
attacked by a predator the feeding response would be produced by a member of a species which evokes a
curbed until the danger passed. Many behavioural res- behavioural response in another member of the same
ponses associated with reproduction have a motivational species. The term ‘releaser’ was introduced by Lorenz and
element. For example, many female mammals are only its role in behaviour was extensively studied by Tinbergen.
receptive to mating attempts by males at certain times of The effect of a releaser was demonstrated during an
the year. These times coincide with the period of oestrous investigation into feeding in herring gulls. Young herring
and have the adaptive significance of ensuring that mating gulls normally peck at a red spot on the yellow lower
coincides with the optimum time for fertilisation and mandible of the parent’s bill to signal the parent to
therefore the production of offspring at the most regurgitate fish which the young then swallow. Ina series of
favourable time of the year. These behavioural patterns are controlled experiments, carried out by Tinbergen and
known as biological rhythms and are described in section Perdeck using cardboard models of adult gulls’ heads, they
16.8.5. In many species the degree of motivation, or found that the releaser of the begging response was the
‘drive’, coincides in males and females, but in other species presence of a contrasting colour on the beak. Such was the
some system of communication between the sexes is
strength of the releaser that a pointed stick with alternating
essential to express the degree of motivation. In many
coloured bands was able to elicit a greater response than
primate species the timing of oestrus is signalled by a
the parent bird, as shown in fig 16.57.
swelling and change of colour of the genital area of the
female and this is displayed to the male. Such behaviour Terminating stimuli. Terminating stimuli, as
reduces the likelihood of a male attempting to mate at a the name implies, complete the behavioural response and
time when the female is not receptive. The signals used to may be external or internal. For example, the external
bring about a change in behaviour are known as sign stimuli visual stimuli of a successfully completed nest will
and, depending upon their origin or function, are classified terminate nest building in birds, whereas the internal
as motivational, releasing or terminating stimuli. satisfaction or ‘satiety’ accompanying ejaculation in the
male will terminate copulation and likewise a full stomach
Motivational stimuli. This type of stimulus will terminate feeding.
may be external, for example increasing day length Further examples of sign stimuli for a selection of be-
inducing territorial and courtship behaviour in birds, or havioural mechanisms are discussed in sections 16.8.4—
internal, for example depleted food stores in the body 16:3:9:
Fig 16.56 Female chimpanzee signalling to the male that she is sexually receptive
614
(a
71
16.8.4 Innate releasing mechanisms
Lorenz suggested that there must exist a
means of filtering out stimuli which are irrelevant from
those that are relevant to producing the correct behaviour-
al response. Investigations suggest that this may occur
peripherally at the receptors or centrally within the central
nervous system. For example, Schneider found that the
chemoreceptors on male moth antennae are only sensitive
to the sex-attracting chemicals (pheromones) produced by
the female of that species and not to those of other species.
Modifications of Tinbergen and Perdeck’s experiments on
the herring gull have been carried out, and the results
suggest, as Lorenz postulated, that centrally situated
neurosecretory mechanisms control the response to sign
stimuli.
34 16.8.5 Biological rhythms

Many behavioural activities occur at regular
intervals and are known as biological rhythms or
biorhythms. Well-known examples of these include the
courtship displays and nesting behaviour of birds in the
spring and the migration of certain bird species in autumn.
The time interval between activities can vary from minutes
to years depending on the nature of the activity and the
HoSooESD
species. For example, the polychaete lugworm Arenicola
marina lives in a U-shaped burrow in sand or mud and
carries out feeding movements every 6-7 min. This cyclical
50
feeding pattern has no apparent external stimulus nor
internal physiological motivational stimulus. It appears
that the feeding pattern rhythm is regulated by a biological
‘clock’ mechanism dependent, in this case, on a ‘pace-
maker’ originating in the pharynx and transmitted through
the worm by the ventral nerve cord.
=S>
—m
Rhythms involving an internal clock or pacemaker are
known as endogenous rhythms, as opposed to exogenous
rhythms which are controlled by external factors. Apart
from examples such as the feeding behaviour of Arenicola,
most biological rhythms are a blend of endogenous and
exogenous rhythms.
In many cases the major external factor regulating the
rhythmic activity is photoperiod, the relative lengths of
day and night. This is the only factor which can provide
Fig 16.57 (a) The horizontal bars indicate the number of a reliable indicator of time of year and is used to ‘set the
pecking responses made by herring gull chicks to a series of clock’. The exact nature of the clock is unknown but the
cardboard models of adult herring gull heads having grey bills clockwork mechanism is undoubtedly physiological and
and spots of varying shade. (From Hinde, R. A. 1966, after may involve both nervous and endocrine systems. The
Tinbergen, N., 1951) effect of photoperiod has been studied extensively in
(b) The artificial bill, coloured red with three white bars evoked
20% more pecks than an accurate three-dimensional model relation to behaviour in mammals, birds and insects and,
of an adult herring gull head and beak, coloured yellow with a whilst it is evidently important in activities such as
red spot. (From Tinbergen and Perdeck, 1950, Behaviour, preparation for hibernation in mammals, migration inbirds
3,7) and diapause in insects, it is not the only external factor
regulating biological rhythms. Lunar rhythms, too, can
influence activity in certain species, such as the palolo
worm of Samoa. The polychaete worm swarms and mates
throughout the whole South Pacific on one day of the year,
the first day of the last lunar quarter of the year, on average
615
Light regime indicate that in the absence of an exernal time-cue the
circadian rhythm persisted even though the onset of
ee activity varied by a small amount each day. These results
are consistent with the idea that circadian rhythms are
controlled by an endogenous mechanism or ‘clock’,
governed or ‘set’ by exogenous factors.
Circadian rhythms are believed to have many species-
specific adaptive significances and one of these involves
orientation. Animals such as fish, turtles, birds and some
insects which migrate over long distances are believed to
use the sun and stars as a compass. Other animals, such as
honeybees, ants and sandhoppers use the sun as a compass
A in locating food and their homes. Compass orientation by
sun or moon is only accurate if organisms using it possess
some means of registering time so that allowances can be
made for the daily movements of the sun and moon. The
increasingly familiar concept of ‘jet-lag’ is an example of a
situation where Man’s internal physiological circadian
rhythm is out of step with the day-and-night rhythm of the
destination.
Fig 16.58 Results of cockroach activity over a 20 day 16.8.6 Territorial behaviour
period. During days 1-10 the cockroach received a light
regime of 12 hour light: 12 hour dark cycle and between days A territory is an area held and defended by an
11-20 the cockroach was kept in constant darkness as
organism or group or organisms against organisms of the
indicated in the figure. The black areas shown for each day
represent the time and duration of each ‘burst’ of activity same, or different, species. Territorial behaviour is
common in all vertebrates except amphibia but is rare in
invertebrates. Much research into the nature and function
2nd November. The influence of lunar rhythms on tidal of territoriality has been carried out on birds and groups of
variations is well known, and these are two exogenous primates. In the latter it forms an important part of their
factors which have been shown to impose a rhythmic social behaviour.
behaviour pattern on the midge Clunio maritimus. The The exact function of territory formation probably varies
larvae of Clunio feed on red algae growing at the extreme from species to species, but in all cases it ensures that each
lower tidal limit, a point only uncovered by the tide twice mating pair of organisms and their offspring are adequately
each lunar month. Under natural conditions these larvae spaced to receive a share of the available resources, such as
hatch, the adults mate and lay eggs in their two-hour-long food and breeding space. In this way the species achieves
life during which they are uncovered by the tide. In optimum utilisation of the habitat. The size of territories
laboratory conditions of a constant 12 h light — 12 h dark occupied by any particular species varies from season to
photoperiod the larvae continued to hatch at about 15-day season, according to the availability of environmental
intervals, demonstrating the apparent existence of an resources. Birds of prey and large carnivores have
endogenous clock programmed to an approximately territories several square miles in area in order to provide
semi-lunar rhythm coinciding with the 14.8-day tidal cycle. all their food requirements. Herring gulls and penguins (fig
The behaviour of many completely terrestrial insects 16.59), however, have territories of only a few square
appears to be controlled by endogenous rhythms related to metres, since they move out of their territories to feed and
periods of light and dark. For example, Drosophila emerge use them for breeding purposes only.
from pupae at dawn whereas cockroaches are most active Territories are found, prior to breeding, usually by
at the onset of darkness and just before dawn. These males. Defence of the area is greatest at the time of
regularly occurring biological rhythms, showing a periodic- breeding and fiercest between males of the same species.
ity of about 24 h, are known as circadian (circa, about; There are a variety of behavioural activities associated with
dies, day) rhythms or diurnal rhythms. In an investigation territory formation and they involve threat displays
of the activity of a cockroach (Periplaneta) under two between owners of adjacent territories. These threat
different light regimes (12 h light and 12 h darkness for 10 displays involve certain stimuli which act as releasers. For
days followed by total darkness for 10 days), the cockroach example, Lack demonstrated that an adult male robin
restricted its activity in the later regime to a time (Erithacus rubecula) would attack a stuffed adult male
approximately related to the period of activity associated robin displaying a red breast, and a bunch of red breast
with the onset of darkness under the former light regime. feathers, but not a stuffed young male robin which did not
The results of this investigation are shown in fig 16.58 and have a red breast. The level of aggression shown by an
616
organism increases towards the centre of the territory. The Fig 16.59 Aerial photograph showing the territories occupied
by penguins
aggressiveness of males is determined partly by the level of
testosterone in the body and this can affect territory size.
For example, the territory size of a red grouse can be
increased by injecting the bird with testosterone. Fig 16.60
shows the changes in territory size of three red grouse
treated in this way. The boundary between adjacent
territories represents the point where neighbouring ani-
mals show equally strong defence behaviour. Despite the
apparent conflict and aggression associated with territory Fig 16.60 The solid lines indicate the territories of a group of
formation, actual fighting, which would be detrimental to male red grouse. The dotted lines show changes which
the species, is rare and is replaced by threats, gestures and occurred after birds A, X and Y had received doses of
testosterone. Birds X and Y had not previously held territories.
postures. Having obtained a territory many species, (After Watson, A (1970). J. Reprod. Fert., Suppl., 11.3.)
particularly carnivores, proceed to mark out the boundary
by leaving a scent trail. This may be done by urinating or
rubbing glandular parts of the body against objects called
scent posts along the boundary of the territory. Although
territorial behaviour involves the sharing of available
resources amongst the population there are inevitably
some organisms unable to secure and defend aterritory. In
many bird species, such as grouse, these weaker organisms
are relegated to the edges of the habitat where they fail to
mate. This appears to be one of the adaptive significances
of territoriality as it ensures that only the ‘fittest’ find a
territory, breed and thus pass on their genes to the next
generation. Thus a further function of territorial behaviour
as a
is associated with intraspecific competition and may act
means of regulating population size.
617
16.8.7 Courtship and mating Many insects, amphibia, birds and mammals use audi-
tory signals in courtship. Some species of female mosquito
There are many elaborate and ritualistic attract males by the sounds produced by the frequencies of
species-specific behaviour patterns associated with their wing beats whilst grasshoppers, crickets and locusts
courtship and mating. In birds, mammals and some fish stridulate. This involves either rubbing the hindlegs against
these two processes often follow the establishment of a each other or the elytron (hardened wing case), or rubbing
territory by the male. Courtship is a complex behaviour the elytra together to produce a ‘chirping’ sound which is
pattern designed to stimulate organisms to sexual activity species-specific and only produces a response from mem-
and is associated with pair formation in those species where bers of that species.
both sexes are involved in the rearing of offspring, as in Some species of spiders employ a mechanical means of
thrushes, or in gregarious mixed-sex groups such as attracting the opposite sex. Male spiders approach the web
baboons. The majority of these species show rhythmic of a female sitting at the centre of the web and pluck a
sexual activity of the type described in section 16.8.5. thread of the web at a species-specific frequency. The
Courtship behaviour is controlled primarily by motiva- plucking ‘serenades’ the female and reduces her natural
tional and releasing stimuli and leads to mating which is the aggressive manner so enabling the male to approach and
culmination of courtship. During mating, the behavioural mate her. Unfortunately, if the male ‘woos’ a female of the
activities are initiated by releasing stimuli and ended by wrong species or ‘plays the wrong tune’ he is attacked and
terminating stimuli associated with the release of gametes killed!
by the male. The secretion and release by organisms of small amounts
The motivational stimuli for courtship in most species of chemical substances, leading to specific physiological or
are external, such as photoperiod, and lead to rising levels behavioural responses in other members of the same
of reproductive hormones and the maturation of the species, is used in courtship and mating and, as described
gonads. In most species this produces striking changes in later, the regulation of behaviour within social groups.
the secondary sexual characteristics and other behavioural These substances are called pheromones and are usually
activities including coloration changes, as in the develop- highly volatile, low relative molecular mass compounds.
ment of a red belly in male sticklebacks; increase in size of Many of these compounds function as natural sex attrac-
parts of the body, as in the plumage of birds of paradise; tants and the earliest to be identified were civetone from the
mating calls, as in nightingales; postural displays, as in
civet cat and muscone from the musk deer. Both of these
grebes (fig 16.61) and the use of chemical sex attractants, as substances are secretions from the anal glands and are used
in butterflies and moths. commercially in the preparation of perfume. Mares, cows
Of the variety of signals used in courtship to attract
and bitches secrete pheromones whilst on ‘heat’. This is
members of the opposite sex, sight, sound and smell play
undetectable by human olfactory epithelium but detectable
important roles. For example, the male fiddler crab, Uca,
by the males of the species concerned. Bombykol, a
uses a visual display and attracts females by waving an
pheromone released by eversible glands at the tip of the
enlarged chela in a bowing movement similar to that of a
abdomen of unfertilised adult female silk moths, is capable
violinist. The vigour of the movement increases as a female
of attracting males of the same species from considerable
is attracted to the male.
distances. The olfactory receptors on the antennae of the
male moths detect the presence of the pheromone
molecules in great dilutions and the moths make a
rheotactic response by flying upwind until they reach the
female. Pheromones are used increasingly as a method of
biological control in insect pest species such as the gypsy
moth. In these cases the artificial release of the pheromone,
gyplure, attracts males to the source of release where they
can be captured and killed. This not only immediately
reduces the number of male moths in the population but
also, in preventing them from breeding, reduces the size of
the next generation.
Pheromones are also used to induce mating as in the case
of the queen butterfly Danaus gilippus. The pheromone is
released by the male and brushed on to the female by a pair
of brush-like structures, called hairpencils, everted from
the tip of the abdomen. The entire courting and mating
sequence is shown in fig 16.62.
Fig 16.61 A courtship behavioural activity in great crested
grebes. Male and female grebes are shown here presenting Courtship in some species is accompanied by conflict
nesting material to each other. (After Huxley, J. S. (1914). behaviour on the part of one or both sexes. In species
Proc. Zool. Soc. Lond., 1914 (2), 491-562) where individuals normally live a solitary existence,
618
female behaviour a or offspring and the establishment of rank in a social
appears
ie hierarchy.
on
male behaviour
The term ‘aggression’ is emotive and suggests an
ner pursues in
existence of unnecessary violence within animal groups;
~S as air the alternative term ‘agonistic’ is preferable. Agonistic
—
flies
a ee
behaviour has the adaptive significance of reducing
intraspecific conflict and avoiding overt fighting which is
not in the best interest of the species. Most species channel
Sic overtakes their ‘aggression’ into ritual contests of strength and threat
a and postures which are universally recognised by the species.
. oe hairpencils
alights on For example, horned animals such as deer, moose, ibex
herbage and chamois may resort to butting contests for which
‘ground rules’ exist. Only the horns are allowed to clash
hairpencils and they are not used on the exposed and vulnerable flank.
while Siamese fighting fish, Betta splendens, resort to threat
hovering
postures involving increasing their apparent size as shown
in fig 16.63.
The threats issued by two organisms in an agonistic
conflict situation are settled invariably by one of the
alights
laterally organisms, generally the weaker, backing down and
withdrawing from the situation by exhibiting a posture of
submission or appeasement. In dogs and wolves an
appeasement posture may take the form of the animal lying
down on its back or baring its throat to the victor.
copulates During actual physical contact animals often refrain
from using their most effective weapons on another
|post-nuptial
flight member of the same species. For example, giraffes will
female male
behaviour ie behaviour
fight each other using the short horns on their heads, but in
defence against other animals they use their very powerful
feet.
For agonistic behavioural activities to be most effective
they must be stereotyped for any species and Tinbergen
demonstrated several of these during investigations carried
Fig 16.62 Courtship and copulation in the queen butterfly.
(The arrows indicate the stimuli and responses involved in the
behavioural activities). (From Brower, L. P., Brower, J. V. Z.
and Cranston, F. P. (1965). Zoologica, 50, 18)
courtship conflict may be associated with changing atti-

tudes to other members of the species as a result of
increasing hormone levels. Other significances of this
behaviour may be the tightening of the pair bond between
the mating pair and the synchronisation of gonadial
development so that gametes mature at the same time. In
certain species of spider, such as wolf spiders, conflict
between male and female only diminishes for the act of
copulation which culminates in the female killing the male.
16.8.8 Aggression (agonistic behaviour)

Aggression is a group of behavioural activities
including threat postures, rituals and occasionally physical
attacks on other organisms, other than those associated Fig. 16.63 Stages in threat displays in Siamese fighting fish.
with predation. They are usually directed towards mem- (a) and (b) fish not showing threat displays, (c) and (d)
bers of the same sex and species and have various functions operculum, o, and fins erected to increase their apparent size
including the displacement of other animals from an area, during threat displays. (From Hinde, R. A., 1970, after
mate Simpson, 1968)
usually a territory or a source of food, the defence of a
619
a territorial dispute may begin nest-building activities, such
as pulling up grass, when presented with a choice between
fighting or fleeing. Such displacement activities act as an
outlet for pent-up activities. Many activities in Man may be
considered displacement activities in certain circum-
stances, for example fist clenching, fist banging, nail biting,
straightening clothes, finger drumming, etc. A similar form
of behaviour is called vacuum activity which occurs when
motivation is high and no releaser presents itself. In this
(a)
case the normal response is produced but is not directed
towards the normal object or situation, and so provides a
means of reducing frustration; for example, showing
(b) irritation towards someone who is not the cause of the
irritation but acts as a substitute.
16.8.9 Social hierarchies

Fig 16.64 (above) (a) Full threat posture in male stickleback
(ob) Reduced threat posture when contained in a tube held Many species of insects and most vertebrates
vertically. (After Tinbergen, N., 1951) show a variety of group behavioural activities associated
with numbers of individuals living together temporarily or
permanently. This is known as social behaviour and the
Fig 16.65 (below) Models used as releasers of aggressive
behaviour in male sticklebacks holding a territory. (a) Accurate coherence and cooperation achieved has the adaptive
model not having a red belly does not elicit aggression from significance of increasing the efficiency and effectiveness of
male stickleback. (b)-(c) are models of sticklebacks not the species over that of other species. In a social group of
having an accurate shape but having a red belly. All these this kind a system of communication is essential, and the
models produced aggressive responses from male
sticklebacks. (After Tinbergen, 1951) efficiency of the organisation is further increased by
individuals carrying out particular roles within the society.
One aspect of social behaviour arising out of these points is
the existence of social hierarchies or pecking orders.
A pecking order is a dominance hierarchy. That is to say
(a) (b)
that the animals within the group are arranged according to
status. For example, in a group of hens sharing a hen-house
a linear order is found in which hen A will peck any other
hen in the group, hen B will peck all hens other than A and
(©) (d) (e) so on. Position in the hierarchy is usually decided by some
agonistic form of behaviour other than fighting. Similar
out on sticklebacks. In one series of experiments he patterns of dominance have been observed in other species
demonstrated that the effectiveness of threat posture of birds and in mice, rats, cows and baboons. The
depended on the stickleback assuming a_ horizontal institutional organisation of all human societies is based on
position with fins and spines outstretched. When trapped in a pattern of dominance hierarchy.
a specimen tube and kept vertical a male stickleback does Pecking orders exist only where animals are able to
not have the same threat potential to ward off other recognise each other as individuals and possess some ability
sticklebacks as it has when free-swimming or held to learn. The position of an animal within a pecking order
horizontally (fig 16.64). usually depends on size, strength, fitness and aggressive-
In another series of experiments he demonstrated that ness and, within bird hierarchies, remains fairly stable
agonistic behaviour in male sticklebacks defending a during the lifetime of the individuals. Lower-order male
territory is triggered off by ‘releasers’ which can take the members can be raised up the hierarchy by injections of
form of almost any object whose underside is coloured red. testosterone which increase their levels of aggressiveness.
These objects act as mimics of male sticklebacks whose The experimental removal of lower-order mice from a
bellies turn red during the breeding season and who appear hierarchy and subsequent provision of unlimited food for
to the territory holder as a potential threat (fig 16.65). them increases their mass, improves their vigour and can
At times of stress, for example during conflict situations raise their position in the hierarchy when reintroduced to
or during courtship and mating, an organism may perform the group. Similarly placing lower-order mice into other
an action which is trivial and irrelevant to the situation. groups where they are dominant appears to give them a
This is known as displacement activity and occurs when degree of ‘self-confidence’ (to use an anthropomorphic
motivation is high but two conflicting ‘releasers’ present term) which stays with them when reintroduced to their
themselves. For example, one of a pair of birds involved in original groups and results in their rank increasing.
620
One advantage of pecking order is that it decreases the hundred fertile male drones. Each type of honeybee has a
amount of individual aggression associated with feeding, specific series of roles determined primarily by whether it
mate selection and breeding-site selection. Similarly it hatched from a fertilised or an unfertilised egg. Fertilised
avoids injury to the stronger animals which might occur if eggs are diploid and develop into females; unfertilised eggs
fighting was necessary to establish the hierarchy. Another are haploid and develop into males. Secondly, the type of
advantage of pecking order is that it ensures that resources food provided for female larvae determines whether they
are shared out so that the fittest survive. For example, if a will become queens or workers. This food is called royal
group of 100 hens is provided with sufficient food for only jelly and is one example of the importance of chemical
50 hens it is preferable, in terms of the species, for 50 hens substances in the organisation of the society. Information
to be adequately fed and the weaker 50 hens die than for within the colony is transmitted either by chemical odours
them all to live and receive only half rations, as this might and pheromones during the many licking and grooming
prevent successful breeding. In the short term, social activities called trophallaxes, or by particular forms of
hierarchies increase the genetic vigour of the group by visual orientation displays known as dances.
ensuring that the strongest and genetically fittest animals Karl von Frisch, a German zoologist and Nobel
have an advantage when it comes to reproducing. prize-winner, investigated the nature of these dances using
marked worker bees in specially constructed observation
Social organisation hives. Worker bees ‘forage’ for sources of nectar and
When animals come together to form a cohesive social communicate the distance and direction of the source to
group individuals often assume specialised roles, which other workers by the nature of a dance generally performed
increases the overall efficiency of the group (fig 16.66). on a vertical comb in the hive. If the distance is less than
These roles include members specialised or designated for about 90 m the worker performs a round dance as shown in
food-finding, reproduction, rearing and defence. Coopera- fig 16.67a which intimates that the source is less than 90 m
tion between members of a society sharing division of from the hive but gives no indication of direction. The
labour depends upon stereotyped patterns of behaviour waggle dance is performed if the source is greater than 90 m
and effective means of communication. These patterns of and includes information as to its distance from the hive
behaviour and methods of communication vary between and its direction relative to the hive and the position of the
species and are vastly different for primate and insect sun. The dance involves the worker walking in a figure-of-
societies. Primate societies are flexible, in that roles are eight and waggling her abdomen during which, according
interchangeable between members of the group whereas in to von Frisch, the speed of the dance is inversely related to
insect societies differences in body structure and reproduc- the distance of the food from the hive; the angle made
tive potential affect their role within the society, a feature
called polymorphism.
(a) round dance we ase
Ants, termites and bees are social insects living in yA food SOUTCE NS
colonies and have an organisation based on a caste system.
- within this se
/ : x
/ radius \
In the honeybee colony there is a single fertile female
/ \
/ \
|e—45 \|
queen, several thousand sterile female workers and a few m—->(J
\\ hive |
(b) waggle dance O Sun O
| !
1 |
food |
° source 1
| 1
! |
|
|
|
|
| direction
A of food
Se 190° | direction
Gia aes of food
waggle Rae 9 source
dance fal Geese p=
hive | hive
Fig 16.67 Honeybee dances

(a) Round dance is performed when food is less than 90
metres from hive
es a (b) Waggle dance shows relation between hive, sun and
Fig 16.66 Social grooming in adult chimpanzees provid direction of food source
means of social cohesion within a group
621
between the two loops of the figure-of-eight and the Evidence based on the latter observation suggests that
vertical equals the angle subtended at the hive by the sun; memory is a biochemical event involving the synthesis of
and the food source and the intensity of the waggles is substances within the brain. Extracts of the ‘brains’ of
related to the amount of food at the source (fig 14.67). It is trained flatworms or rats injected into untrained flatworms
thought that allowances for movement of the sun are made or rats reduce the time taken by the latter organisms to
by the use of an inbuilt ‘biological clock’ and that bees learn the same task as compared with control groups. The
orientate on cloudy days by substituting polarised light active substance in all the experiments appears to be RNA.
from the sun for the position of the sun. Further evidence exists which suggests that the composi-
Recent evidence has suggested that bees may use tion of the RNA of neurones changes during learning and
high-frequency sound to communicate sources of food to that this may result in the synthesis of specific ‘memory
other workers, but whether this is the main means of proteins’ associated with the learned behaviour. Investiga-
communication has yet to be demonstrated clearly. This tions have shown that injections of the protein-inhibiting
does not, though, invalidate the data and interpretations of drug, puromycin, also interfere with memory. For ex-
von Frisch and may be an associated communication sys- ample, injecting puromycin into the brains of mice recently
tem which augments the visual dance displays. However, trained to choose one direction in a maze destroyed their
it is known that the returning worker may communicate ability to retain this learning, whereas a control group,
the type of flower visited by feeding the other workers with injected with saline, retained the learned behaviour.
some of the nectar collected. In conclusion, it would appear that the nature of memory
is far from being clarified but it seems probable that
changes in electrical properties of neurones, the permeabil-
ity of synaptic membranes, enzyme production associated
16.9 Learned behaviour with synapses and synaptic transmission are all concerned
with the formation of a ‘memory trace’. Certainly it seems
16.9.1 Memory that memory is associated closely with events occurring at
Memory is the ability to store and recall the synapses.
effects of experience and without it learning is not possible.
Past experiences, in the form of stimuli and responses, are
16.9.2 Learning
recorded as a ‘memory trace’ or engram, and since the
extent of learning in mammals is proportional to the extent Learning is an adaptive change in individual
of the cerebral hemispheres it would appear that these are behaviour as a result of previous experience (fig 16.68).
the site of engram formation and storage. The degree of permanence of newly acquired learned
The nature of the engram is not known and it exists only behaviour patterns depends upon memory storing the
as a hypothetical concept backed up by conflicting data. Of information gained from the experience. In Man, acquiring
the two broad areas of thought on the nature of the engram or learning ‘facts’, for example for examinations, may be
one is based on changes in neuronal structure and short lived whereas the ability to carry out coordinated
organisation within the central nervous system and the motor activities such as toilet training, riding a bicycle or
other based on permanent changes in brain biochemistry.
Histological examination of brain tissue shows the
existence of neurones arranged in loops, and this has given 12
rise to the concept of ‘reverberating circuits’ as units of the £ 10
engram. According to this view these circuits are conti-
aes
nuously active carrying the memory information. It is °
doubtful if this activity could last for any length of time and B 6
experiments suggest that memory has greater performance Ea4
and stability than could be achieved by this mechanism 3s 2
<
alone. For example, cooling the brains of rats down to 0 °C = 0 { [eee af Sew eee: J
causes all electrical activity in the nervous system to cease, 1 2 3 4 5 6 7 8
but on restoring the rats to normal temperatures there is no Trial numbers
impairment in memory. However, it is thought that such
Fig 16.68 The graph above shows a typical ‘learning curve’.
circuits may play a role in short-term memory, that is
The graph shows the results of the number of times a cuttlefish
memory lasting at most for minutes such as memorising strikes at a prawn kept in a glass tube. The prawn was
six-digit telephone numbers in Man, and in facilitating presented to the cuttlefish on eight successive occasions
particular neural pathways. Events associated with short- lasting three minutes each time. As the cuttlefish
term memory take longer to be recalled following unsuccessfully attacks the prawn the number of attacks
decreases as the cuttlefish ‘learns’ that it cannot capture the
concussion or amnesia and gradually disappear in old age.
prawn. (The results are based on data obtained from forty
Long-term memory is more stable and suggests that some cuttlefish, from Messenger, J. B., (1977), Symp. Zoo. Soc.
mechanism for permanent change exists in the brain. Lond., 38, 347-76)
622
swimming, lasts throughout life. Learning is generally ways in different species and different contexts. The
thought of in terms of vertebrates, and mammals in classification and features of learned behaviour presented
particular, but has been demonstrated in all groups of in this chapter are artificial and must be recognised as such.
animals except protozoa, coelenterates and echinoderms They do, however, cover the spectrum of current thinking
where neural organisation is absent or primitive. Psycho- on types of learning and are backed up by experimental
logists have attempted to establish general ‘laws of evidence. A summary of the major types of learned
learning’ but all attempts so far have failed. It would appear behaviour is given in table 16.18 and is designed to provide
that learning is an individual event and occurs in different only an introduction to the topic of learning.
Table 16.18. Summary of the major types of learned behaviour based onaclassification proposed by Thorpe (1963)
Learned behaviour Features of the learned behaviour
Habituation Continuous repetition of a stimulus not associated with reward or punishment (reinforcement) extin-
guishes any response to the stimulus, e.g. birds learn to ignore a scarecrow. Important in development
of behaviour in young animals in helping to understand neutral elements in the environment, such as
movements due to wind, cloud-shadows, wave-action, etc. It is based in the nervous system and is not a
form of sensory adaptation since the behaviour is permanent and no response is ever shown to the
stimulus after the period of habituation.
Classical conditioning Based on the research of Pavlov on dogs. It involves the development of a conditioned salivary reflex in
(conditioned reflex) which animals learn to produce a conditioned response (salivation), not only to the natural unconditioned
stimulus (sight of food) but also to a newly acquired conditioned stimulus (ticking of a metronome) which
was presented to the dog along with the unconditioned stimulus. Animals learn to associate uncondi-
tioned stimuli with conditioned stimuli so either produces a response. For example, birds avoid eating
Associative black and orange cinnabar moth larvae because of bad taste and avoid all similarly coloured larvae even
learning though they may be nutritious.
Based on the research of Skinner on pigeons. Trial motor activities give rise to responses which are
reinforced either by rewarding (positive) or punishment (negative). The association of the outcome of a
Operant conditioning response in terms of reward or punishment increases or decreases respectively future responses.
(trial-and-error learning) Associative learning efficiency is increased by repetition as shown in investigations carried out on
learning in cuttlefish (fig. 16.68).
Latent learning Not all behavioural activities are apparently directed to satisfying a need or obtaining a reward (i.e.
(exploratory learning) appetitive behaviour). Animals explore new surroundings and learn information which may be useful
at a later stage (hence latent) and mean the difference between life and death. For example in mice,
knowledge of the immediate environment of its burrow may help it escape from a predator. At the time
of acquiring this knowledge it had no apparent value. This appears to be the method by which
chaffinches learn to sing, as described in section 16.8.2.
Insight learning Probably the ‘highest’ form of learning. It does not result from immediate trial-and-error learning but
may be based on information previously learned by other behavioural activities. Insight learning is based
on advanced perceptual abilities such as thought and reasoning. Kohler’s work on chimpanzees sug-
gested ‘insight learning’: when presented with wooden boxes and bananas too high to reach the chimps
stacked up the boxes beneath the bananas and climbed up to get them. Observations revealed that this
response appeared to follow a period of ‘apparent thought’ (previous experience of playing with boxes
[latent learning] may have increased the likelihood of the response).
Imprinting A simple and specialised form of learning occurring during receptive periods in an animal’s life. The
learned behaviour becomes relatively fixed and resistant to change. Imprinting involves young animals
becoming associated with, and identifying themselves with, another organism, usually a parent, or some
him and
large object. Lorenz found that goslings and ducklings deprived of their parents would follow
have a
use him as a substitute parent. ‘Pet lambs’, bottle-fed, show similar behaviour and this may
forming
profound and not always desirable effect later in life when the animal finds difficulty in
imprinting has obvious adaptive
normal relationships with others of the species. In the natural situation
learning to fly
significance in enabling offspring to acquire rapidly skills possessed by the parents, e.g.
in birds, and features of the environment, e.g. the ‘smell’ of the stream in which migratory salmon were
hatched and to which they return to spawn.
Oe
623
Chapter Seventeen
Movement and support
Movement can occur at the cellular level, for instance

cytoplasmic streaming and swimming of gametes, at organ
17.1 Skeletal systems
level, such as heartbeat and movement of a limb, or at the
The vast majority of animals possess some
level of the organism. Movement of the whole organism
form of supportive structure. It may be from simple rods of
from place to place is termed locomotion. Plants exhibit
strengthening material in the protozoans to the highly
cellular and often organ movement, but they do not complex skeleton of arthropods and vertebrates. Generally
locomote, that is move from place to place in search of food
speaking, the design of the supporting structure contri-
or water.
butes towards the specific shape of the organism. This, in
Whilst a few animals can survive successfully by turn, is dictated by the particular requirements of the
remaining attached to one place, the vast majority have organism concerned. Structures of different design are
complex locomotory systems which presumably evolved to needed for aquatic or terrestial animals, quadripedal or
enable them to search for and acquire food. However, even bipedal animals, and for those that move over the ground
sessile animals exhibit a great degree of mobility of their or through the air.
bodily parts. The general functions of a skeleton are as follows.
Apart from the need for food, some animals use (1) Support. All skeletons provide a rigid framework for
locomotion to avoid capture by predators. It is also used for the body and are resistant to compression. They help to
dispersal purposes and locating new favourable habitats as maintain the shape of the body. For terrestial organ-
well as for bringing together individuals for reproductive isms the skeleton supports the weight of the body
activity. against gravitational force and in many cases raises it
Locomotion is the result of the arrangement, interaction above the ground. This permits more efficient move-
and coordination of the nervous, muscular and skeletal ment over the ground. Within the body, organs are
systems. Muscles promoting locomotion are attached to attached to, and suspended, from the skeleton.
the skeleton and are therefore called skeletal muscles (2) Protection. The skeleton protects the delicate internal
(section 17.4.1). They act as machines, converting chem- organs in those organisms with an exoskeleton (arthro-
ical energy into mechanical energy. They have the ability to pods), and parts of the endoskeleton are designed for a
contract, and when they do they serve to move the systems similar function. For example, in Man the cranium
of levers comprising part of the skeleton. Coordinated protects the brain and the sense organs of sight, smell,
movement of the levers enables the animal to move about. balance and hearing; the vertebral column protects the
The posture of the animal is also maintained by the spinal cord, and the ribs and sternum protect the heart,
musculo-skeletal system which is under the overall control lungs and large blood vessels.
of the central nervous system. (3) Locomotion. Skeletons composed of rigid material
Other muscles within the body serve not to move the provide a means of attachment for the muscles of the
whole organism but to move materials from place to place body. Parts of the skeleton operate aslevers on which the
within it. Cardiac muscle (section 8.5) of the heart pumps muscles can pull. When this occurs, movement takes
blood round the body, whilst smooth muscle (section place. Soft-bodied animalsrely onmusclesacting against
17.5.3) located in the walls of various blood vessels body fluids to produce their form of locomotion.
constricts or dilates them and this alters the blood flow. Three major types of skeleton are generally recognised:
Smooth muscle in the wall of the gut propels food along the hydrostatic, exoskeleton and endoskeleton.
intestinal tract by means of peristalsis (section 1O%4r 1).
These are just a few of many such activities constantly 17.1.1 Hydrostatic skeleton
occurring within the body.
This is characteristic of soft-bodied animals.
In this chapter we will be primarily concerned with
locomotion, and two systems will be discussed in detail, Here fluid is secreted within the body and enclosed by the
body wall muscles. The fluid presses against the muscles
namely the skeletal and muscle systems. This will be
which in turn are able to contract against the fluid. The
followed by a review of the types of locomotion that occur
muscles are not attached to any structures and thus they can
in a wide variety of organisms.
625
only pull against each other. The combined effect of muscle
internal exoskeletal
contraction and fluid pressure serves to maintain the shape projection used for
and form of the animal. Generally there are two muscle muscle attachment
(apodeme)
layers, one longitudinal and the other circular in orienta-
tion. When they act antagonistically against each other extensor muscle
locomotion is effected. If the body of the organism is relaxed
flexor muscle
unsegmented (as in nematodes), when a particular muscle contracted
contracts, the pressure on the fluid will be transmitted to all
other parts of the body. If the organism is segmented (such
as Lumbricus terrestris, the earthworm) then such pressure
is localised and only certain segments will move or change
shape. A detailed account of the function of the hydrostatic
skeleton in locomotion is given for the earthworm in
section 17.6.5.
WT a Exoskeleton
This is a particular characteristic of the
arthropods. Epidermal cells secrete a non-cellular cuticle, thin, flexible articular oo e
membrane
or exoskeleton, composed mainly of chitin. It acts as a hard
outer covering to the animal and is made up ofa series of
articulated plates or tubes covering or surrounding organs. exoskeleton
It also extends into the anterior (stomodeum) and posterior

(proctodeum) ends of the alimentary canal, and into the
tracheae. Chitin is very tough, light and flexible; however,
it can be strengthened by impregnation with ‘tanned’ Fig 17.1 VS of joints and musculature of an arthropod limb
(hardened) proteins, and, particularly in the aquatic
crustaceans, by calcium carbonate. Where flexibility is
required, as at the joints between plates or tubes, chitin exoskeleton is extended and moulded into a larger form
remains unmodified. This combination of a system of plates which often incorporates a change of shape. The new
and tubes joined together by flexible membranes provides exoskeleton finally hardens. The animal is vulnerable to
both protection and mobility. predators whilst the new exoskeleton is hardening. At this
Arthropods are the only invertebrate group to possess stage the skeleton is unable to support the weight of the
jointed appendages. The joints are hinges, and the levers animal and movement is virtually impossible. This is less of
on either side are operated by flexor and extensor muscles a problem for aquatic species as their body weight is
which are attached to inward projections of the exo- supported by the water, but aquatic and terrestial organ-
skeleton (fig 17.1). Chitin is permeable to water and this isms usually hide themselves away during this time in an
could lead to desiccation of terrestial animals like insects. attempt to decrease their chances of being devoured by
This is prevented by the secretion of a thin waxy epicuticle predators. Moulting is quite expensive in terms of the
via ducts from gland cells in the epidermis (fig 4.33). energy expenditure involved in building the exoskeleton in
Therefore, the exoskeleton supports and protects the the first place and material loss when it is shed.
delicate inner parts of the animal and in addition prevents
their drying up. 17.1.3 Endoskeleton
The hollow tubular form of the exoskeleton is very This is found in the protozoan order Radio-
efficient as a supporting and locomotive device for small laria, where it consists of a skeleton of siliceous spicules,
animals, such as most arthropods, and can support a much the molluscan class Cephalopoda, where some organisms
greater weight without giving way than a solid cylindrical such as the cuttlefish possess an internal shell, and in
strut of the same mass. However, it loses this efficiency vertebrates. Typically the vertebrate skeleton is made
when organisms become bigger and their weight increases. either of cartilage or bone, is located within the organism
Here, to do the same job just as efficiently, the exoskeleton and is internal to the muscles. A further difference between
would have to increase in weight and thickness. The end this form of skeleton and the exoskeleton is that the
product would be very heavy and cumbersome. endoskeleton is composed of living tissue and so can grow
Growth takes place by ecdysis (moulting) in juvenile steadily within the animal thus avoiding the necessity for
stages (larvae and nymphs) in insects and throughout adult ecdysis. A number of different types of joint exist and
life in crustaceans. At various times the exoskeleton is shed bones that form them are maintained in their correct
(ecdysis), thus exposing a new, soft and extensible relative position by elastic ligaments. Skeletal design in
exoskeleton. Whilst still soft, growth takes place as the quadrupeds and bipeds is essentially the same, but there
626
are slight differences in mobility at the shoulder and the 17:2:2 Bone
hip. This is correlated with the type of locomotion adopted
by the animals concerned and will be discussed in detail Bone is a hard, tough connective tissue
later. composed mainly of calcified material. Details of its
histology can be found in section 8.4.4. When a vertical
longitudinal section of a long bone, such as a femur, is
17.1.4 The vertebrate skeleton examined microscopically it is seen to be made up of
several distinct components. Such a bone consists of a
The vertebrate skeleton is composed either of
hollow shaft or diaphysis, with an expanded head or
cartilage or bone. Both tissues provide an internal
epiphysis at each end. Covering the entire bone is a sheath
supporting framework for the body. Only elasmobranch
of tough connective tissue, the periosteum. The diaphysis is
fish (such as dogfish and sharks) possess a wholly
composed of compact bone whilst the epiphyses are
cartilaginous endoskeleton. All other vertebrates have a
composed of spongy bone overlain by a thin layer of
bony skeleton in their adult form, but with cartilage also
compact bone. The layout of the bony material is designed
present in certain regions, such as at the joints or between
to withstand compression forces and to give maximum
the vertebrae. In the embryo stage the skeleton of bony
strength to the bone (fig 17.2).
vertebrates is initially laid down as hyaline cartilage
Fatty yellow marrow occupies the marrow cavity of the
(section 8.4.4). This is of great biological significance, as
diaphysis, whilst red marrow is present amongst the bony
cartilage is capable of internal enlargement, and so
struts (trabeculae) of the epiphyses. Numerous small
different parts of the skeleton are able to grow in
openings penetrate the surface of the bone, through which
proportion with each other during the development of the
nerves and blood vessels traverse into the bony tissue and
organism. Bone is different from cartilage in this respect as
marrow.
it can grow only by addition of material to its outer surface.
Apart from the functions listed earlier in the chapter, a
If this were to occur during development, the articulating
bony skeleton also functions to produce red blood
surfaces of bones at joints, and their respective points of
corpuscles and white granulocytes. Further it participates
muscle attachment would be unable to retain their correct in the maintenance of constant calcium and phosphorus
spatial relationships. levels in the bloodstream (see chapter 16) by providing a
store of calcium and phosphate ions which can be mobilised
by the action of parathyroid and calcitonin hormones of the
17.2 Skeletal tissues parathyroid and thyroid glands respectively.
17.2.1 Cartilage licate for a bone Sich as the :

cle
ts structure, articular cartilage, mustion
Three types of cartilaginous material are
recognisable: hyaline, white fibrous and yellow elastic
ligaments are adapted for the func s
cartilage. A detailed account of their histology can be
found in section 8.4.4. All types consist of a firm matrix
penetrated by numerous connective tissue fibres. The
matrix is secreted by living cells called chondroblasts.
These later become housed in spaces (lacunae) scattered head (epiphysis)
throughout the matrix. In this condition the cells are of femur
termed chondrocytes. Hyaline cartilage is the most
common type and is found particularly at the ends of bones
articulating to form joints. Its matrix of chondroitin
sulphate is compressible and elastic, and is well able to
withstand heavy weight and absorb sharp mechanical
shocks such as might take place at joints. Within the matrix trabeculae (bony
struts)
are embedded fine collagen fibres which provide resistance
to tension and compression. Dense connective tissue,
called the perichondrium, surrounds the outer surface of periosteum
this cartilage at all places except where it passes into the
cavity of a joint. compact bone
White fibrous cartilage contains a dense meshwork of diaphysis
collagen fibres and is found as discs between vertebrae and marrow
as a component of tendons. It is very strong yet possesses a
degree of flexibility. Yellow elastic cartilage possesses
many yellow elastic fibres and is located in the external ear, Fig 17.2 VS of femur head showing arrangement of
trabeculae in spongy bone
epiglottis and pharyngeal cartilages.
627
17.2.3 Development of the mammalian bony increase their surface area for muscle attachment. Under-
skeleton stressed bone atrophies. This is the subject of much current
research because it occurs during prolonged weightlessness
The cartilaginous skeleton of a mammalian in space and presents problems for returning astronauts
embryo is gradually replaced by bone in a growing and cosmonauts. Bone structure may also be weakened by
organism. This involves the removal of existing cartilage, nutritional deficiencies such as lack of vitamins A and D
and construction of bone in its place, a process called and by lack of growth hormone.
ossification. In a long bone the diaphysis is the first area to
be ossified. Here a layer of membrane bone (section 8.4.4) 17.2.5 Support in vertebrates
is laid down in the perichondrium forming a complete ring
of bone around the diaphysis. When complete, this layer is Amphibia evolved from lobe-finned fish and
termed the periosteum. Internal to the periosteum, as they migrated from water to land they were faced with
chondrocytes increase in size, the matrix between them the problem of gravity and of holding their bodies off the
becomes calcified and the cells eventually disintegrate. ground in the absence of any support by the air. As a
This leads to the appearance of a series of hollow corridors consequence their vertebrae evolved to become complex
in the cartilage. They are gradually filled by embryonic structures linked together by interlocking processes.
bone marrow cells and blood vessels arising from the layer Collectively the vertebrae formed a strong but reasonably
of membrane bone in the outlying regions of the diaphysis. flexible girder that supported the weight of the body.
Some of the bone marrow cells differentiate into bone The legs of early amphibians splayed out from the sides
forming cells called osteoblasts. They position themselves of their bodies so that the animals were able to drag
around the remaining bone marrow and secrete layers of themselves over the ground. This type of stance and
bony material. The end product is a strong, hollow tube of locomotion is also seen in primitive reptiles (fig 17.3).
bone which surrounds the marrow cavity. When in motion most of the muscular energy is used to
Ossification extends towards the epiphysis, but the hold up the trunk from the ground. Such is the effort
cartilage is not completely replaced until the adult structure required to maintain this position that the animals spend
is attained. Even then, portions of hyaline cartilage remain the majority of their time whilst on land resting their bellies
at the articulating surfaces of the bone. The epiphyses on the ground.
become ossified after the diaphysis. There may be one or Later the trend in reptilian evolution was towards
several ossification centres present in each epiphysis and bringing the limbs into a position beneath the body and
they always remain separate from the diaphysis. raising the body well clear of the ground (fig 17.3b). This
Growth in length of the bone occurs at the same time as stance provides greater efficiency in locomotion and means
ossification. It takes place in growth regions towards each that the weight of the body is transmitted through the four
end of the bone between the epiphyses and diaphysis. Until relatively straight limbs.
the adult form is reached, cartilage continues to be Some reptiles and mammals have evolved a bipedal gait,
produced and ossified into new bone on either side of these walking, running or hopping on their hindlimbs. This
regions. Ultimately growing regions become ossified. releases the forelimbs for developing manipulative skills
When this happens, no further cell division occurs, and such as feeding, building and cleaning. A special type of
growth in length ceases. Increase in girth of the bone is locomotion, called brachiation characterises some mon-
produced by further deposition of bone in the periosteum. keys and apes. These animals swing from tree to tree using
17.2.4 Factors controlling bone deposition

Even after growth is complete, bone absorp-
rK
tion, carried out by cells called osteoclasts, and bone
synthesis and reconstruction still occur. This is important as
it permits the shape of the bones to be modified according
to the mechanical stresses placed upon them. It is of
particular significance in animals that exhibit a marked
change in their mode of locomotion during their develop-
ment into adults. This also generally incorporates a shift in
the position of the load that their skeletons have to bear. If
continuous pressure is exerted on a specific region of a
bone, then that region is absorbed. However, if part of a
(6) (c)
bone is subjected to periodic stress, bone deposition is
stimulated in that region. Such pressures and stresses Fig 17.3 Types of stance in vertebrates:
mould the skeleton into its definitive shape, and in (a) a primitive amphibian stance — legs projected laterally from
body and then down; (ob) modern reptilian stance —
particular, intermittent stresses are responsible for the intermediate between amphibian and mammals; (c) mammalian
development of projections and ridges on bones which stance — legs project straight down from beneath the body
628
their long arms and elongate hands to grasp the branches. Table 17.1. Number and types of vertebrae in a range of
Other animals that climb and move about in trees are too mammals
small to brachiate; instead they jump from branch to
branch. The most specialised form of aerial locomotion is
Types of Region Number of vertebrae
true flight. This evolved simultaneously during the Jurassic vertebra
period in the flying reptiles (pterodactyls) and in the first rat rabbit cat cow Man
birds (which were descended from reptiles). The forelimbs
were modified and adapted into wings. Flying reptiles cervical neck of, 7 7 i i
thoracic chest 13 12-13 13 13 12
eventually became extinct, but birds survived and evolved lumbar abdomen 6 6-7 7 6 5
into many highly varied forms. sacral hip 4 4 3 5 5
caudal tail 30+ 16 18-25 18-20 4
17.3 Anatomy of the skeleton of a

mammal (the rabbit)
All mammalian skeletons possess the same tion to the spinal cord. On the vertebrae are numerous
basic divisions. These can be divided into two main groups: projections for the attachment of muscles. When the
the axial skeleton, which consists of the skull, vertebral muscles are active, they may bend the vertebral column
column, ribs and sternum, and the appendicular skeleton, ventrally, dorsally or from side to side.
which consists of an anterior pectoral and posterior pelvic The total number of vertebrae varies in different
girdle, attached to each of which is a pair of limbs. mammals. Nevertheless, in all mammals five regions of the
vertebral column can be distinguished. The number and
17.3.1 The axial skeleton types of vertebrae in a variety of mammals are given in
table 17.1.
The skull consists of the cranium to which the
Vertebrae from different regions of the vertebral column
upper jaw is fused, and a lower jaw which articulates with
all conform to the same basic design. The structure of a
the cranium. Muscles connect the lower jaw to the skull and
typical vertebra is shown in fig 17.4. Note that two facets
cranium. The cranium is composed of a number of
(articulating surfaces) called prezygapophyses are present
flattened bones tightly interlocking forming a series of
at the anterior end of the vertebra, whilst two more, the
immovable joints. Besides enclosing and protecting the
postzygapophyses occur at the posterior end. The prezy-
brain, it protects the olfactory organs, middle and inner ear
gapophyses of one vertebra fit against the postzygapo-
and the eyes. At the posterior end of the cranium are two
physes of the vertebra immediately anterior to it. This
smooth, rounded protuberances, the occipital condyles,
arrangement enables the vertebrae to articulate with each
that articulate with the atlas vertebra to form a hinge joint
other, but it is not a completely rigid arrangement, for the
which permits the nodding of the head.
smoothness of the articulating surfaces permits their slight
The vertebral column is the main axis of the body. It movement over each other. Below each pre- and post-
consists of a linear series of bones called vertebrae, placed zygapophysis is a small notch. When adjacent vertebrae are
end to end, and separated by cartilaginous intervertebral fixed closely together the anterior notch of one vertebra is
discs (fig 17.5). The vertebrae are held together by placed against the posterior notch of the vertebra im-
ligaments which prevent their dislocation, but permit a mediately in front of it. This arrangement forms a hole
degree of movement, so that the vertebral column as a through which a spinal nerve can pass. Other structures
whole is flexible. The vertebral column also gives protec- characteristic of all vertebrae are the neural spine and
transverse processes for muscle attachment. The centrum
forms a central rigid body to the vertebra over which the
neural arch encloses the spinal cord.
Whilst there is a great degree of similarity between
neural spine
vertebrae, their design varies in different regions of the
neural arch vertebral column. This is because of uneven distribution of
SS process body weight along the length of the column, and the
vertebrae are modified and adapted to perform those
specific functions required by each region.
neural canal
prezygapophysis When a rabbit stands up, its vertebral column is
supported by the fore- and hindlimbs, with the bulk of the
body weight suspended between them. The centra of the
vertebrae withstand compression whilst ligaments and
muscles which overlay the dorsal parts of the vertebrae
Anterior view of a typical mammalian vertebra
withstand tension (fig 17.5)
Fig 17.4
629
as compression members w hilst the
Fiqi 17.5 Skeleton of rabbit seen from left side. The centra of the vertebrae serve
The abdominal musculature prevents the
ieaione and muscles which link one vertebra to another form the tension members.
weight of the body from forcing the girdles apart
upthrust of upthrust 0 f
forelimb hindlimb
skull
i J —_ at ENSION ay
cervical ;
) thoracic 7
Ye vertebrae vertebrae *Nsy,,
% |
1 lumbar
iver \ Ayd YQ vertebrae
Sues WYER
NySOS} Bay
, 9
weight ae floating a
ribs
ee eee i) |
Da ie me
ee / Con vertebrae
1ON : sess peer
head ars. rt STon
humerus
h rn vy vertebrae
wu ulna tibia
pubis
sternum
radius fibula ischium
weight
carpals tarsals
metatarsals
metacarpals
phalanges weight
(a) neural spine
17.3.2 Structure and functions of the ver-

prezygapophysis \ neural arch
tebrae of a rabbit
Cervical vertebrae
neural canal vertebrarterial
Cervicals 3-7 are very similar in structure (fig 17.6a). They canal
possess a small centrum which is able to withstand
transverse
compressional forces, and a short neural spine to which the process
neck muscles are attached. Some of these muscles run from
the cervicals to the thoracic vertebrae and are used for cervical rib centrum
holding up the neck, whilst others run to the back of the (b) neural spine
skull and serve to maintain the head in position. On each aN neural arch
neural canal aTveIse
side of the centrum is a single hole, the vertebrarterial
process
canal, formed by fusion of a cervical rib with the transverse So) _svertebrarterial
process. As its name implies, it serves as a channel for the canal
vertebral artery to pass through to the brain. Thus this
important blood vessel is protected as it traverses the anterior facet
for articulation
vulnerable region of the neck. ligament with occipital
condyles of
skull
(°C)
neural spine
postzygapophysis
Fig 17.6 (right) (a) Fifth cervical vertebra of a rabbit, anterior odontoid cervical rib
process z
view. Note the characteristic vertebrarterial canal; (b) Anterior
a centrum
view of atlas vertebra of a rabbit. Note absence of centrum articular P
and anterior facets; (c) Side view (left) of axis vertebra of a surfaces for vertebrarterial
rabbit. Note odontoid process and forwardly projecting neural atlas canal
spine <+—— anterior posterior ———>
630
The first two cervical vertebrae possess a quite different Fig 17.8 (a) Left side view of thoracic vertebra of a rabbit.
design and are modified to support the head and enable it Note long neural spine and demi-facets. (bo) Lumbar vertebra
to move in various directions. The first cervical vertebra is of a rabbit from left side. No hypapophysis is shown. Where it
does occur (first and second lumbar vertebrae) it exists as a
the atlas (fig 17.6b). Zygapophyses and a centrum are small projection from the ventral surface of the centrum. (c)
absent and the neural spine is reduced to a very small crest. Dorsal view of sacrum of a rabbit
On its anterior surface are two concave depressions, the
articular facets which articulate with the curved convex (a)
occipital condyles of the skull to form a hinge joint. This tubercular facet neural spine
supports the skull and permits it to be nodded up and
transverse
down. Wide, flattened transverse processes provide a large process © S“<—~
surface area for the attachment of those muscles that bring
about the nodding action. prezygapophysis
postzygapophysis
The second cervical vertebra is the axis (fig 17.6c). It
possesses a peg-like structure called the odontoid process capitular
demi-facet capitular
demi-facet
which projects forwards from the centrum. The process is
formed by the fusion of the centrum of the atlas to that of
the axis, and it fits into the cavity of the atlas below the
ligament (fig 17.6b) thus being separated from the neural <— anterior centrum
canal. This arrangement gives a pivot joint which enables
the head to be rotated from one side to the other (that is to
(b) neural spine
be shaken). Such activity is brought about by muscles on
the left and right sides of the neck. They run forwards from metapophysis postzygapophysis
the neural spine of the axis to attach to the transverse
processes of the atlas (fig 17.7). No prezygapophyses are anapophysis
present.
<———-anterior centrum
neck muscles
transverse
axis process
atlas
vertebrarterial
odontoid process (c) anterior
canal
prezygapophysis
Fig 17.7 The arrangement of the neck muscles between the
atlas and axis vertebrae in a rabbit transverse
‘ :
rocess
E region fused with
pelvic girdle
neural spines of
four fused
Thoracic vertebrae vertebrae
These possess long, backwardly pointing neural spines and
short transverse processes. Also present on the transverse
processes are small, rounded pro jections called tubercular
facets. Anterior and posterior half or demi-facets are
located on the sides of the centrum. Both types of facet are
for articulation with the ribs (fig 17.8a). The end of the rib
which joins to a thoracic vertebra branches into two vertebra in front of it to form a common depression. The
projections, one being called the capitulum and the other capitulum of the rib fits into this depression thus effectively
articulating with two vertebrae. As a result the thoracic
the tuberculum. The tuberculum articulates with the facet
vertebrae serve to support the ribs, but because of the
of the transverse process whilst the capitulum articulates
with two demi-facets of the centrum. Here the arrange-
complex arrangement between the vertebrae and ribs,
movement between them is strictly limited. Some forward
ment is quite complex. When two thoracic vertebrae are
and sideways movement can occur, but in general the
closely applied to each other, the anterior demi-facet of
thoracic vertebrae are the least flexible of all.
one vertebra fits closely to the posterior demi-facet of the
631
Lumbar vertebrae Fig 17.9 Anterior view of thoracic vertebra of a rabbit
attached to a pair of ribs neural spine
The vertebrae of this region are subject to the greatest
process of rib prezygapophysis
stress in terms of gravity and locomotion. Not only must for ligament \
they provide rigidity for the body, but they must also attachment
transverse
permit bending, sideways movement and rotation of the tubercle process
trunk. Therefore, not surprisingly this is the region where
the large muscles of the back are attached and where there capitulum of rib
u
centrtrum
are many adaptive modifications of the vertebrae. The
centrum and neural arch are massive, although the centrum rib
is quite short. This arrangement provides greater flexibility
between the lumbar vertebrae. The transverse processes
‘ sternum
are long and wide. They point forwards and downwards. costal cartilage
Extra muscle bearing projections called meta-, ana-, and
hypapophyses are present on the vertebrae (fig 17.8D).
They also interlock with each other and keep the vertebrae 17.3.3 The appendicular skeleton
in their correct positions relative to each other when this Limb girdles
part of the vertebral column is placed under stress.
These provide a connection between the axial skeleton and
the limbs. The width of the pectoral girdle separates the
Sacral vertebrae (sacrum) forelimbs, and that of the pelvic girdle the hindlimbs, and
The sacral vertebrae are fused together to form a broad both contribute to the stability of the animal. A number of
structure, the sacrum (fig 17.8c). The most anterior sacral areas are modified for muscle attachment and articulation
vertebrae possess well-developed transverse processes with the limb bones.
which are fused to the pelvic girdle. It is through the sacrum Pectoral girdle. This is composed of two
that the weight of the body of a stationary animal is
distinctly separate halves. Each half consists of the scapula,
transmitted to the pelvic girdle and the legs. When an
coracoid process and clavicle. It is not fused to the axial
animal moves forwards, the thrust developed by the
skeleton but flexibly attached to it by ligaments and
hindlimbs is transmitted via the pelvic girdle through the
muscles. This arrangement enables the girdle and its
sacrum to the rest of the axial skeleton.
associated limbs to be moved through a great variety of
planes of movement and angles. The girdle is strong
Caudal vertebrae enough to support the majority of the weight of a
The number of caudal vertebrae varies greatly from one quadruped when it is stationary. It also acts as a shock
mammal to another (table 17.1) and is related to different absorber when the animal lands at the end of a jump.
lengths of tails in such mammals. In general, as they pass The scapula is a flat, triangular-shaped bone which
towards the posterior end of the animal, transverse overlies a number of the anterior ribs (fig 17.10a). At its
processes, neural arches and zygapophyses all become apex is a concave depression, the glenoid cavity, which
reduced in size and gradually disappear. This results in the articulates with the head of the humerus to form a
terminal vertebrae only consisting of small centra. Man ball-and-socket joint. A spine runs along the outer surface
possesses four caudal vertebrae which are fused to form the of the scapula, and at its free end, close to the glenoid
coccyx. It is not visible externally. cavity are two projections, the acromion and metacromion
which are both used for muscle attachment. The coracoid
Ribs and sternum
process is all that remains of a small bone, the coracoid,
which has fused with the scapula to form a projection above
Each rib is a flattened, curved bone. Its dorsal end is forked the glenoid cavity.
into the capitulum and tubercle which provide points of The clavicle is variable in size and shape in different
articulation with the thoracic vertebrae. The joints formed mammals. In Man it is well developed with one end
permit movement of the ribs by the intercostal muscles articulating with the acromion process and the other with
during breathing. All of the ribs, thoracic vertebrae and the the sternum. It is used for muscle attachment and aiding the
sternum form a thoracic cage which protects the heart, complex movements of the arms. It is sometimes referred
lungs and major blood vessels (fig 17.9). to as the collar bone in Man. Its removal has no serious
In the rabbit the ventral ends of the first seven pairs of consequences. In quadrupeds it is much smaller and
ribs are attached to the sternum, a flattened, kite-shaped relatively less important. It forms the ‘wishbone’ in birds.
bone, via costal cartilages. These are called true ribs. The
next two pairs of ribs are also attached ventrally to the 17.2 What advantages are there tomam-
cartilage of the seventh rib. The ventral ends of the mals in possessing a flexible connection between ee
remaining three or four pairs of ribs are unattached and are pee we and vertebral column?
called floating ribs (fig 17.5).
632
Fig 17.10 (a) Left scapula of a rabbit. (b) Ventral view of Fig 17.11 Vertebrate pentadacty! limb
pelvic girdle of a rabbit. Note how sacrum is fused to the ilium
FORELIMB ;
(a) spine
humerus femur
coracoid
process
glenoid cavity supra-scapula

tibia
radius fibula
acromion
ulna
metacromion nr
cpa —____{ a = =
WRIST ee ANKLE
metacarpals__ff I [ [ metatarsals
(0) ilium
PALM FOOT
digits - s 4 LH i - digits
sacrum
FINGERS B&BS
a ar | B TOES
(phalange — name given to each bone of a digit)
acetabulum
Forelimb. The upper part of the forelimb
consists of a single bone, the humerus. At its upper end is
obturator
pubis foramen (hole) the head which articulates with the glenoid cavity of the
ee pubic scapula to form a ball-and-socket joint at the shoulder
symphysis allowing universal movement. Near the head are two
(line of fusion) roughened projections, the greater and lesser tuberosities,
between which is a groove, the bicipital groove. It is along
this groove that the tendon of the biceps muscle passes. At
Pelvic girdle. Again this consists of two its lower end is the trochlea which articulates with the
halves, each half comprising three bones, the ilium, forearm to form a hinge joint at the elbow. A hole, the
ischium and pubis. They are fused to each other forming a supra trochlear foramen, perforates the humerus just above
single structure, the innominate bone. The ilium is fused to the trochlea in the rabbit, but is absent in Man. Also visible
the sacrum of the vertebral column on each side. On the is the characteristic deltoid ridge running anteriorly along
outer edge of each half is a depression, the acetabulum, the upper half of the humerus (fig 17.12a). The lower part
which articulates with the head of the femur to form the of the forelimb, the forearm, is composed of two bones, the
ball-and-socket hip joint (fig 17.10b). The ilium is above ulna and radius. The ulna is the longer of the two. A notch,
the acetabulum. Dorsally it possesses a large crest to which the sigmoid notch at its upper end articulates with the
the thigh muscles are attached. trochlea of the humerus. Beyond the elbow joint is a
Between the ischium and pubis is a large hole, the projection, the olecranon process. This is a most important
obturator foramen. Except for a small aperture through structure, for when the arm is straightened it prevents any
backward movement of the forearm; hence
which blood vessels and nerves pass to the legs, it is covered further
by a sheet of tough inflexible connective tissue which dislocation does not occur. On the anterior surface of the
provides yet another surface for muscle attachment. Sucha humerus, above the trochlea, is a hollow, the supra
design could be an adaptation to reduce the weight of the trochlear fossa, into which the radius fits when the arm is
pelvic girdle and so lighten the load that has to be bent (fig 17.125).
supported by the hindlegs. The radius is a flattened, slightly curved bone which is
Ventrally a line of fusion can be seen where the two relatively simple in design. In Manitisnot firmly bound to the
halves of the pelvic girdle meet. This is the pubic ulna; muscles are able to rotate the radius about the ulna so
symphysis. Flexible cartilage in this region permits a that the palm of the hand can be turned downwards or
widening of the female’s girdle at the time of giving birth. upwards, contributing to Man’s manipulative skills. This
freedom of movement is not apparent in the rabbit where
Limbs both bones are tightly bound and the palm always faces
The limbs of all mammals are designed on the same basic downwards. However, this is not disadvantageous as the
plan, that of the pentadactyl limb, so named because each limb is in the best position for burrowing and running.
limb terminates in five digits (fingers or toes) (fig 17.11). Distally the ulna and radius articulate with a number of small
There are numerous variations of the general plan, which carpal bones which form the wrist. In turn the carpals articu-
late with five long metacarpals which finally articulate with
are adaptations to the different modes of life of different
five digits. The first digit, on the inside of the limb is com-
animals. In some cases the number of digits per limb has
posed of two phalanges whereas all others contain three.
been reduced during evolution (section 24.7).
633
Fig 17.12 (a) Left humerus of a rabbit, anterior and posterior with the tibia to form a hinge joint at the knee. A patella
views. (b) Ulna and radius of a rabbit, side view groove separates the two condyles. The patella bone (knee
cap) is located here.
Co
bicipital
lesser
tuberosity
The tibia and fibula bones form the shank of the
5
groove
ee
(projection) greater hindlimb. Two slight depressions at the upper end of the
; tuberosity tibia represent the articular surfaces at the knee joint (fig
(projection)
deltoid
ridge 17.13b). The fibula is not a component of this joint. It is a
supratrochlear head
thin bone, and in the rabbit is fused to the tibia at its lower
foramen (hole) end. At the lower ends of the tibia and fibula are a number
olecranon of tarsal bones. The two longest tarsals, the astragulus and
trochlea fossa calcaneum (heel bone), articulate with the tibia and fibula
anterior posterior to form the ankle joint. The tarsals articulate distally with
view view
long metatarsal bones to form the foot, whilst in turn the
metatarsals articulate with digits composed of phalanges
(b) radius sigmoid
notch forming the toes. It is interesting to note that the rabbit
hindlimb possesses only four digits.
17.3.4 Joints
distal surface In bony vertebrates, where a bone meets
articulating
with wrist bones another bone, or bones, a joint is formed. Movement of
skeletal elements over each other is only possible if there is
olecranon a joint between them. A variety of different types of joint
ulna process exist in the mammalian skeleton. They are summarised in
table 17.2.
Hindlimb. The upper part of the hindlimb Synovial joints are essentially similar to each other in
consists of a single bone, the femur. At its upper end is a design. The end surface of each articulating bone is
large round head which articulates with the acetabulum of overlain by a smooth covering of hyaline cartilage. Though
the pelvic girdle to form a ball-and-socket joint at the hip a living tissue, it contains no blood vessels or nerves. The
(fig 17.13a)). Three processes called trochanters protrude nutrients and respiratory gases it requires diffuse from the
below the head and provide points of attachment for the synovial membrane and fluid. The cartilage serves to
thigh muscles. The lower end of the femur possesses two reduce friction between the bones during movement.
curved convex surfaces, called condyles, which articulate Because of its elastic properties, the cartilage also acts asa
shock absorber.
great trochanter
The bones of the joint are held in position by a number of
ligaments which collectively form a strong fibrous capsule.
third trochanter They run from one side of the joint to the other and are
orientated in such a way as to cope effectively with the
particular stresses suffered by the joint. The inner surface
trochanter
of the capsule is lined by a thin, cellular synovial membrane
which secretes synovial fluid into the synovial cavity (fig
rounded 17.4). Synovial fluid, containing mucin, acts as a lubricant
articulating
surfaces patella groove
(condyles) for
tibia
spongy bone ;
(b) periosteum
cnemial crest
tibia compact bone

synovial cavity
\for fibrous capsule
Drie iseia at containing
synovial fluid
|
hyaline == ===
cartilage
es EG (
synovial
membrane
articulating
surface for ankle
17.13 (a) Left femur of a rabbit, anterior view. (0) Anterior

view of left tibia and fibula of a rabbit Fig 17.14 Diarthrodial/synovial joint of a mammal
634
Table 17.2. A variety of joints in the endoskeleton of a mammal
————
et
—=——eee——eee——ooooooooooooeoeleeaeoeaeaeaeaeaeaeaeaeaeaeaeaeaae
Type of joint General characteristics Examples Function

Immovable/suture/ A thin layer of fibrous connective Between bones of Provides strength and support for the
synarthrodial tissue exists between the bones, holding skull; between body, or protection of delicate
them firmly in position sacrum and ilia of structures which cannot withstand
pelvic girdle; any kind of deformation
between bones of
pelvic girdle
Partially movable/ Bones are separated from
amphiarthrodial each other by cartilaginous
' pads
(a) Gliding Joints between Bones glide over each other to a
vertebrae; wrist limited extent. Collectively they
and ankle bones provide a wide range of movement
and confer strength on the limb.
(b) Swivel/rotating/ Joint between Permits shaking of head from
pivot atlas and axis side to side
vertebrae
Freely movable/synovial/ Articulating bone surfaces are
diarthrodial covered with cartilage and separated from
(fig 17.14) each other by a synovial cavity containing
synovial fluid
(a) Hinge Relatively few muscles operate this joint Elbow, knee and Permits movement in one plane about
finger joints one axis. Capable of bearing
heavy loads
(b) Ball and socket Variety of muscles attached to the bones Shoulder and hip Permits movement in all planes, and
of the joint joints some rotation. Unable to bear very
heavy loads
for the joint surfaces and serves to reduce friction between It contracts slowly and fatigues slowly. It is spon-
them. The synovial membrane acts as a waterproof seal taneously active and innervated by the autonomic
preventing escape of synovial fluid. Therefore the joint nervous system.
effectively requires no maintenance. (3) Cardiac muscle. Muscle found only in the heart. It
contracts spontaneously and without fatigue. It is
innervated by the autonomic nervous system.
17.4 The muscle system
17.4.1 Skeletal muscle in detail
Muscles are composed of many elongated cells
called muscle fibres which are all able to contract and relax. A skeletal muscle is attached to bone in at
During relaxation they are capable of being stretched, but least two places, namely the origin, a firm non-movable part
they exhibit the property of elasticity which permits them of the skeleton, and the insertion, a freely movable part of
to regain their original size and form after being stretched. the skeleton. Attachment is by means of tough, relatively
Muscles are well supplied with blood which conveys to inextensible tendons made up of connective tissue com-
them nutrients and oxygen, and takes away metabolic prised almost entirely of collagen (section 5.5). At one end
waste products. The amount of blood arriving at a muscle a tendon is continuous with the outer covering of the
at any one time is able to be adjusted according to its need. muscle, while the other end combines with the periosteum
Each muscle possesses its own nerve supply. Histologically of the bone to form a very firm attachment.
three distinct types of vertebrate muscle can be identified. As muscles can only produce a shortening force (that is
contract), it follows that at least two muscles or sets of
(1) Skeletal muscle (section 17.4.1) (also called striated,
striped, voluntary). Muscle which is attached to bone. muscles must be used to move a bone into one position and
It is concerned with locomotion, contracts quickly and back again. Pairs of muscles acting in this way are termed
fatigues quickly. It is innervated by the voluntary antagonistic muscles and they may be classified according
nervous system. to the type of movement they bring about (table 17.3).
It is rare that a movement will involve a single pair of
(2) Smooth muscle (section 17.5.3) (also called unstriated,
muscles. Generally, groups of muscles work
unstriped, plain, involuntary). Muscle which is found antagonistic
in the walls of tubular organs of the body and is together to produce a particular individual movement, and
concerned with movement of materials through them. such groups are known as synergists.
635
Table 17.3. Types of movement brought about by pairs the muscle to skeletal elements. Each muscle fibre is
of antagonistic muscles enclosed by a membrane, the sarcolemma. This is very
ee
————————————
SSS
———————ooeqeq=$?$eoesSsSsSsSS\<woo similar in structure to a typical plasma membrane.
Muscle Type of movement brought about Within the muscle fibres are numerous thin myofibrils
classification (myo, muscle) which possess characteristic cross striations.
Each myofibril is composed of two types of proteinaceous
Flexor Bends a limb by pulling two skeletal elements
towards each other myofilaments, actin and myosin. Numerous mitochondria
are interposed between the myofibrils. The cytoplasm of
Extensor Extends a limb by pulling two skeletal elements
away from each other the myofibril is called sarcoplasm and contains a network of
Adductor Pulls a limb towards the central long axis of internal membranes termed the sarcoplasmic reticulum.
the body Running transversely across the fibre and between fibrils is
Abductor Pulls a limb away from the central long axis of a system of tubules known as the T system, which is in
the body contact with the surface of the sarcolemma (fig 17.15). At
Protractor Pulls distal part of a limb forwards certain points the T tubules pass between pairs of vesicles
Retractor Pulls distal part of a limb backwards which are components of the sarcoplasmic reticulum. A
Rotator Rotates whole or part of a limb at one of its T tubule together with a pair of vesicles is called a triad. The
joints tubule and vesicles are held together by membranous
cross-bridges. The vesicles are involved in the uptake and
release of Ca’* ions. Their activity raises or lowers Ca”* ion
concentration in the sarcoplasm, which in turn controls
17.4.2 Striated muscle ATPase activity and hence the contractile behaviour of the
A striated muscle consists of numerous phy- muscle fibre.
siological units called muscle fibres or muscle cells. They Under a light microscope only the striated nature of the
are cylindrical in shape and arranged parallel to each other. myofibrils can be observed. This is seen as a regular
They are between 0.01 and 0.1 mm in diameter, several alternation of light and dark bands called the I and A bands
centimetres long and multinucleate. The nuclei are located respectively, traversed by thin, dark lines. Electron
near the surface of each fibre. Bundles of muscle fibres are microscope studies clearly indicate that the bands are due
enclosed by collagen fibres and connective tissue. Collagen to the regular arrangement of actin (thin filaments) and
also occurs between fibres. At the ends of the muscle the myosin (thick filaments). Fig 17.16 shows this clearly.
collagen and connective tissue forms tendons which attach Traversing the middle of each I band is a dark line called
myofibrils
transverse tubule
glycogen granules
longitudinal tubules of
sarcoplasmic reticulum
outer vesicles of
a =n sarcoplasmic reticulum
‘terminal cisternae’
CHIILLTUIL
VMI
Fig 17.15 Sarcoplasmic reticulum and T system.
636
Fig 17.16 Fine structure of skeletal muscle
(a) muscle
tendon
IA
(b) muscle fib eS striations
visible
myofibril_ gq
p fcad Whee ee |
a]
nucleus
pea
(c) three ——— a Fig 17.17 Longitudinal section of fish muscle (roach —
sarcomeres =) = Rutilus rutilus). Note the triads and clear myofibrilar structure
= ————
froma (X< 7650)
myofibril Z L
(d) myosin H zone band actin

(thick filament) Ld _(thin filament)
A band
+ tropomyosin
e * —*o + troponin
Fie —e
, )|F-actin
myosin rod myosin G-actin
(each myosin heads
has two heads)
(f) transverse through through through overlap
sections through H zone I band region
sarcomere to
illustrate cet Oo 9o
arrangement of
Oo Oo
5 0000
o.0°o"*o
Diets ie
OR One "010
Fig 17.18 Insect flight muscle (giant water bug). Transverse
filaments in a Qo © 0 ©
°
00°00 oe
Sven
section of fibril in rigor state (x 137 000).
© 6 0 e*s eake
o* oo
eae tennis
myofibril
the Z line. The section of a myofibril between two Z lines is sides of the filament except in a short region halfway along
called a sarcomere. From the Z line actin filaments extend its length. This central portion of the filament, which
in both directions, whilst in the centre of the sarcomere are possesses no heads, is termed the bare zone. Where the
found myosin filaments. These are aligned side by side in a actin and myosin filaments overlap the myosin heads can
hexagonal lattice (fig 17.17 & 17.18). In certain regions of attach to neighbouring actin filaments, and while attached
the sarcomere, actin and myosin filaments overlap. Where they can generate the force which may cause the muscle to
they do, transverse sections in these regions indicate that shorten. The energy for this force production is derived
six actin filaments surround each myosin filament. This from ATP hydrolysis since each myosin head is capable of
arrangement of actin and myosin filaments results in a ATPase activity. As described later, the attachment of
number of other bands being recognisable in the sarco- myosin heads to actin is controlled by the level of Ca** ions
mere. Myosin and actin filaments constitute the A band in the sarcoplasm. The myosin ATPase is activated by
whilst actin filaments alone constitute the I band. The attachment of myosin to actin. This activity may be
centre of the A band is lighter than its other regions in a inhibited by Mg’* ions. The importance of this will become
relaxed sarcomere as there is no overlap between actin and evident when we deal with the actual contraction mechan-
myosin in this region. It is called the H band. The H band ism of the sarcomere.
itself may be bisected by a dark line, the M line. The M line Actin (thin filaments)
joins adjacent myosin filaments together at a point halfway
Each actin filament is made up of two helical strands of
along their length.
globular actin molecules (G-actin) which twist round each
Myosin (thick filaments) other. The whole assembly of actin molecules is called
A molecule of myosin consists of two distinct regions, a F-actin (fibrous actin). It is thought that an ATP molecule
long rod-shaped region (myosin rod) on one end of which is is attached to each molecule of G-actin. Neither form of
a globular region. This globular region consists of two actin exhibits any ATPase activity. Actin filaments consist
similar globular parts, each called a myosin head. The of F-actin together with two accessory proteins, tropomy-
globular heads are regularly spaced and project from the osin and troponin. Tropomyosin is a rod-shaped fibrous
637
17.4.4 Mode of action of vertebrate skeletal
muscle
When a muscle is stimulated it exhibits
mechanical activity (that is it contracts) and this may either
produce a shortening of the muscle, or if the muscle is fixed
rigidly at both ends, it may develop tension within the
muscle without the muscle changing length. When a muscle
shortens against a constant load, this is called isotonic
contraction, but when there is no change in length, it is
isometric contraction.
Investigation into the nature of the contractile response
of a muscle can be carried out by the use of a kymograph.
The gastrocnemius muscle of a frog is often used for such
investigations. Muscle is excised from afreshly killed frog,
Fig 17.19 /llustration of placed in a trough on a platform and bathed in a
changes in actin filament well-oxygenated saline solution. Under these conditions it
structure
(a) ‘off’ state — low Ca?* may perform its contractile activity adequately for several
level: tropomyosin blocks hours. The origin of the muscle is fixed to a rigid station on
myosin attachment site. (ob) the platform, whilst its insertion is hooked up to a movable
‘on’ state — high Ca?* lever. It is then suitably stimulated by an electric current
a
level: tropomyosin moves either directly or via a nerve. When the muscle contracts or
to expose attachment sites
(arrows). A, actin; T,
develops tension it moves the lever, and such movement is
tropomyosin, Troponin, recorded as a trace on the revolving drum of the
which is not shown, lies kymograph. The record of events that take place is called a
Fi nearer to ‘grey’ actins. myogram. The tension developed by a muscle is a force and
(a) (b) is generally measured in terms of grams mass.
protein and these rods link end to end to form two helical
strands which are wrapped around the F-actin in a 17.4.5 Contractile response
longitudinal fashion. Tropomyosin functions to switch on, Single stimulus. When asingle stimulus is
or off, the contractile mechanism. Troponin is a globular applied to a muscle, there is a very short period of about
protein with three subunits. Each subunit has a particular 0.05 s, called the latent period, before the muscle responds.
function. Troponin-T binds troponin to tropomyosin, Then contraction takes place rapidly and a force is
troponin-C is sensitive to, and can reversibly bind to, Ca developed. This phase of contraction lasts for about 0.1 s.
ions, whilst under certain conditions troponin-I is able to Following this is a longer period of relaxation where the
inhibit any interaction between actin and myosin. Collec- force declines and the muscle returns to its relaxed state
tively both of these accessory proteins serve to inhibit the over a period of 0.2 s. A single contraction by a muscle is
actin-myosin interaction in the absence of Ca’ ions (fig called a muscle twitch (fig 17.20).
17.19).
Two stimulations. If a long interval of time
17.4.3 The ‘all-or-nothing’ response elapses before a second stimulus is applied, two identical
myograms for the muscle are recorded. However, if the
When askeletal muscle fibre is stimulated by time between two stimuli is reduced so that the second
an impulse, the fibre will only contract if the stimulus is at stimulus is applied when the muscle is still contracting in
or above a certain threshold level (resting potential). This response to the first stimulus, a second contraction occurs
contraction is maximal for any given set of conditions, and
even if the strength of stimulus is considerably increased phase of phase of
contraction relaxation
there will be no increase in the shortening of the muscle or
the force that it develops. This phenomenon is referred to
e
as the muscle’s ‘all-or-nothing’ response. A stimulus that is &
op
too weak to provoke muscle fibre contraction is referred to SS
cS
as a subliminal stimulus. 2
5 latent
After response, the muscle endures an absolute refrac- fe period
tory period when no contraction is possible. This is
\ I
succeeded by a relative refractory period. Only strong 0 ‘ 0.1 0.2 0.3 0.4
stimuli can provoke a response during this time. The STIMULUS Time/s
refractory period is the time it takes for ionic activity to Fig 17.20 Single muscle twitch of frog gastrocnemius
return the muscle to its resting potential. muscle
638
together (or summate). A smooth trace is drawn which
reaches a steady level or plateau and remains there for a
relatively long time. When the muscle is in this condition it
r oa
is said to be in the state of tetanus. The plateau of tension
ee _
A
developed during tetany is the maximum tension that the
(b)
muscle can produce. Tetany cannot continue indefinitely as
L
the muscle becomes fatigued.
17.4.6 The sliding filament theory of muscle

contraction
f ()
In 1954, two independent research groups,
namely H. E. Huxley & J. Hanson, and A. F. Huxley &
A R. Niedergerke, formulated the sliding filament theory of
L muscle contraction. They discovered independently that
(d) the A band of a sarcomere always remained the same
i"
length whether the sarcomere was stretched or shortened.
This gave rise to the suggestion that there are two
A interdigitating sets of filaments, actin and myosin, which in
A moment of
Time/0.1s intervals stimulation some way slide past each other when the sarcomere
changes its length. Observations indicated that during
Fig 17.21 Tracings recorded on a kymograph using frog
contraction the actin filaments move inwards towards the
gastrocnemius muscle.
(a) Single twitch in response to single stimulus centre of the sarcomere (fig 17.23). The heads of the
(ob) | Mechanical summation occurring myosin filaments were thought to operate as ‘hooks’
(c) 4 when frequency of stimulation attaching to F-actin in a particular way to form cross-
(d) | is increased bridges, and then changing their relative configuration such
that the actin molecules were pulled further into the A
band. After the process was completed, the myosin heads
detached from the actin and hooked up to another site
further along the actin filament. A sarcomere could
contract up to 30% of its length, and the cross-bridge
attachment/detachment cycles could be repeated many
times depending on the speed of shortening. The energy
required for this process was provided by the splitting of
ATP, one ATP molecule being split for each cross-bridge
cycle.
I A
Far
myosin
—
Z M : Z
actin
Time intervals 0.5s
Fig 17.22 Myogram indicating development of tetany in frog

gastrocnemius muscle
(a) Unfused tetany, 8 stimulations per 0.5s
(b) Fused tetany, 18 stimulations per 0.5s
y’
which is superimposed on the first. This results in a ‘bump
ops
myogram (fig 17.21). The second contraction also devel
is called
a greater force than the first one. This effect
HEHEHE
mechanical summation. HEHE HH
Frequent stimulations. If the frequency of
ram is Diagrammatic representation of how a sarcomere
stimulation is increased, the bumpiness of the myog Fig 17.23
hes fuse contracts by the actin filaments sliding over myosin filaments
gradually lost (fig 17.22) and the individual twitc
639
(a)| actin filament G-actin
Today the theory is almost universally accepted. How-
ever, the actual force-generating process, often called the |
excitation—contraction-coupling mechanism, is still little |
] myosin
understood and remains one of the major problems to be
resolved by muscle physiologists. Much progress has been
made in this field and the following section provides an
ae tte
Sapir picture of sarcomere contraction. (b) |
; 17.3 What happens to the length : cross-bridge

formation
:
|
H — 1bands as the sarcomere contrac ! myosin
— ATA Explain how any chang filament
|
and Iband lengths are brought. about in ter
_ happens to ihe a ae myosin filaments
| .
(©) | movement of actin

change in
orientation
17.4.7 Excitation—contraction—coupling of myosin
head |
|
At rest a sarcomere possesses Mg” ions and |
ATP in certain concentrations, but Ca’* ions are present
only in very low concentrations. Under these conditions (d)|
the actin filament is in the ‘off’ position. This is achieved by breaking of
See
tropomyosin being positioned on each actin molecule in cross-bridge

such a way that it blocks the sites on actin to which myosin
will attach. Also the myosin heads are held away from actin
in a position close to the long axis of the myosin filament.
When a muscle is stimulated by a nerve impulse, the
Fig 17.24 Excitation—contraction coupling. (Tropomyosin and
wave of depolarisation spreads over the muscle and passes troponin not shown.)
from the outside of the muscle fibre membrane into the
sarcomere along the T system. As the impulse reaches the
17.4.8 The energy supply
triad vesicles it stimulates them to release Ca’* ions into the
sarcoplasm, and Ca’* ion concentration consequently rises. Within the body the ultimate source of energy
Ca’** ions bind to the troponin-C, which in turn interacts for muscle contraction is usually glycogen, but may be fatty
with troponin-I and reverses the normally inhibitory effect acids. When these substrates are metabolised (during
of the troponin system on actin—myosin interaction. Actin respiration), ATP is generated. It is the hydrolysis of ATP
is switched ‘on’ when the tropomyosin moves to a new that liberates the energy necessary to promote actual
position on each actin molecule (fig 17.19) so that the muscle contraction:
myosin-binding sites are exposed. In some muscle, not ATP—————» ADP +P; + energy for muscle contraction
vertebrate, the ATPase activity of myosin is also stimulated
by the presence of Ca”* ions. It is further enhanced by the In resting muscle the level of ATP is low, being sufficient
presence of actin. In all muscles, when the actin and myosin only to power about eight muscle twitches. This condition
have been activated, the myosin head moves out from its is maintained by normal aerobic respiration. The ATP is
resting position and links to actin to form an actomyosin soon used up when a muscle contracts, and has to be
cross-bridge. Release of energy by ATP hydrolysis accom- quickly restored by other processes.
panies cross-bridge formation and leads to a change in the Restoration of ATP involves a substance located in the
angle of the cross-bridge such that the myosin head pulls muscle called phosphocreatine (PCr). The ADP produced
the actin filament over itself towards the centre of the during muscle contraction is reconverted to ATP at the
sarcomere. With all myofilaments acting in this way during expense of phosphocreatine:
sarcomere stimulation the end result is the generation of a
ADP +PCr ——————» ATP+Cr
force (fig 17.24) which may lead to a shortening of the creatine
sarcomere length. phosphotransferase
When excitation of the sarcomere ceases, Ca”* ions are
This ensures that there is always a constant supply of ATP
actively pumped back into the triad vesicles by an
in the muscle which it can utilise for immediate contraction.
ATP-driven calcium pump. Ca”* ion concentration soon
At some stage, the phosphocreatine level has to be
decreases below the threshold for contractile activity and
replenished. This is brought about by oxidation of fatty
relaxation of the sarcomere begins. The tropomyosin—
acids or glycogen. The ATP produced here enables
troponin complex inhibits ATPase activity, cross-bridges
phosphocreatine to be resynthesised from creatine:
are broken, actin and possibly myosin too are switched ‘off’
and the sarcomere reverts to its normal resting tension. Cr+ ATP —————_» PCr + ADP
640
(a) fatty
ey acid
wd
oxidation
: Dake +40
pizae ATP Cr ATP
eveaes aerobic
OIC energy liberated for
pyruvic respiration muscular contraction
acid ; ADP PCr ADP (b)
anaerobic
ct 1
N
glycogen lactic oxygen actual
in muscle [glucose requirement oxygen
oxygen y
blood _
oxygen/dm?
min
Extra resting level
repayment of
oxygen debt Exercise Recovery/min
Fig 17.25 (a)Relationship between ATP, phosphocreatine and respiration in the process of muscle contraction. LA, lactic
acid. (b) Oxygen requirements during exercise showing the relationship between oxygen intake and oxygen debt
When a muscle becomes very active its oxygen supply fully removed from the body represents the time it takes
rapidly becomes insufficient to maintain adequate oxida- the body to repay the oxygen debt incurred during
tive phosphorylation (section 11.3.6) of its respiratory strenuous muscular activity (fig 17.25).
substrates. Under these conditions pyruvic acid, the
end-product of glycolysis, is converted to lactic acid by
addition of H* ions. This occurs because there is 17.5 innervation of skeletal muscle
insufficient oxygen present to attach to the H” ions,
Each muscle is innervated by many motor
produced during glycolysis, to form water. Whilst this is
nerve fibres, all of which branch to supply a group of
happening the muscle is said to be incurring an oxygen debt.
muscle fibres. This group, together with its motor supply, is
CH,COCOOH +2H; ————> CH,CHOHCOOH called a motor unit, and all muscle fibres in it will contract
pyruvic acid ey lactic acid © simultaneously when suitably stimulated. The number of
Lactic acid formation is relatively inefficient as an muscle fibres in a motor unit is variable and depends on the
energy-liberating process, yielding only about 7% of the sophistication of control that the unit is required to exert.
energy available from the complete oxidation of glucose. For example, there are about ten in eyeball muscle but over
Lactic acid is toxic and sooner or later has to be removed 1 000 in a biceps muscle. The fewer the number of muscle
from the body. This occurs when muscular activity slows fibres in a unit, the greater the nervous control over them.
down or ceases. Where a motor nerve fibre makes contact with a muscle
When this happens the oxygen supply once again fibre, a neuromuscular junction, or motor end-plate, is
becomes sufficient to oxidise lactic acid and aid the formed. Here the axon of the motor nerve fibre loses its
reconversion of some of it into glycogen. This process myelin sheath, and its terminal dendrites are sunk into
generally occurs in the liver where one-fifth of the lactic grooves which ramify over the end-plate.
acid is fully oxidised to carbon dioxide and water, providing The stimulus for muscle contraction to take place is
energy which is utilised for the conversion of the remaining delivered by the central nervous system (fig 17.26).
lactic acid into glucose. Some of the glucose is transported Impulses are propagated along the motor nerve fibre to the
back to the muscle where it is finally transformed to motor end-plate. Here, in response to the nervous signal,
glycogen, whilst the remainder is converted to glycogen acetylcholine is released into the synaptic gap which
and stored in the liver. The time taken for lactic acid to be separates the motor fibre and sarcolemma of the muscle
folds in axon
synaptic knob of terminal
sarcolemma
dendrite myelin sheath
sarcoplasm
motor end-plate nucleus of muscle
synaptic vesicles containing mitochondrion

acetylcholine
Fig 17.26 Motor end-plate at a myofibrils

neuromuscular junction
641
Table 17.4. Structure, location and general properties of slow and fast skeletal muscle fibres
a
Slow/tonic muscle fibres Fast/twitch muscle fibres
Structure Many mitochondria Few mitochondria

Poorly developed sarcoplasmic reticulum Well-developed sarcoplasmic reticulum ;
Red — due to presence of myoglobin and cytochrome White — little or no myoglobin or cytochrome pigments
pigments
Low in glycogen content Abundance of glycogen granules
Capillaries in close contact with fibres to facilitate fast
exchange of materials
Location Deeply seated inside the limbs Relatively superficial
Innervation Associated with small nerve fibres of 5 wm diameter. Associated with large nerve fibres of 10-20 ym diameter.
A number of end-plates are distributed along the length Usually one or possibly two end-plates per fibre
of the fibre. This is called multi-terminal innervation.
Velocity of impulse conductance 2-8 m s ' 840 ms
axons
_, end-plates
—fibre
Excitability Membrane electrically inexcitable. Each impulse Membrane electrically excitable. Exhibit ‘all-or-none’
-causes release of only small amount of acetylcholine. response when an action potential is elicited.
Therefore amount that membrane is depolarised
depends on frequency of stimulation
Response Slow graded muscular contraction of long duration. Fast contraction (3 times faster than slow fibres)
Relaxation process slow (up to 100 times slower than Fatigues quite quickly
twitch fibre)
Physiological Depend on aerobic respiration for ATP production Depend on the anaerobic process of glycolysis for ATP
activity supply
Many continue to function anaerobically if oxygen in Oxygen debt quickly built up
short supply in which case lactic acid is formed and an
oxygen debt incurred
Carbohydrate or fat store mobilised at same rate as Glycogen used extensively as respiratory substrate
respiratory substrate is oxidised
Heat transported away from muscle as soon as it is Heat produced is absorbed by the fibres as the
produced circulatory system does not immediately remove it
Steady state between muscle activity and its needs Muscle contraction occurs during a period when the
is set up circulatory system has not had time to increase the
oxygen supply to the muscle
Function Enable sustained muscle contractions to occur. This is Immediate, fast muscle contraction is permitted at a time
used for the maintenance of posture by the organism when the circulatory system is still adjusting to the needs
of the new level of muscle activity. Therefore of great
importance during locomotion.
fibre. What happens at the end-plate is remarkably similar 17.5.1 Gradation of response by skeletal
muscles
to what happens at a neurone-neurone synapse (section
16.1.2). The acetylcholine diffuses towards the sar- In order for fine control of muscular activity to
colemma and effects a temporary increase in its permeabil- be exerted, it is important that the degree of tension
ity to ions, especially Na* and K*. Asa result, an end-plate developed by each muscle should be closely controlled.
potential is developed and this generates an action This is achieved in two ways, either separately or
potential which passes rapidly along the length of the collectively.
muscle fibre, also inwards via the T-system, eliciting (1) The number of muscle fibres that are actually excited at
contraction which is of an all-or-nothing nature. any one time may be varied. It follows that the force
Acetylcholinesterase, located in large quantities in the generated by a muscle will increase if an increased
region of the sarcolemma, rapidly hydrolyses acetylcholine number of fibres are stimulated, and vice versa. This is
to choline and ethanoic acid, and thus prevents over- what generally happens in vertebrate skeletal muscle.
stimulation of the muscle fibre. The motor end-plate (2) The frequency of nerve impulses received by muscle
rapidly reverts to its resting state once the end-plate fibres may be varied. Repetitive stimulation in thisway
potential has passed. can increase the force developed by the muscle.
642
The whole process of muscle contraction within an spindle-shaped connective
muscle es
organism is a smooth, orderly affair. This is achieved by the el
asynchronous contractions of different groups of muscle
fibres in antagonistic muscles.
17.5.2 Types of skeletal muscle fibre

There are two major types of skeletal muscle
fibre, each with its own specific physiological properties. faint striations
They are the slow or tonic fibres, and the fast or twitch of actin nucleus
fibres. Table 17.4 indicates their structure, location and Fig 17.27 Vertebrate smooth muscle
general properties. Whilst some muscles may contain
purely tonic, or twitch, fibres, some muscles contain
proportions of both.
arranged in a longitudinal fashion within each cell (fig
Collectively the two types of fibre endow the organism
17.27). It is now generally accepted that vertebrate smooth
with the ability to move about and to maintain posture. The
muscle has myosin filaments in its normal state, although
twitch fibres enable fast muscle contraction. Predators these may be different from those in striated muscle
possess many twitch fibres and use them for fast reactions filaments. Cross-striations are not seen here because the
to capture prey. On the other hand, would-be prey can also myosin and actin filaments through the cells are not in axial
react quickly in order to avoid capture by predators. In register. It is thought that the contractile mechanism in
both cases speed of body movement would influence the smooth muscle is essentially similar to that of striated
probability of survival of the organism concerned. When an muscle, although regulation of activity may be quite
animal is still it has to maintain a particular posture. This is different.
achieved by contraction of the tonic muscle fibres. They Conduction between cells is relatively slow and this
generate a slower, more sustained contraction, whilst at the results in prolonged slow contraction of the muscle and an
same time consuming less fuel than the twitch muscle equally slow relaxation period. The muscle is also capable
fibres. The nature of the contraction is usually isometric, of spontaneous rhythmic contractions and these may vary
and muscles and limbs are held at a constant length, thus both in intensity and frequency. Stretching of smooth
resisting the force of gravity. muscle, caused by the distension of the hollow organ which
In Man both types of fibre enter into the composition of the muscle surrounds, is generally followed by an immedi-
all muscles, but one or other usually predominates. The ate contraction of the muscle. This particular property
functional significance of this is that the predominantly again aids the propulsion of contents within the organ.
tonic muscles are suited to long-term slow contractions, The cells are not under voluntary nervous control, but
and consequently are found in the postural extensor instead are innervated by two sets of nerves from the
muscles, whilst twitch muscle fibres predominate in the autonomic nervous system. One set is from the parasym-
flexor muscles, which are designed to react at speed. pathetic and the other from the sympathetic system (section
16.2.3). The general opposing effects that they exert on the
17.5.3 Smooth muscle organs they innervate means that the activity of the organs
can be quickly controlled to meet any changing conditions
Vertebrate smooth muscle is located in the that might occur. Smooth muscle activity may also be
walls of many hollow structures of the body. These include modified by adrenaline and other specific hormones.
the intestinal tract, bladder, blood vessels, ureter, uterus
and vas deferens. The individual cells are uninucleate and
spindle-shaped. Connective tissue, consisting largely of 17.5.4 Muscle spindles and the stretch reflex
collagen, holds them together. They are oriented parallel
Within skeletal muscles are a number of
to each other and form a distinct muscle layer. An example
proprioceptors called muscle spindles (fig 17.28). The
of this is the smooth muscle of the intestine where there is
centre of each muscle spindle is composed of several
an outer longitudinal, and an inner circular, layer. When
modified non-contractile muscle fibres called intrafusal
the longitudinal muscle contracts, it shortens and dilates
whilst contraction of the circular fibres, enclosed within a connective tissue sheath. These
the intestinal lumen,
and constrict the lumen. Such are surrounded by a number of annulo-spiral nerve endings
muscle will lengthen
movement which unite to form an afferent nerve which passes to the
coordinated activity, called peristalsis, aids the
interesting spinal cord. Surrounding, and in parallel with, the
of the contents of the gut and provides an
to move intrafusal fibres are the muscle fibres of the muscle proper
example of the function of smooth muscle, namely
the body. (extrafusal fibres). The two ends of the spindle consist of
along materials within the hollow organs of
50-200 ym contractile muscle fibres innervated by efferent nerves
Each smooth muscle cell is approximately
. Actin is coming from the central nervous system.
long and 2-5 ym in diameter in the extended state
643
Fig 17.28 (a) Vertebrate muscle, muscle spindle and tendon active at different times according to the body’s needs.
organ; (b) Muscle spindle in detail. Arrows indicate the There are always some spindles operative at any given
direction in which nerve impulses travel time, being stimulated by a constant stream of signals from
(a) nerve the central nervous system. This causes a degree of partial
muscle contraction within the body called muscle tone.
Constant use of the muscles, through, for example,
muscle physical fitness, is required to maintain good muscle tone.
(extrafusal
fibres) The two ends of the spindle are important in maintaining
muscle tone. When stimulated by impulses from their
non-contractile efferent nerve supply they contract, and this ensures that
region of there is a degree of tension in the muscle before a load is
muscle spindle
added to it. If the muscle was not maintained in this
contractile
muscle spindle region of condition, there would be a danger that it would be
muscle spindle overstretched and damaged by the sudden application of
such a load.
tendon organ
tendon 17.5.5 Inhibitory reflexes
rl
For a limb to be moved to and fro it must be
(b) sensory neurone operated by at least two opposing muscles or sets of
118 from central
4 Ni region of muscles. When one contracts the other must relax. This is
annulo-spiral f muscle spindle achieved by a simple inhibitory reflex mechanism. It will be
nerve endings f
recalled that impulses generated by muscle spindles of a
ES non-contractile
— AY central portion muscle arrive at the spinal cord via an afferent neurone and
lymph-filled ES of muscle are eventually transmitted to the same muscle causing it to
capsule spindle
contract. The afferent neurone also synapses with inter-
intrafusal motor fibre to neurones in the grey matter of the spinal cord (fig 17.29).
muscle fibre contractile end
of muscle spindle When suitably stimulated, these inhibit the efferent
neurones leading to the antagonistic muscle, which is
therefore unable to contract and thus remains relaxed.
When a muscle is stretched, the annulo-spiral nerve A good example of this is the mechanism of walking.
endings of the spindle develop a tension and are stimulated Initially the limb flexes in order to lift the foot off the
to discharge a volley of impulses to the spinal cord via ground. During flexion, the antagonistic extensor muscles
afferent neurones. These synapse directly with efferent are stretched but are reflexly inhibited from contracting.
neurones which convey impulses to the extrafusal muscle After flexion, the limb is straightened and the foot is again
fibres causing a reflex contraction. This is the stretch reflex. brought into contact with the ground. With the flexor
The greater the degree of stretching incurred by the muscles no longer contracting, inhibition of the extensor
spindle, the more impulses that are discharged and this muscles ceases and the stretch reflex now proceeds
leads to a greater degree of muscle contraction. In this way culminating with the contraction of the extensor muscles.
the stretch reflex tries to maintain the constant length of the When the limb is straight no stretching in the extensor
muscle when the load on it is altered. muscle spindle is detected and the stretch reflex ceases. The
Muscle spindles act asynchronously, different ones being whole process is then free to be repeated.
spinal cord
MUSCLE
CONTRACTION
inhibitory pair of
interneurone antagonistic
muscles
MUSCLE
RELAXATION
|
motor axon
Fig 17.29 = Interneural inhibition. When muscle A contracts, imp ulses pass to the spinal cord where
they meet an inhibitory
interneurone. Inhibition of the appropriate motor neurone to muscle B causes it to relax
644
17.5.6 Tendon organs plasm. Locomotion is thought to be brought about by altern-
ate changes in the colloidal state of the cytoplasm effect-
Stretch receptors are also found in tendons.
ed by sol-gel-sol transformations, and the cytoplasmic
However they only respond when tension becomes severe.
streaming of plasmasol into the pseudopodia (section 4.1).
Upon stimulation impulses are discharged to the central
nervous system where they meet inhibitory interneurones. Where a pseudopodium is about to form, the plasmagel
Stimulation of these causes reflex inhibition of contraction liquefies into plasmasol. Plasmasol from within the cell now
flows towards this point and outwards into the newly
and inhibition of active resistance to stretching. This is
forming pseudopodium. Plasmasol in the outer regions of
significant because it protects the muscle from being
the pseudopodium is rapidly transformed into plasmagel
damaged when suddenly overloaded, or overcontracting
which thus forms a rigid collar around the pseudopodium.
and damaging itself in response to the application of a
At the posterior end of Amoeba the plasmagel is rapidly
oe load.
converted to plasmasol which then flows forwards (fig
17.30).
| aA ine fig 17.28 and answerr the
A number of theories exist to explain amoeboid
Ucowing. questic movement. One suggests that the posterior end of the
1 What happens to‘the muscle spindle whee the
animal contracts and drives plasmasol forwards into the
_ whole muscle (extrafusal fibres) is stret ched? newly forming pseudopodium. It is thought that variations
in the viscosity of the cytoplasm accompanied by slight
pressure changes in different parts of the cell stimulate
streaming of the cytoplasm from one region of the cell to
another. A more favoured theory, the ‘fountain-zone’
hat occurs. when the:intrafusal
theory, states that the cell is pulled forwards by contraction
cote; of the anterior end of the animal. Here, protein molecules
in the plasmasol exist in an extended state. When the
plasmasol is converted to plasmagel they contract and coil
17.6 Locomotion in a variety of inverte- up. As this occurs, protein molecules at the posterior end of
brates the animal are automatically extended, thus being trans-
formed into plasmasol which is then pulled towards the
17.6.1 Amoeboid movement anterior end.
Amoeboid movement is characterised by the These theories may have to be further modified because
formation of temporary projections of the cell called it has now been established that the protein molecules actin
pseudopodia. It occurs in protozoans of the class Rhizo- and myosin are present in all eukaryotic cells and that actin,
poda and in vertebrate white blood cells. During move- existing as fine filaments binds reversibly to myosin. This
ment the cell produces a definite anterior end which is at reaction is accompanied by the hydrolysis of ATP.
the point where new pseudopodia are forming. Analysis of Therefore possibly the whole mechanism of amoeboid
the cytoplasm of an Amoeba indicates that there is a movement, and possibly cytoplasmic streaming within
peripheral layer of viscous plasmagel, the ectoplasm, which cells, may be essentially similar to that of muscle
encloses a more fluid cytoplasm, the plasmasol or endo- contraction itself.
Direction of movement of Amoeba
plasmagel
forward movement
of sol
a >
anterior
posterior hyaline cap
of
plasmagel
eae: pseudopodium
=
ee + per ska fluid

solation layer moving
lasma towards posterior zone of
Cmembrane gelation
‘fountain-zone’ theory, plasmasol gelates and contracts, thus

Fig 17.30 Pseudopodial formation in Amoeba. According to the
pulling more plasmasol towards it
645
associated material as
Fig 17.31 Structure of a cilium or flagellum. (a) An interpretation of the arrangement of tubules and
Two arms are attached
seen from outside. (b) The structure seen in cross-section. Each doublet consists of an A and a B tubule.
wall of the B tubule is shared
to the A tubule. Note that the A tubule is a complete circle in cross-section, whereas part of the
of each doublet; they connect the doublets to the sheath surrounding
with the A. Spokes occur at intervals along the length
the central tubules. (c) An enlarged microtubule doublet.
(a) radial doublet __. (b) arm (c) shared
protofilaments
enlarged
microtubule
doublet
central
tubule
arm
membrane
17.6.2 Cilia and flagella The end of the cilium or flagellum attached to the cell or
organism terminates in a basal body which is essentially
Electron micrographs indicate that cilia and identical in structure to an axoneme but has a ‘9 + 0’
flagella possess identical internal structures. Cilia are structure, and is derived from a centriole. It differs from a
simply shorter versions of flagella and, unlike flagella, are centriole only in possessing a complex organisation at its
more commonly found in groups than in isolation. A basal end known as a cartwheel structure. The basal body is
transverse section of either organelle shows it to consist of a
thought to act as a template for the assembly of
pair of central filaments surrounded by nine peripheral
microtubules during development of cilia or flagella. There
filaments (fig 17.31), the so called ‘9 + 2’ array. This
are often fibres which extend into the cytoplasm from the
bundle of filaments, called an axoneme, is surrounded by a
basal body which act to anchor the basal body in position.
membrane that is continuous with the plasma membrane.
Whilst cilia and flagella possess fundamentally similar
Each peripheral filament is composed of the protein internal structures, their mode of action is quite different.
tubulin and consists of an A and a B microtubule. Each A A flagellum possesses a symmetrical beat with several
microtubule has a pair of ‘arms’ composed of another undulations occurring along its length at any given moment
protein called dynein which is capable of hydrolysing ATP,
(fig 17.32). This wave-like motion may be in one plane, or
in other words it is an ATPase. The central filaments are
less commonly such that it beats in a corkscrew (helical)
connected to the A microtubules of the peripheral
manner which spins the body of the organism about its
filaments by radial spokes of material.
longitudinal axis as well as propelling it forwards along a
helical path (fig 17.33). In some flagellates the flagellum is
at the anterior end of the body and beats in such a way as to
pull the organism through the water. This sort of flagellum
generally possesses minute lateral projections called
mastigonemes on its surface which aid this type of
locomotion. The flagellum itself is sometimes called a
‘tinsel’ flagellum. More usually, the flagellum is at the rear
of the organism or cell and pushes it through the water, as
in the spermatozoan tail. Fig 17.34 shows a summary of the
types of movement induced by flagella activity.
The beat of a cilium is asymmetric (fig 17.32), there being
an active, fast, straight downstroke followed by a slower,
bent, limp recovery action at the end of which the cilium
returns to its original position. With so many cilia occurring
together it is essential that some sort of mechanism exists to
coordinate their activity. In the ciliate Paramecium, a
traditional view is that this might involve fibres called
Fig 17.32 Successive stages in the motion of a flagellum (a) neuronemes which interconnect the basal bodies. Gener-
and a cilium (0). The effective stroke of the cilium begins at 1. ally the cilia beat in a synchronised fashion which results in
646
Sa
S
Euglena rotates
about its own axis Q
path traced by
anterior end of
Euglena
AN
uz > S. = S Ss ey euglenoid movement

Be promoted by myonemes
general direction E changes Euglena’s
——————
of locomotion direction
Fig 17.33 (above) Euglena’s path of movement waves of ciliary activity passing along the length of the body
in one particular direction. This is called metachronal
rhythm.
There has been much debate concerning the mechanism
Fig 17.34 (below) Summary of types of movement induced involved in the actual beat of a ciltum or flagellum. Current
by flagella activity. evidence suggests that a process exists that is essentially
similar to that of the sliding filament theory for muscle
Planar beat Helical beat contraction. A flagellum is said to begin to bend when the
(beat in one plane) (less common)
two dynein arms of the A microtubule of a peripheral
filament link up with an adjacent B microtubule. When this
cell pulled through
(a) water (flagellum at happens ATP is hydrolysed and the A and B microtubules
anterior end) slide over each other causing movement of the flagellum. It
is thought that five peripheral filaments on one side operate
y thrust y thrust
in this manner to produce the initial movement, whilst the
remaining four on the other side slide fractionally later thus
movement e.g. Spermatozoa | movement e.g. Euglena inducing the recovery action (fig 17.35). The radial spokes
of the cilium tend to resist this sliding process, which
instead is converted into a local bending action. It is
(b) cell pulled
through water thought that the central filaments may transmit the signals
for sliding from the basal body along the length of the
|
Y thrust cilium or flagellum. It has also been proved that ciliary
activity will only proceed when Mg” ions are present, and
movement e.g. Trypanosoma
that the direction of the beat of a cilium is dictated by
specific levels of intracellular Ca** ions. Indeed it is
interesting to note that the avoiding action of Paramecium
is controlled in this way. When Paramecium encounters an
© water pushed towards
ensel cell by minute obstacle, it reverses the beat of its cilia before moving
Fisccllumn: projections, the forward again. Reversal of ciliary beat is stimulated by a
e mastigonemes, on
flagellum sudden influx of Ca’* ions into the cell due to increased
permeability of the organism to Ca’* ions.
A thrust
| Both organelles are used extensively by small eukaryotic
e.g. some organisms to propel themselves through water. The cilia or
movement unicellular algae,
some zoospores
flagella provide forward propulsion for the organism by
thrusting against the surrounding viscous watery medium.
Locomotion brought about in this manner is only effective
NB thrust= direction of
propagation of waves for small organisms which have a much larger surface
647
Fig 17.35 The sliding of doublets during motion of a cilium change its direction by the action of contractile myonemes
in the gills of a mussel. One of the two tubules of each which lie along the length of its body. When they contract,
doublet protrudes further into the tip of the cilium than does the shape of the body is changed as well as its direction.
the other. Thus, cross-sections at an appropriate constant This is called euglenoid motion (fig 17.33).
distance from the tip will show a change in tubule pattern
during bending, from all doublets to doublets and single
tubules. The entire doublets slide with respect to one another; 17.6.4 Locomotion in Paramecium
but the two members of each doublet retain their positions
relative to each other. (After P. Satir, from Novikoff & Surface cilia, arranged in rows, beat diagon-
Holtzman.) ally backwards from left to right causing the animal to
rotate about its longitudinal axis. At the same time,
strongly beating oral groove cilia tend to cause the anterior
() {
ass
S ©C@ 9
00
:
cross-section
end of the animal to move in a spiral fashion about its
® ® at indicated posterior end. The cilia exhibit metachronal rhythm and
level
their coordinated activity may be governed by the
motorium, a body which is interconnected with the
neuronemes and basal bodies. Paramecium swims at a
speed of about 1 mm s_', or four times its body length.
17.6.5 Locomotion in the Annelida

An oligochaete, Lumbricus terrestris (the
earthworm)
The coelom of an earthworm is enclosed by a body wall
composed of two antagonistic muscles, an outer circular
layer and an inner longitudinal layer. The circular muscle is
area:volume ratio than large animals, where the minute divided into separate units along the length of the animal by
forces generated would be quite inadequate. Cilia also septa between the segments, but the muscle fibres of the
frequently occur within the bodies of multicellular organ- longitudinal layer generally extend over several segments.
isms where they serve a number of important functions. Locomotion is brought about by the coordinated activity of
They may propel fluid through ducts such as in the the two muscle layers and those of the chaetae.
nephridia of annelids where metabolic waste is removed. When an earthworm begins to move forwards, contrac-
They also propel eggs along mammalian oviducts, and tion of the circular muscles begins at the anterior end of the
move materials over internal surfaces, such as mucus body and continues, segment by segment, as a wave along
through the respiratory passages, where this activity helps the length of the body. This activity exerts pressure on the
to keep them free from dust particles and other debris. coelomic fluid in each segment, stretching the relaxed
They can also create feeding currents, as in Paramecium, longitudinal muscle and changing the shape of the
from which food particles can be filtered and removed, segments such that they become longer and thinner. This
often by other cilia. causes the anterior end of the worm to extend forwards.
Some bacteria possess flagella, but these differ markedly Chaetae, present in all segments except the first and last,
from eukaryotic flagella, being shorter, thinner and quite are retracted during the activity of the circular muscles and
stiff. The bacterial flagellum is extracellular (not sur- therefore do not impede the forward movement (fig 17.36).
rounded by a plasma membrane) and resembles a single Whilst the anterior end of the worm is moving forward,
microtubule of a eukaryote flagellum. It is moved by forces
longitudinal muscle in more posterior segments contracts,
that emanate from the point where it is joined to the
causing this region of the worm to swell and press against
bacterium.
the surrounding soil substratum. Chaetae in this region are
protruded and help the worm to grip the substratum. This is
17.6.3 Locomotion in Euglena
particularly useful during burrowing for the worm can exert
The locomotory flagellum is at the anterior a powerful thrust against the surrounding soil particles
end of the body and pulls the organism forward. Waves of during forward locomotion.
activity are generated by the flagellum itself, and as they Contraction of the circular muscle is quickly followed by
pass in a spiral fashion from its base to its tip they increase contraction of the longitudinal muscles throughout the
in amplitude and velocity. This activity of the flagellum length of the body, and this means that different parts of
causes the body of Euglena to rotate about its axis, at about the worm may be either moving forward (when the circular
one complete body turn per second, as well as making it muscles contract) or static (when the longitudinal muscles
describe a corkscrew pathway through the water. The contract) at any given moment. The net effect is a smooth
organism can travel forward at arate of 0.5 mm s_' whichis peristaltic wave of activity along the length of the worm as
approximately four times its body length. Euglena is able to it progresses forwards. The worm is also able to crawl
648
movement
‘ forwards
extension (E)
circular muscles ‘
contract, longitudinal
muscles relax, chaetae
anterior retracted
retraction (R)
circular muscles relax, 4
longitudinal muscles
stationary , contract, chaetae
enero stationary (S) retracted
both sets of muscles
relaxed, chaetae
extended and anchor
worm to substratum
posterior
1 2
Fig 17.36 (above) Locomotion in the earthworm

Fig 17.37 (below) Segmental innervation ofdongitudinal
muscle in the earthworm (dorsal view). A similar arrangement
is present in the circular muscle
nerve cord longitudinal A polychaete, Nereis (the ragworm)
—_—_—7 ae ve
muscle
In contrast to the earthworm, the longitudinal muscle layer
circular.
muscle is divided into a pair of dorsal and a pair of ventral muscle
blocks (fig 4.26). The circular muscle is relatively weak and
segmentally specialised into sets of oblique muscles which
sensory intermediate
run into the parapodia and function to move the parapodia
neurone neurone
and their associated structures. Each parapodium is hollow
motor internuncial and ramified by an extension of the coelom which is filled
neurone neurone
with coelomic fluid. It is subdivided into a dorsal
septum projection, the notopodium, and a ventral projection called
the neuropodium. Both of these structures possess a
strengthening rod, the aciculum, and a bundle of chaetae.
bilo
Interaction between the body wall muscles, parapodia
and coelomic fluid brings about several different types of
locomotion. Slow creeping movement is achieved by using
backwards by reversing the direction of contraction of the the parapodia as a system of levers. During its effective
muscles. stroke a parapodium protrudes its aciculum and chaetae to
Control of muscle contraction is brought about by a make contact with the substratum and push against it in a
complex network of inter- and intrasegmental neurones. backward direction, thus propelling the animal forwards.
Allsegments are in contact with the longitudinal nerve cord Protrusion of these structures is achieved by increased
and also possess their own set of segmental nerves. This hydrostatic pressure in the coelom of the parapodium. The
means that localised control of each segment is possible, as parapodium moves forward when it lifts from the substra-
well as control of overall activity of the animal (fig 17.37). tum and its aciculum and chaetae are retracted. Retraction
The ventral nerve cord possesses a giant axon which runs is achieved by contraction of the segmental oblique
centrally along its length and conducts impulses in an muscles.
anterior-to-posterior direction. Also present in the nerve The movements of the right and left parapodia of a
cord are two longitudinally oriented lateral fibres which segment are coordinated so that whilst one is moving
convey impulses from the tail to the head. When sensory forwards the other is moving backwards. Collectively the
receptors in the head are stimulated, impulses pass along parapodia of the worm move asa series of peristaltic waves
which pass along the length of the body. In contrast to the
the giant axon stimulating the longitudinal muscles to
earthworm these waves move in a direction from the tail to
contract, thus causing the anterior end to be recoiled from
the head with a group of parapodia on one side all moving
the stimulus. If the tail of the worm is similarly stimulated,
generally forwards whilst at the same time their opposite
impulses pass along the jateral fibres from tail to head and
numbers move backwards.
cause the tail to be withdrawn. This is the basis of the
Rapid creeping occurs when the longitudinal muscles on
worm’s escape reaction.
649
either side of the body contract alternately throwing the When an insect begins to walk, three legs remain on the
body into a series of lateral undulations. Here the ground to support the animal whilst the other three move
parapodia again come into contact with the substratum. forward. The first leg on one side pulls the insect, whilst the
When this occurs, the longitudinal muscles work against third leg of the same side pushes. The second leg on the
these points of contact and effectively pull the animal other side serves as a support for this activity. The process
forwards. is then repeated but with the role of each trio of limbs
When swimming freely in the water the right and left reversed.
longitudinal muscles contract alternately to produce waves Many insects possess a pair of claws and a sticky pad at
of lateral movement that pass forward from the tail to the the distal ends of their legs. The pad consists of minute
head. The broad, paddle-like parapodia move in a hollow tubes which secrete a sticky fluid that helps the
backward direction during their effective stroke. In doing insect to adhere to smooth surfaces. Thus, these insects are
this they push against the water and thrust the animal able to walk up vertical surfaces as well as upside down.
forwards. Flight. The wings of insects are flattened
extremities of the exoskeleton and are supported by an
17.6.6 Locomotion in the Arthropoda intricate system of veins. Their movement is controlled by
Crustacean walking and swimming, two main groups of muscles, direct and indirect muscles. In
Astacus fluviatilis (the crayfish) insects with large wings (such as butterflies and dragonflies)
the muscles are actually attached to the bases of the wings
The crayfish possesses eight thoracic and six abdominal (fig 17.38); these are the direct muscles. They elevate and
segments each of which bears appendages of various shapes depress the wings as well as controlling the angle of the
and sizes which are used for specific functions (fig 4.34). wing stroke during flight. When the angle of one wing is
The last four thoracic segments bear walking legs called adjusted with respect to the other, turning in the air is
pereiopods, whilst forked paddle-like appendages called accomplished. They are also used to fold up the wings when
pleopods are found on abdominal segments 2-5 in the the insect is stationary.
female and 3—5 in the male. The sixth abdominal segment, In insects with smaller wings, the main driving force for
called the uropod, is modified as a flattened tail fin which is flight is developed by two sets of antagonistic, indirect flight
most frequently used in swimming. muscles; the dorso-ventrals are attached to the tergum
When walking, the chelipeds of the fourth thoracic (roof) and sternum (floor) of the thorax, and the
segment (section 4.10.1) are raised above the substratum longitudinals are attached to the anterior and posterior
and the abdomen is extended horizontally. Only two legs, aspects of the thorax (fig 17.39). There is no direct
one on either side of the body, are out of contact with the attachment of the wings to these muscles. However the
ground at any one time, the other six firmly grip the base of each wing is attached to the tergum and pleura
substratum with those of thoracic segments 5-7 pulling the
(side) of the thorax. This arrangement acts as a highly
crayfish along while the pair on the eighth thoracic segment
efficient lever system, for when either of the indirect
push it.
muscles shortens fractionally it distorts the shape of the
Muscle which promotes movement of the limbs in the
thorax, which in turn produces a large degree of movement
crayfish is striated; that in the region of the abdomen is
by the wings. Because muscle contraction takes place over
particularly strong. Here thick antagonistic muscles oper-
a very small distance it can be repeated rapidly. This is
ate to flex and extend the abdomen up and down. When the
particularly important for insects that possess very fast
abdomen is flexed downwards the force exerted on the
wingbeat speeds.
water by the extended tail is sufficient to cause the crayfish
to shoot backwards through the water very quickly. This is body (only one a
the basis of its escape mechanism. side shown)
Locomotion in insects WING

DEPRESSED elevator
Walking. This is achieved by the coordinated muscle
relaxes
activity of three pairs of legs, one pair being attached to depressor muscle
contracts
each of the three thoracic segments of the animal. Each leg
consists of a series of hollow cylinders whose walls are
composed of rigid exoskeletal material. The cylinders are
linked together by joints and soft pliable membranes. WING
ELEVATED
Where the coxa (the basal segment of the insect leg) joins
to the body, a form of ball-and-socket joint occurs, but all elevator
other joints in the leg are hinge joints. Bending and depressor muscle muscle
relaxes contracts
straightening of the legs is achieved by antagonistic flexor
and extensor muscles attached to the inner surface of the Fig 17.38 Action of direct flight muscles in a large winged
exoskeleton on either side of a joint (fig 17.1). insect, such as a butterfly or dragonfly
650
Fig 17.39 Cross-section of the thorax of an insect from the anterior showii ing the relationsh ] |
tergum and muscles during insect flight ? ee D
t pleural tergal process
UPSTROKE
process
dorso-ventral muscles relaxed
longitudinal muscles relaxed
dorso-ventral muscles contracted
ee :
contracted it
S leading
Sane
cde
win
pleuron
thoracic leg
S sternum DOWNSTROKE
In large, winged insects such as the butterfly and locust The following account of asynchronous flight is based on
the rate of wing beat is between 5-50 beats per second detailed observations made by Boettiger on the fly
(table 17.5). Here the flight muscles contract each time as a Sarcophaga bullata. Whilst flying in a straight line the wings
result of a single nerve impulse. Hence impulses are describe a pathway through the air in the form ofa figure of
generated at the same rate as the wings beat. Insect flight eight. The downstroke provides the majority of forward
muscle which responds in this way is called synchronous thrust and lift for the insect. During this phase each wing
muscle. In the housefly, which has a wing beat frequency of beats forward and downward with its anterior margin
120-200 beats per second, contraction of the flight muscles inclined at a lower level than its posterior one. On the
is much too fast to be triggered by individual nerve upstroke the wing moves upward and backward with its
impulses. This muscle is termed asynchronous and receives anterior margin raised above the posterior one. In this
roughly one impulse per 40 wing beats, which is necessary position the wings move through the air with minimum
to maintain the muscle in an active state during flight. It can resistance whilst at the same time providing more lift for
contract further and generate more power than synchro- the body.
nous muscle. Asynchronous muscle can also automatically In order to raise the wings, the dorso-ventral muscles
contract in response to being stretched. This is called the contract. When this occurs the tergum is lowered and the
stretch reflex and occurs faster than the speed of a nerve tergal attachment of each wing is pulled into a position
impulse. below that of its pleural attachment (fig 17.39). During this
process the resistance of the tergum to this distortion
Table 17.5. Wing speeds of a variety of insects increases. However, at a critical point the resistance
disappears and the wings click into an elevated position. At
Wing speeds the same time that this is happening the longitudinal
muscles are stretched, considerably stimulating a stretch
Large butterfly
5 times per second reflex which makes them contract instantaneously. The
e.g. swallowtail
Locust Sars ee tergum now arches upwards and the tergal attachment of
Hawkmoth A() ee x the wing is raised above the pleural one. Once again
Housefly 20 a at this speed resistance to such a movement is suddenly overcome and
a a humming
Bee 180 ,,
sound heard
the wings click downwards. This action in turn stretches
,, - at this speed and stimulates the dorsoventral muscles to contract, and
Midge 700-1000
a high pitched the whole cycle is repeated.
whine heard
oOaS
In general, the smaller the insect the faster it beats its wings. 17.6 The sarcoplasmic reticulum of in-
NB Some insects have two pairs of wings (such as locusts and dragonfl
ies). sect flight muscle is modified to increase its surface
In some cases both pairs of wings beat together , as in bees; in others the area by being perforated at intervals. Can you
of the front pair, for example
back pair of wings beats slightly ahead suggest a reason for this?
beetles.
locusts. Some insects have one pair of wings, such as flies and
In houseflies the hindwin gs are reduced and modifie d to form a pair of 17.7. Would synchronous or asynchro-
oscillat e rapidly
club-shaped halteres which are sensory in function . They
tion for the nous muscle be expected to contain more sarcoplas-
during flight, detect aerodynamic forces and provide informa mic reticulum? Give a reason for your answer.
in flight. Some insects (very few) have no wings,
maintenance of stability
for example fleas.
651
iform locomotion
Fig 17.40 Comparison between (a) ostraciform, (b) carangiform and (c) anguill
/forward component (F)
reaction (R)
movement of
undulations
forward
forward
component _
component reaction
reaction
lateral lateral
drag drag
push of tail
against water
pushof tail
against water
(a) (b)
The myotomes contract and relax alternately-on each side

17.7 Locomotion in vertebrates
of the vertebral column beginning at the anterior end of the
17.7.1 Swimming in fish fish and travelling towards its tail. This activity bends the
body of the fish into a series of waves, the number
Water, particularly sea water, has a high increasing the longer and thinner the fish.
relative density, many hundreds of times greater than air. Very compact fish such as Ostracium (the tunny) show
As such it represents a comparatively viscous medium to little evidence of this wave-like action with as much as 80%
move through. However, its density is made use of by fish of their forward thrust being achieved purely by the
as it supports them and also provides a medium against side-to-side lashing of the tail and caudal fin. This
which the fish can thrust during swimming movements. locomotion is called ostraciform. Longer fish, such as the
Any successful organism shows many adaptive features dogfish and the majority of bony fish, exhibit carangiform
suited to the environment in which it lives and afish is no locomotion; here the posterior half of the fish is thrown into
exception. The body of most fish is highly streamlined, a series of waves. Anguilliform locomotion, as demons-
being tapered at both ends. This means that water flows trated by eels, is where the body is very long and thrown
readily over the body surface and that drag is reduced to a into many waves so that different parts of the body are
minimum. Apart from the fins no other structures project moving simultaneously to the left and to the right.
from a fish, and it seems that the faster the fish, the more
perfect is the streamlining. The dermal denticles of
_
cartilaginous fish and the scales of bony fish are moistened
by slimy exudation from mucus or oil glands and this also lateral drag
counteracted
considerably reduces friction between the fish and the by pressure of
water. Other general adaptations possessed by fish for water against
head and
moving efficiently through the water are the fins. Dorsal median dorsal
and ventral, unpaired, median (along the midline of the fin
body) fins help to stabilise the fish, the paired pectoral and
pelvic fins are used for steering and balancing the animal,
and the caudal or tail fin, in concert with the paired fins,
provides the forward movement of the fish through the
water. Details of precisely how the fins operate will be
discussed later.
reaction : et
direction in
Liha Propulsion in fish which tail
moves as it
Movement is brought about by sets of pushes against
the water
segmentally arranged antagonistic muscle blocks called
myotomes located on either side of the vertebral column.
Superficially each myotome possesses a zig-zag shape, and lateral drag
component
internally it traverses the joint formed by two adjacent
vertebrae. The vertebral column isa long, flexible rod and, Fig 17.41 Force components exerted by the caudal fin of a
when myotomes on one side of it contract, it bends easily. dogfish as it is lashed from side to side
652
Fig 17.42 Action of paired pectoral and pelvic fins and heterocercal tail in the dogfish to provide lift
anterior posterior
lifting force Peace lifting force reaction dorsal fin chains5re fin
4
reaction lifting force

J
/
/
backward
backward
drag
ead oe forward
weight
§ thrust
of body
push of paired pectoral and heterocercal
pelvic fins against the water anal fin a
Forward propulsion is generally effected by the side-to- this the paired pectoral and pelvic fins act as hydrofoils.
side movement of the tail to which is attached the caudal They are held at an angle to the long axis of the body, and
fin. As the caudal fin is moved in this way it bends slightly when the fish wishes to swim up or down in the water their
and exerts a backward pressure on the relatively viscous angle is altered accordingly by muscular activity. When
water. This force can be resolved into a forward and lateral held at an angle, the force exerted by the fins can be
component. The forward one thrusts the fish through the resolved into an upward component, which forces the head
water whilst the lateral one tends to swing the head of the of the fish upwards (positive pitch), and a backward or drag
fish sideways in a direction opposite to that of the tail. This component (fig 17.42). However the drag is considerably
is called lateral drag (fig 17.41). Fortunately, it is less than the forward thrust produced by the tail and is
counteracted by the inertia of the water against the relatively unimportant. The dogfish possesses a heterocer-
relatively massive anterior end of the body (when com- cal tail, the ventral lobe being proportionately larger than
pared to the tail) and the large surface area posed by the its dorsal counterpart. As the tail thrusts from side to side,
dorsal median fin. Also it would require a much greater once again the force produced by it can be divided into an
force to move the body laterally through the water than to upwards thrust, termed negative pitch, as well as a lateral
propel it forwards. The magnitude of the force that the tail component. The positive pitch at the anterior end is
and caudal fin apply to the water depends on their speed of balanced by the negative pitch at the rear.
action, surface area and the angle at which they are held As the fish swims along in a level forward manner it may
with respect to the water. be subjected to three kinds of displacement or instability.
These are yawing, pitching and rolling. The various fins of
17.7.3 Locomotion ina cartilaginous fish, the the body are generally responsible for maintaining control-
dogfish led forward locomotion and the way in which they
counteract any displacement of the fish can be seen in fig
A dogfish is heavier than sea water and will
1743s
begin to sink to the bottom if it ceases to swim. To avoid
(a) = A vA (b) YAW
\\Z
) (resisted by anterior and posterior dorsal
fins, and ventral anal fin)
ROLL
yaw (turning) (regulated by
all fins)
YAW
itch
(up and down)
ae.
DO
FIG Fig 17.43 (a) Yaw, pitch and roll.
Y (regulated by pectoral (b) Functions of the fins to control
roll and pelvic fins)
yaw, pitch and roll
(leaning or tilting)
653
Fig 17.44 Closed swim bladder of a bony fish; (a) location, (b) bladder and circulation
(a) (2) Vy dorsal aorta —>
i.
-La the heart
swim bladder
reabsorption
©)) area
i
portal vein
DP
<—to the liver y y gas gland
The fins respond almost immediately to any form of automatically increasing or decreasing the amount of
instability. This is because of the activity of a set of three gas in its bladder, the fish can match the density of the
semicircular canals located on either side of the head of the surrounding water and thus preserve ‘neutral buoy-
fish. Should the body yaw, pitch or roll, then the ancy’.
appropriate semicircular canal is stimulated to fire nerve The closed swim bladder is under nervous control.
impulses which pass to the brain. The brain, in turn, At its anterior end is a structure, the gas gland, rich in
immediately sends the appropriate motor impulses to the blood capillaries, which secretes gas rich in oxygen into
fin muscles which adjust their position accordingly. the bladder. Posteriorly is another heavily vascularized
region which can absorb gases from the bladder (fig
17.7.4 Locomotion in a bony fish, the herring 17.44). In the gas gland, arterial and venous capillaries
are interspersed amongst each other. It is here that a
Bony fish possess a structure called the swim
countercurrent system operates in order to facilitate
or air bladder. Whilst lobe-finned fish used the swim
secretion of oxygen into the bladder. The gas gland
bladder to perfect the air-breathing habit, bony fish
produces lactic acid when gas secretion occurs, and as
adapted it into a hydrostatic device. The swim bladder is a
the acid enters the venous capillaries it reduces the
sac lying between the vertebral column and gut and
affinity of haemoglobin for oxygen (this is called the
functions to provide the fish with ‘neutral buoyancy’. When
root effect). This phenomenon increases the oxygen
this occurs the fish possesses a density equal to that of the
tension of the blood in the venous capillaries leaving
surrounding water and therefore does not need to expend
the gas gland, and paradoxically the venous blood
energy to keep itself from sinking, so concentrating its
efforts on moving through the water.
With the development of such a swim bladder in bony
fish, the paired fins were released from their lifting
function. Now, they are much smaller than those of the swim bladder
dogfish and are used instead as stabilisers or brakes, in the 10.1 MPa O,
10 cm? O, per 100 cm? blood
latter case being spread vertically at 90° to the body. Each
arterial capillary
pectoral fin may be used independently of its opposite
number, and in this way they act as pivots round which the
fish can turn rapidly. When the fish is swimming in a
straight line the paired fins are pressed firmly against the
sides of the body thus improving its streamlined shape.
venous capillary
Possession of a swim bladder has also permitted the
development of a symmetrical homocercal tail which 9 cm?O, per 100 cm? blood
transmits most of the force it develops against the water ina
forward direction.
Two types of swim bladder exist.
(1) Open swim bladder (as in goldfish, herrings). The Fig 17.45 Countercurrent system in the closed swim bladder
bladder is connected to the pharynx by a duct. Air is of a bony fish. The gas gland produces lactic acid which
taken in or expelled from the bladder via the mouth increases the oxygen tension in the venous capillary. Thus
and duct, thus decreasing or increasing the relative gas diffuses from the venous to arterial capillaries, and
remains within the loop. As venous blood leaves the gland it
density of the fish respectively.
contains less oxygen than the incoming arterial blood. NB For
(2) Closed swim bladder (as in codfish). The bladder has simplification only one venous and one arterial capillary are
completely lost its connection with the pharynx. By shown. The gas gland itself contains numerous capillaries.
654
possesses a higher oxygen tension than that in the Fig 17.46 Crawling in a frog viewed dorsally
arterial capillaries (fig 17.45). Therefore oxygen (a) Left forelimb retracts, right hindlimb extends, right forelimb
constantly diffuses from the venous to the arterial extends, left hindlimb placed forwards.
(b) Left forelimb extends, right hindlimb is placed forwards,
capillaries and is secreted into the swim bladder. At the right forelimb retracts, left hindlimb extends
time of gas secretion by the gas gland the blood vessels
of the posterior region of the swim bladder are closed
off so that no gas can escape from it. (a) right forelimb
ethe adaptati
mming.
17.7.5 Locomotion in the frog
right hindlimb
In the frog the paired limbs are no longer used
as fins for stabilising and steering the animal, but are
modified to operate as elongated jointed levers, raising the
body off the ground whilst at the same time moving the (b)
animal over land. The forelimbs are relatively short, with left forelimb
the radius and ulna united. The hindlegs are very long and
held underneath the body when the animal is at rest, by
being bent at the knee and ankle. The tibia and fibula are
fused together and the tarsals greatly elongated. The
girdles, which articulate with the limbs, are greatly
left hindlimb
enlarged when compared with those of fish and support the
weight of the animal against the effects of gravity. This is
especially true of the pelvic girdle.
When a frog crawls over the substratum its mode of
locomotion is effected by diagonally opposite limbs
operating in unison with each other. If we begin with a
stationary frog with all four limbs on the ground, the straightened by rapid contraction of strong extensor
following sequence of events takes place when it begins to muscles. The force exerted against the ground during this
crawl. The left forelimb retracts at the same time as the activity is transmitted from the limbs to the vertebral
right hindlimb extends. Both limbs thrust against the column via the pelvic girdle and is at least three times
substratum and so propel the animal forwards. This is greater than the weight of its body. This is sufficient to
followed by retraction of the right forelimb and extension propel the animal upwards and forwards into the air against
of the left hindlimb, the whole process being repeated as the downward pull of gravity. On landing, the short,
long as the animal is crawling. A full explanation is given in compact forelimbs, attached in a flexible manner to the
fig 17.46. pectoral girdle, dampen down the shock of impact (fig
At take off, during a jump, each joint in the hindlimb is 17.47).
Fig 17.47 The take-off and jump of a frog
a 655
(a) (b)
(c)
Fig. 17.48 (a) Quill feather, (b) contour feather, (c) down
feather
656
Fig 17.49 Pigeon wing extended
primaries to show primary and secondary
feathers
alula
secondaries
coverts
down
17.7.6 Bird feathers and flight lower region, the quill, embedded in the skin, and an upper
solid rachis. The rachis supports the vane, which consists of
The possession of feathers, and considerable rows of barbs on each side, which themselves bear two rows
modification of their skeleton has enabled most birds to of barbules. The barbules interlock with each other by
develop structural adaptations which have provided them means of small hooks and grooves (fig 17.50) thus
with the ability to fly, and gain substantial mastery of the providing a firm, strong and light, air-resistant surface to
air. the wing. At the junction between the quill and radius is a
Feathers are epidermal structures developed within the smaller tufted structure, the aftershaft or afterfeather. It
skin of birds, evolved from scales of reptiles. Whilst they consists of a group of separated barbs.
provide body covering for birds, they are not spread evenly
over the body surface. They occur in tracts called pterylae
(ptero, flying) which are separated by featherless areas
called apteria. Four distinct kinds of feather are recogni-
sable, namely the quill, contour, down and filoplume.
Details of the structure of each of these can be seen in fig
17.48a-c.
Each bird wing possesses 23 quill feathers called remiges.
They are positioned in a backward direction over the body
and overlap each other. This arrangement provides an
and
upper convex, and lower concave, surface to the wing
of
therefore enables it to act as an aerofoil, the significance
ed
which will be discussed later. There are 11 remiges attach hooklet on
calle d
to the hand (metacarpals and digits), these are
anterior row
of barbules
to the
primaries, whilst the remaining 12 are attached posterior row
(fig
forearm (radius and ulna) and are termed secondaries of barbules
retric es.
17.49). The quill feathers of the tail are called Vane of quill feather showing barbs and barbules
w Fig 17.50
Typically a quill feather is composed ofastiff hollo
657
Contour feathers, though similar in structure, are streamlined, pointed bill. The arrangement of the
smaller and more flexible than quill feathers, and their feathers provides a smooth surface for the air to flow
barbs are less firmly attached. They provide the main body over. The tail is short. This cuts down drag and
covering and wind-proof layer of the bird and establish its increases manoeuvrability. The legs of birds are tucked
body outline. up to the body during flight.
Down feathers are the only body covering of nestlings. In (4) The possession of a number of hollow spongy bones
older birds they are closely applied to the skin in amongst helps to lighten the body. Many of the cavities are filled
the contour feathers. The barbs of these feathers are long by air sacs. These also help to improve ventilation in
and pliable and their barbules do not possess hooks. the bird by being able to supply the lungs with fresh air
Filoplumes are small hair-like structures. There is no during inspiration and expiration (section 11.6.10).
vane as such, just a small tuft of flexible barbs and barbules The insides of the bones of large birds have struts for
at the tip of the rachis. These feathers are spread all over strength. The beak represents toothless, pointed jaws
the bird and help to retain air close to its body. covered with a horny bill. The absence of teeth
Apart from the development of feathers, many other precludes the need for heavy jaw muscles for grinding
modifications associated with flight have taken place within food. This function is carried out by the gizzard which
birds. The major ones are summarised below. is nearer to the centre of gravity of the bird. It has been
(1) The pectoral girdle is well developed and provides a estimated that in a pelican weighing 11 kg, and
firm base for the wings. There is a sizeable keeled standing 1.5 m tall, its skeleton weighs only 0.65 kg,
sternum providing a large surface area for the and that the frigate bird (a sea glider) witha wing span
attachment of the powerful pectoral flight muscles (fig of 2 m possesses a skeleton weighing just 110 g. In fact
17.51). Flight muscles may approach half the body here the feathers weigh more than the skeleton
weight in a strong flying bird such as a racing pigeon. (1.4-1.8 kg)! The sex organs of birds are very small,
(2 Considerable fusion and elongation of forelimb bones
—
only developing fully during the breeding season. This
has provided a large wingspan. Three carpals have again aids flight economy in terms of keeping the body
fused with three metacarpals to form the carpo- as light as possible.
metacarpus, a strong supportive structure for the (5) Birds have a high body temperature and good
attachment of the primary flight feathers. Only three insulation which combats the cooling effect of the air as
digits remain. Loss of the others has helped to lighten the birds move through it. It also ensures more efficient
the wing. The skeleton is rigid in places; for instance muscle contraction for flight. The heart of a bird is
some of the vertebrae are fused, and the vertebral large and strongly beating thus maintaining high blood
column is fused to the pelvic girdle. This transmits the pressure.
force of the wings more efficiently and provides better (6) Birds have excellent visual acuity. A third eyelid, the
lift. The hindlimbs have been lengthened, which means nictitating membrane, is present which removes par-
that the wings do not flap against the ground during ticles from the eye during flight. Many birds possess a
take-off. long flexible neck which enables the head to reach the
(3 —
The body is highly streamlined so that when the bird ground for feeding purposes, and permits it to be
flies through the air its shape provides little resistance. rotated, providing good all-round vision.
The skull is compact and generally possesses a (7) As would be expected, all birds that fly possess
remarkable powers of muscular coordination.
fulcrum pectoral
girdle
17.7.7 Flight in birds
Consider the horizontal flow of air over an
wing mn wing pulled inclined wing surface which has its leading edge raised
raised
above its trailing edge. In this context the wing acts as an
aerofoil. Air flowing over the upper surface meets less
resistance than that flowing over the lower surface (fig
17.52) and therefore develops a greater velocity. The
combined effect of these two phenomena is to cause air
pressure above the wing to be reduced, and that below to
be increased, thus providing the wing with a lifting force.
The power of this lifting effect depends on the size and
pectoralis
sternum
(keel) shape of the wing, the angle at which it is inclined to the
major long axis of the body (the angle of attack) and the bird’s air
Fig 17.51 Section of a bird (pigeon) skeleton viewed from speed. There is also a force acting on the bird, exerted by
the front to show how flight muscles and bones of the wings the air, which tends to push the wing backward in the
and pectoral girdle operate together direction of the airstream. This is called drag. The
658
Fig 17.52 , The bird wing acting as bird wing LIFT
an aerofoil
RESULTANT
reduced air
pressure (negative
> pressure)
a air flows over upper wing surface at a

flow of > greater velocity than over the lower surface
air
>
+ DRAG
>
\
increased
air pressure
(positive angle of attack
pressure)
mechanical efficiency of a wing depends on its ability to and twists like a propellor so that its front edge is tilted
develop a big lifting force for a small relative increase in upwards. In this position the wing develops lift for the
drag. body. The lifting effect of the air as it moves between the
There are three major types of flight; flapping, gliding primary feathers tends to separate them and bend them
and soaring, and hovering. upwards (fig 17.53).
The upstroke starts before the downstroke has been
Flapping flight completed. The inner part of the forearm is moved sharply
In a bird such as the pigeon, which beats its wings two times upwards and backwards with its front edge inclined above
per second, the main powerstroke is delivered by the that of the trailing edge. This action is produced by the
downbeat of the wings and is brought about by contraction pectoralis minor muscles attached to the dorsal surface of
of the large, powerful pectoralis major muscles which are the humerus and to the sternum. It provides lift for the
attached at one end to the ventral surface of the humerus body. As the wing moves upwards it bends at the wrist and
and to the keel of the sternum at the other. At take-off the the hand twists round so that the primary feathers are
first part of the downstroke is almost vertical with the suddenly jerked backwards and upwards until the whole
leading edge of the wing lower than the trailing edge. The wing becomes more or less straight above the body. On
primary feathers are bent upwards due to the pressure of their upward journey the primaries are separated, permit-
the air. They are tightly closed to provide maximum ting the air to pass between them thus minimising air
resistance to the air and therefore maximum lift. During resistance. The backward movement of the primaries
the latter part of the downstroke the wing is moved forward provides the bulk of the forward propulsive thrust
primaries -
for forward
movement
secondaries — for lift
es wing flexed
(bent) to
reduce
(a) resistance
wing tilts and

pushes air back
(i) beginning of causing forward -
downstroke (ii) movement (ili) upstroke
feather air passes

downstroke = 92 ———) between
ge Tee feathers,
o | _——. { Oxe ea ee
Pee st
i a y a resistance
upstroke
ee
Mh Zs hc
air resistance closes feathers
n ofwings during flapping flight in a pigeon. (bo)

Fig 17.53 Action of wings and feathers during flapping flight. (a)Positio
during flapping flight
Position of feathers during downstroke and upstroke of a bird wing
659
developed by the animal. Before the primaries have Fig 17.54 Comparison of the structure of the wing of a fast
flying bird (such as a swift) and a slow flying bird (such as a
reached their highest point, the pectoralis major muscles sparrow)
begin to contract again, eliciting a new downward beat and
the whole process is repeated. slow flier
During sustained flapping flight the wing action is

considerably modified and requires far less energy than
that needed at take-off. Here the wings do not flap so hard,
meet each other over the bird’s back or move forwards short wide wing
during the latter part of the downstroke. The wings are wingspan
generally held out straight, and the up and down flapping
movements occur mainly at the wrists (junction of lower
limb bones and carpals). The active backward lift of the eee
wrist has virtually disappeared and the upstroke is passive, secondaries
large te
being achieved by air pressure acting against the underside wing for lift
of the wing.
At the end of flight the bird lands by lowering and fast flier
spreading out its tail. This acts both as a brake andalifting
flap. As lift is achieved, the legs are lowered and the bird large hand
section for
comes to rest. The tail is also used to steer the bird in flight, forward
narrow wing
and stability in the air is accomplished by nervous control movement
for low drag
large
via the semicircular canals. Impulses are generated here wingspan
which stimulate accessory muscles to alter the shape and
position of the wings and the number of beats of one wing \ well-developed
primaries
relative to the other. 1 \
Different birds fly at different speeds. Such differences ’

inner wing
: NS
Xe
:
fewisecondaies
are achieved by the shape of the wing, its changing form
during flight and the frequency of the wing beat. A NB wing is flattened for minimum resistance
comparison of the structure of the wing ofa fast flier, the
swift, and that of a slow flier, the sparrow, is given in fig
17.54.
17 .9 ,
swift to fly so fast L
. s t t h e f e a t u r e s h en
i
the
able
Fig 17.55 Drawing

; showing forces
ee exerted against the bird when gliding in still air. A, A, direction of airflow over r bird;
bird; B, iv
force; C, weight of body; D, sinking force; E, drag; F, resultant aerodyne force; G, lift; H, direction of movement of bird. neers
(Modified after Gray.)
660
Gliding and soaring forward position. The new position of the wings and the
During gliding the wings are held out at more or less 90° to forward velocity of the body provide the bird with a lifting
the body and the bird gradually loses height. As it force which enables it to gain height in preparation for
descends, the force acting on it is called the driving force another rapid descent. Albatrosses are also able to glide for
(fig 17.55) and the pull of gravity acting in a downward long distances close to, and parallel to, wave ridges, using
direction is called the sinking force. As the gliding speed the small upcurrents of wind from the waves as land birds
increases so the lifting force increases and when lift equals do in slope soaring.
the sinking force, and the driving force equals drag, the Hovering
bird glides through the air at a constant velocity. The speed
and angle of the glide depends on the size, shape and angle Hovering is flapping the wings while at the same time
of attack of the wings and the weight of the bird. staying in one place. The wings move backwards and
Land birds make use of thermal upcurrents when gliding. forwards at speeds of up to 50 beats per second, and the
They occur when a horizontal wind is deflected upwards on upward thrust developed counterbalances the bird’s
weight. Birds that hover possess enormous flight muscles
encountering an obstruction such as a mountain (this is
known as slope soaring), or when warm air moves upwards relative to their size (3 body weight) and are consequently
and is replaced by colder air, such as over towns. Birds with very strong flappers. Their wings can tilt at almost any
angle. The majority of the wing feathers are primaries
light bodies and large wings such as buzzards and eagles are
(only six are secondaries) and are used for developing
good thermal gliders and they manoeuvre gently upwards
in a series of small circles. Rising in the air without flapping
thrust.
is called soaring.
17.7.8 Flight in bats
Marine birds, such as the albatross, possess a different
shape and glide in a different fashion (fig 17.56). The Bats are the only mammals capable of true
albatross has a large body, and very long narrow wings, and flight. Each wing is composed of a thin layer of heavily
makes use of gusts of wind that arise above the waves. vascularised skin, stretched between the bat’s forearm, all
During its upward glide it ascends to a height of about fingers of the hand except the thumb, the sides of its body
7-10 m. Then it turns downwind and descends at great and the ankle of the hindlimb. The wrist bones are fused
speed with its wings angled in a backward direction. At the and give the wing strength whilst those of the forearm and
bottom of the descent the albatross decribes an arc as it hand are considerably elongated and function as a
turns back into the wind and its wings are placed in a more supportive framework for the wing. The index and middle
slow gliders e.g. vultures, gliding position to lose height

buzzards, eagles, hawks
soaring position — wings stretched
out
broad wing for maximum lift
secondaries — many present for lift \____ primaries - spread out for extra lift
tail for balance
slow gliders are mostly land birds; they can

use light winds and weak currents of air
fast gliders e.g. albatross,

seagull, gannet large wingspan (e.g. albatross 3.0-3.4 m)
narrow sharp tip reduces

wing drag
long narrow wing

fast gliders are mostly sea birds; they can
use strong winds
gliders
Fig 17.56 A comparison between the body and wing shapes of slow and fast
661
bones of forelimb
fingers of the hand support the leading edge of the wing Fig 17.57 (above) Bat wing extended showing attachment of
whilst the other two fingers extend backwards to the wing to forearm, body and ankle
trailing edge which eventually links up with the fifth digit of
the ankle (fig 17.57). The actual movement of the wings
occurs in a similar fashion to the wings of birds, and flight Fig 17.58 (below) Running sequence in a dog (such as a
can be powerful and fast. greyhound)
(a) Backbone fully arched and feet immediately under the
body. (b) Backbone fully extended and somewhat concave,
17.7.9 Locomotion in quadrupeds, the dog limbs fully extended
Walking (a)
When a dog walks, its vertebral column remains rigid, and
forward movement is achieved by the activity of the
hindlimbs. They are moved forwards and backwards by
ZBfa A
alternate contraction of flexor and extensor muscles
respectively. () vertebral
pelvis column
When its extensor muscle contracts, each hindlimb,
acting as a lever, extends and exerts a backward force
against the ground, thus thrusting the animal forward and
slightly upwards. When the flexor contracts, the limb is
lifted clear of the ground and pulled forward. Only one
limb is raised at any one time, the other three providing a
tripod of support which balances the rest of the body. anki ewe ny) wrist
Beginning with the left forelimb in a stationary dog, the joint joint
sequence of leg movement is as follows when it walks
forward: left forelimb; right hindlimb; right forelimb; left knee elbow
hindlimb; and so on. Joint joint
662
Running Fig 17.59 Successive positions of the right leg during a
As a dog begins to run, it loses its tripod means of single pace
suspension and develops a type of movement where the 5 a 3 2 1
forelimbs move together, followed by the hindlimbs. The
feet are in contact with the ground for much less time than
in walking, and usually one forelimb touches the ground a
split second before the other. This also occurs with the
hindlimbs. Therefore the sequence of limbs touching the
ground is: left forelimb; right forelimb; right hindlimb; left
hindlimb.
As the dog reaches maximum speed, leg movement
quickens even further, and as they extend, all four legs may
be off the ground at the same time. The strong trunk
muscles arch the flexible backbone upwards when all four
limbs are underneath it, and downwards when the limbs are
fully extended. In this way the thrust of the limbs is
increased and the stride of the dog considerably leng-
thened, both of which enable the dog to increase its speed
(fig 17.58). over the right toe, backward pressure against the substra-
tum generally being exerted through the right big toe.
17.7.10 Bipedal gait, Man With the weight of the body now over the right leg, the
left heel is raised and the whole sequence repeated. This
In the standing position the weight of the body alternating sequence of right and left, heel-and-toe action
is balanced over two legs. When astride is taken by the continues until walking ceases.
right limb the first thing to happen is that the right heel is
raised by contraction of the calf muscle. This action serves 17.10 Why do sprinters generally run on
to push the ball of the right foot against the ground and thus their toes? — |
exert a forward thrust. The right limb pushes further
against the ground as it is pulled forwards, slightly bent at
the knee (fig 17.59). As this occurs, the weight of the body
is brought over the left foot which is still in contact with the 17.8 | Support in plants
ground and acting as a prop for the rest of the body. When
the right limb extends, the heel is the first part of the foot to Four tissues, the parenchyma, collenchyma,
touch the ground. The weight of the body is gradually sclerenchyma and xylem contribute towards the support of
transferred from the left side to a position over the right plants. A full account of their supporting role can be found
heel, and then as the body continues to move forwards, in chapter 8.
663
Chapter Eighteen
Homeostasis
An organism may be defined as a physico-chemical system organism with a degree of independence of the environ-
existing in a steady state with its external environment. It is ment which is determined by the effectiveness of the
this ability to maintain a steady state within a constantly mechanisms. Independence of the environment is used as a
changing environment that contributes towards the success criterion of the success of an organism and on this basis
of living systems. In order to maintain this condition mammals are seen as successful since they are able to
organisms, from the morphologically simplest to the most maintain relatively constant levels of activity despite
complex, have developed a variety of anatomical, phy- fluctuations in environmental conditions.
siological and behavioural mechanisms designed to achieve In order to achieve a degree of stability the activities of
the same end, that is the preservation of a constant internal organisms need to be regulated at all levels of biological
environment. The advantage of a constant internal en- organisation, from the molecular to the population. This
vironment in providing optimal conditions in which necessitates organisms employing a range of biochemical,
organisms can live and reproduce most efficiently was first physiological and behavioural mechanisms as are most
proposed by the French physiologist Claude Bernard in appropriate to their level of complexity and mode of life. In
1857. Throughout his research he had been impressed by all these respects mammals are seen to be better equipped
the way in which organisms were able to regulate than protozoa to cope with changes in environmental
physiological parameters, such as body temperature and conditions.
water content, and maintain them within fairly narrow Investigations have shown that the mechanisms of
ranges. This concept of self-regulation leading to physio- regulation found in organisms show many features in
logical stability was summed up by Bernard in the now common with mechanisms of regulation in non-living
classic statement, ‘La fixité du milieu interieur est la systems, such as machines. The systems of both organisms
condition de la vie libre.’ (The constancy of the internal and machines achieve stability by some form of control, to
environment is the condition of the free life.) which Wiener, in 1948, applied the term cybernetics
Bernard went on to distinguish between the external (cybernos, steersman). Cybernetics is the science of control
environment in which organisms live and the internal mechanisms and is commonly referred to as control theory.
environment in which individual cells live (in mammals, Plant and animal physiologists have used many of the very
this is tissue, or interstitial, fluid). He realised the precise mathematical models of control theory to explain
importance of conditions in the latter being continuously the functioning of biological control systems, hence before
stable. For example, mammals are capable of maintaining studying some of these self-adjusting mechanisms, such as
a constant body temperature despite fluctuations in the body temperature regulation, gaseous exchange rates,
external temperature. If it is too cold the mammal may blood metabolite levels and water and ionic balances, it is
move to warmer or more sheltered conditions; if this is not necessary to have some appreciation of the theoretical
possible, self-regulating mechanisms operate to raise the considerations underlying control systems.
body temperature and prevent further heat loss. The
adaptive significance of this is that the organism as a whole
will function more efficiently because its constituent cells 18.1 Control systems in biology
are maintained at optimum conditions. In this respect
biological systems are seen to operate not only at the level 18.1.1 Introduction to control systems
of the organism but also at the level of the cell. An
The rigorous application of control theory to
organism is the sum of its constituent cells and the optimum
functioning of the whole depends upon the optimum biology has led to a deeper understanding of the functional
relationships between the components of many physiolo-
functioning of its parts.
In 1932 the American physiologist Walter Canno
n gical mechanisms and has clarified many concepts which
stasis, were previously obscure. For example, living systems are
introduced the term homeostasis (homoios, same;
now seen to be open systems; that is they require a
standing) to describe the mechanisms whereby Bernard’s
. continuous exchange of matter between the environment
‘constancy of the internal environment’ is maintained
ity and themselves. Living systems are, in fact, in steady state
Homeostatic mechanisms function to maintain the stabil
provi de the with their environment and require a continuous input of
of the cellular environment and in so doing
665
REGULATOR
oven temperature by the use of a thermostat. In the electric
oven the control system includes an input (an electric
input—— detector reference point effector ——» output
current flowing through an element which acts as a source
of heat), an output (the oven temperature) and a
(set-point)
thermostat which can be set to a desired level, the

set-point. The thermostat acts as a modulator. If the
thermostat is set to read 150 °C, an electric current will
modulator
provide a source of heat which will flow until the oven
Fig 18.1 Basic components of a control system temperature passes the set-point of 150°C, then the
thermostat will cut out and no more heat will be supplied to
energy in order to prevent them coming to equilibrium with the oven. When the oven temperature falls below 150 °C
the environment. This equilibrium is achieved only upon the thermostat will cut in and the electric current will
the death of the organism when it becomes thermodynami- increase the temperature and restore the set-point. In this
cally stable with respect to its environment. The basic system the thermostat is functioning as an error detector
components of any control system are summarised in fig where the error is the difference between the output and
18.1. the set-point, The error is corrected by increasing the
The efficiency of the control system is measured in terms input. This is an example of a steady-state closed-loop
of how little displacement from the reference point system which is typical of many of the physiological control
(optimal level) occurs and the speed with which the level is mechanisms found in organisms.
restored. Homeostatic mechanisms must be free to Examples of biological negative feedback mechariente
fluctuate, as it is the fluctuations themselves which activate include the control of gas tensions in the blood, heart rate,
the control systems and return the parameter towards its arterial blood pressure, hormone and metabolite levels,
optimal level. Such control systems rely upon their water and ionic balances, the regulation of pH and body
components being linked together so that the output can be temperature. Fig 18.3 illustrates the role of negative
regulated in terms of the input, a concept known as feedback in the control of thyroxine release by the thyroid
feedback. In most feedback systems the output and input gland. In this example the modulator has three compo-
are identical. nents, a detector (the hypothalamus), a regulator (the
Feedback requires the action of the system to be referred pituitary gland) and an effector (the thyroid gland).
back to a reference point or set-point, which is the optimal Positive feedback is rare in biological systems since it
level of the parameter (or controlled variable), so that leads to an unstable situation and extreme states. In these
subsequent action may be modified to restore the set-point. situations a disturbance leads to events which increase the
There are two forms of feedback, negative and positive. disturbance even further (fig 18.2). For example, during
The former is most common in the homeostatic mechan- the propagation of a nerve impulse, depolarisation of the
isms of organisms. neuronal membrane produces an increase in its sodium
Negative feedback is associated with increasing stability permeability. Sodium ions pass into the axon through the
of systems (fig 18.2). If the system is disturbed, the membrane and produce a further depolarisation which
disturbance or error sets in motion a sequence of events leads to the production of an action potential. In this case
which counteract the disturbance and tend to restore the positive feedback acts as an amplifier of the response whose
system to its original state. The principle of negative extent is limited by other mechanisms as described in
feedback may be described in terms of the regulation of section 16.1.1.
further increase
/
p positive feedback
disturbance we corrective
A ¢ mechanism
»\
a
Increase K
eee
XN
\
steady-state normal situation steady-state

condition (no disturbance) condition
1H
negative feedback
4
/
decrease.
XN
disturbance , corrective
J mechanism
positive feedback
\
‘\
4
further decrease
>
Fig 18.2 Homeostatic control system. The directions of the lines on the diagram indicate the directions of stimulus and response
666
DETECTOR
(hypothalamus) INPUT
(thyroxine)
REGULATOR
Sea
oe) (thyroxine in blood vessels)
(pituitary gland)
ire
ley
le)
leleics!
ty
Felis
OUTPUT
EFFECTOR (thyroxine)
(thyroid gland)
Fig 18.3 A biological example of a simple control system, the control of thyroxine production. TRF, thyroid releasing factor;
TSH, thyroid stimulating hormone; see section 16.6
There are several control mechanisms in the body which nature and amounts of materials synthesised within the cell
are more complex than those previously described. are controlled by the rates of protein synthesis. Metabolic
Broadly they either involve the use of additional detectors activities in cells are determined by enzymes produced by
(physiological early-warning systems) or effectors (‘fail- the transcription and translation of the base sequence
safe’ systems) operating at different levels. For example, codes of DNA into the primary structure of proteins which
temperature detectors situated externally and internally act as enzymes. Regions of DNA carrying the code for a
enable homeotherms to maintain an almost constant ‘core’ specific protein are known as genes and the ‘switching on
body temperature. Temperature receptors in the skin, and off’ of genes is thought to be controlled by systems of
acting as disturbance detectors of changes in the external induction and repression which are described in detail in
environment, send impulses to the hypothalamus which section 22.7. In terms of control systems the maintenance
acts as a modulator and initiates corrective measures of a steady state within the cell depends upon the rates of
before any change in blood temperature occurs. Other supply and utilisation of cellular material, in other words
examples of this system include the control of ventilation the input and output, and the activity of the modulators (fig
during exercise, appetite and thirst. Similarly multiple 18.4).
detectors and effectors provide fail-safe mechanisms for Both single-celled organisms and cells of multicellular
many vital processes, such as regulation of arterial blood organisms control their internal environments as described
pressure where stretch receptors in the carotid sinus and above. In the case of single-celled organisms their
aorta, and baroreceptors in the medulla, respond to blood immediate environment is the external surroundings over
pressure changes and produce responses in various effec- which they have no control. They may, if they possess
tors including the heart, blood vessels and kidneys. Failure locomotory mechanisms, move to more favourable sur-
of any one of these is compensated for by the others. roundings, but for the most part they are at the mercy of the
environment. They adapt to, and tolerate, conditions as
best they can. The sheer numbers of these minute species
Gakic The nature and control of the internal reflect their degree of success in surviving in spite of their
environment structural simplicity.
The internal environment of an organism and The immediate environment of the cells of multicellular
its control may be considered at two levels: the cellular plants and animals is an extracellular fluid. This is called
level and the tissue level. sap in plants, haemolymph in insects, water in echinoderms
The cell is composed of cytoplasm whose constituents and tissue fluid in most other animals. The composition of
this extracellular fluid can be regulated by the organisms to
are modulated by the selective permeability of the cell
varying degrees depending upon the efficiency and effec-
membrane and by enzyme activity under the control of
tiveness of their homeostatic organs. In mammals the
protein synthesis. The cell membrane only permits certain
immediate environment of all living cells is tissue fluid. The
molecules to enter and leave the cell and the rates at which
mechanism of formation of this remarkably stable fluid is
exchange is permitted are strictly controlled by diffusion
described in chapter 14. Since the time of Bernard, the
gradients, osmotic gradients, electrical gradients, active
nature of the extracellular fluid and the mechanisms of its
mechanisms involving membrane-bound carrier systems
by control have been widely studied. It is birds and mammals
and changes in membrane distribution as shown
that show most control over the composition of the fluid
pinocytosis and phagocytosis (section 7.2.2). Similarly the
667
INPUT
(metabolite)
selective
uptake
MODULATOR
(cell membrane)
selective
secretion
OUTPUT
(metabolite)
———— peOUP (ee > OU TR Ue

TN eee ODA
(enzyme) (product)
(DNA) (protein synthesis) 1
| aT
Fig. 18.4 Summary diagram of the mechanisms involved in maintaining a steady state in the internal environment of a cell
and its parameters of water, gases, ions, nutrients, and produce local changes in pH which would affect the
hormones, excretory products, pH and temperature. In all rates of enzyme reactions. The body maintains a fine
cases these parameters have one or several tissues, organs control over the concentration (or tension) of carbon
or organ systems to maintain control within narrow limits. dioxide in the blood despite variations in oxygen availabil-
The mechanisms of control in most animals involve ity and metabolic demands which may increase twenty-fold
responses by endocrine glands, or the nervous system during vigorous activity.
coordinated by control centres in the brain and spinal cord The rate and depth of ventilation (inspiration and
(regulators) as shown in fig 18.5. expiration) are controlled by respiratory centres situated in
In conclusion it is worth re-emphasising the adaptive the pons and the medulla oblongata at the base of the brain
significance of homeostatic mechanisms. All metabolic (fig 18.6). These centres generate rhythmic impulses to the
systems operate most efficiently if maintained within diaphragm and intercostal muscles which bring about
narrow limits either side of optimal conditions. It is the role ventilation movements. This rhythm is basically involun-
of homeostatic organs and systems to operate both tary but can be overridden within limits by higher centres of
separately and together to buffer against fluctuations from the brain, as shown by the ability to hold the breath. The
these optimal conditions caused by variations in the rate and depth of ventilation directly governs the composi-
external and internal environments. Some of these tion of the alveolar air which, in turn, determines the
mechanisms are discussed in the following sections. oxygen and carbon dioxide tensions in the arterial blood
supplied to the body tissues. The mean partial pressures of
oxygen and carbon dioxide in alveolar air and the arterial
18.1.3 Control of respiratory gases in the blood are 100 mm Hg and 40 mm Hg respectively in a
blood
resting subject at sea level. The maintenance of the above
The cells of the body require a continuous levels is achieved by negative feedback to the respiratory
supply of oxygen from their immediate environment, the centres in the medulla of impulses from two types of
tissue fluid, in order to carry out respiration. Carbon receptors: mechanical stretch receptors (proprioceptors) in
dioxide, produced by this process, must not be allowed to the walls of the trachea and lungs, and chemoreceptors in
accumulate in the cells or in the tissue fluid, as it would the wall of the aorta, the carotid bodies (situated in the
upset the equilibria of the reactions involved in respiration walls of the carotid arteries) and in the medulla itself.
peripheral
internal nerves
=a
REGULATOR
autonomic
control centres INTERNAL
ENVIRONMENT sensory input
in the CNS ENVIRONMENT
external — glands ——__—_» hormones

negative feedback
STIMULUS—————_——— IN PUT ——Y_—_—_—__® MODULA TOR——————_»> OUTPUT————————————» RES PONSE
Fig 18.5 Homeostatic mechanisms for the control of the internal environment
668
higher centres trachea and lungs are no longer stimulated there is no
of brain
inhibition of the inspiratory respiratory centre and the
ventilation cycle repeats. During vigorous activity in-
creased carbon dioxide in the blood stimulates the
expiratory centre expiratory centre and impulses pass to the internal
respiratory
centres of medulla intercostal muscles which contract more frequently and this
medulla oblongata leads to deep or forced breathing.
The rate and depth of ventilation is governed by the
activity of chemoreceptors in response to changes in
branch of oxygen and carbon dioxide tensions in the blood. Experi-
vagus nerve
ments involving subjects breathing air composed of varying
concentrations of oxygen and carbon dioxide have shown
that the effects of increasing carbon dioxide levels are more
important in controlling ventilation than oxygen levels.
Reducing the oxygen concentration of air from 20% to 5%
carotid body produces a doubling of the normal resting ventilation rate,
whereas an increase of only 0.2% in carbon dioxide
a
C oie. | tco,
concentration produces the same response.
The influence of carbon dioxide concentrations and
ee lees
O
reduced blood pH in regulating ventilation rate is mediated
ho |S almost entirely through chemoreceptors in the aorta,
le Lit carotid bodies and in the medulla itself. Chemoreceptors in
| aortic body foo,
right left | pH the aorta and carotid bodies are also sensitive to changes in
aorta oxygen concentration and are vital when oxygen concen-
trations are low, because a fall in oxygen concentration
depresses activity of the medulla. Increased ventilation
facilitates loss of carbon dioxide from the blood by
diffusion into the alveolar air which is brought to a new
es
re
ee lower equilibrium with atmospheric air. The same effect is
: ——» neurones achieved by deliberate deep breathing or hyperventilation.
phrenic nerve
——-> blood vessels
diaphragm
external intercostal muscles * increased level 18.1 Explain why the ventilation rate
internal intercostal muscles i decreased level decreases and subjects feel dizzy and faint after
hyperventilating.
Fig 18.6 Summary diagram of mechanisms involved in the
control of respiratory gases in the blood
At high altitudes (in excess of 3 000m) the partial
pressures of oxygen and carbon dioxide in the atmosphere
are reduced. This results in the oxygen and carbon dioxide
Higher centres in the brain can override this feedback and
concentrations in the alveolar air coming to a new
voluntarily inhibit or increase the activity of the respiratory
equilibrium with those of the atmosphere. Consequently
centres, such as during breath-holding, forced breathing,
more carbon dioxide is lost from the blood into the alveolar
speech, singing, sneezing and coughing.
The effect of impulses from the stretch receptors 1s air due to the lower concentration here. As there is
mainly concerned with the mechanics of ventilation insufficient carbon dioxide in the blood to stimulate the
carbon dioxide chemoreceptors the stimulus for ventilation
movements. Impulses generated by the respiratory centres
cannot be carbon dioxide. Instead, low oxygen concentra-
pass down efferent neurones in the spinal cord; some
the tions stimulate chemoreceptors in the carotid body which
emerge in the cervical region as the phrenic nerves to
activates the respiratory centres.
diaphragm, other neurones emerge from the thoracic
There is a close relationship between the respiratory
region of the spinal cord as the external intercostal nerves.
The centres and the cardiovascular centre which are both
Together they produce the inspiratory movements.
of the lungs situated in the medulla. Changes in blood pressure, which
lungs inflate, and stretch receptors in the walls
which are maintained by the cardiovascular centre, affect the
and trachea activate afferent neurones of the vagus
centre, ventilation rate, for instance ventilation rate increases as
temporarily inhibit the inspiratory respiratory
propert ies the blood pressure falls and vice versa. Equally, changes in
preventing further inspiratory activity. Elastic
relaxes respiratory gas concentrations, as monitored by the
of the lungs produce recoil as the diaphragm
of air respiratory centres, will produce changes in blood pressure
reducing the volume of the thorax and thus expiration (section 14.12.6).
from the lungs occurs. As the stretch receptors in the
669
18.1.4 Control of metabolites in the blood demands imposed on tissues. The mechanisms of control
involve the activity of the autonomic system and details
One of the most important metabolites in the of these mechanisms are described in section 14.12.5.
blood whose level must be strictly controlled is glucose. There are two major conditions of abnormal heart rate,
This sugar is the principal respiratory substrate and must be tachycardia and bradycardia. Tachycardia (tachys, swift,
continuously supplied to cells. The brain cells are especially cardia, heart) is a general term describing an increased
sensitive to glucose and are unable to utilise any other heart rate and may be caused bya variety of factors
metabolites as an energy source. Lack of glucose results in including emotional states, such as anxiety, anger and
fainting. The normal level of glucose in the blood is about laughter, and overactivity of the thyroid gland. Severe
90 mg 100 cm~* blood, but may vary from 70 to 150 mg tachycardia is often the result of changes in the electrical
100 cm™ blood during fasting and following a meal, activity of the heart. Regions of the heart, other than the
respectively. The sources of blood glucose and its metabo- S—A node, can act as foci for the origin of the stimulus
lic interrelationships with other metabolites are described producing contraction of the heart muscle and augment the
in section 18.5.2. normal heart activity, based on the S-A node, producing
The control of blood glucose level is an example of a tachycardia.
complex, endocrine-regulated homeostatic mechanism Bradycardia (bradys, slow) describes the condition
involving the integrated secretion of at least six hormones where the heart rate is reduced below the mean level. It is
and two negative feedback pathways. A rise in blood common in athletes who develop an increased stroke
glucose level (hyperglycaemia) stimulates insulin secretion volume as a result of training. Therefore, in order to
(section 16.6.7) whereas a fall in blood glucose level maintain a constant cardiac output at all times their resting
(hypoglycaemia) inhibits insulin secretion and stimulates heart rate is reduced. Underactivity of the thyroid gland,
the secretion of glucagon (section 16.6.7) and other hor- and changes in the electrical activity of the S-A node and
mones which raise blood glucose levels (hyperglycaemic other conductile regions of the cardiac tissue, can also give
factors). The control mechanism is summarised in fig 18.7. rise to bradycardia.
The antagonistic regulatory effects of the sympathetic
18.1.5 Control of heart rate and blood pres- and parasympathetic nervous systems on the S-A node
sure
tend to counteract temporary conditions of tachycardia and
Variations in heart rate and blood pressure bradycardia and restore the normal heart rhythm.
have an indirect effect upon the composition of tissue fluid Blood pressure is the force developed by the blood
and enable it to be kept within narrow limits despite pushing against the walls of the blood vessels. It is usually
fluctuations in the external environment and varying measured in the brachial artery using a sphygmomano-
MUSCLE
uptake of glucose by cells
insulin secreted
(regulator) a glucose ——» CO, + H,O
<> glucose ——» glycogen
glucose ——» fat
islets of Langerhans (effector)
(detector) > fallin glucose level
of
rise in glucose level
Te Ray
$f ~Redbag
diet (input) =
normal blood normal blood

glucose level glucose level tissues
(set-point) (set-point) (output)
assimilation —_ —_
ae
(input) —
fall in glucose level eae @
feedba k
4 ae
rise in glucose level
islets of Langerhans,
adrenal cortex and
medulla, hypothalamus LIVER
(detectors) glycogen——_ glucose
protein ————» glucose
glucagon, glucocorticoids, (effector)
adrenaline, growth hormone
and thyroxine secreted Fig 18.7 Summary diagram of the dual
(regulator)
control of blood glucose level
670
meter. The systolic pressure is produced by the contraction temperature on surface of Sun 6000-4]
of the ventricles and the diastolic pressure is the pressure in |
|
the arteries when the ventricles relax. Blood pressure is |
|
affected by age, sex and state of health, and the mean water boils (100 °C)
pressures for a healthy young man are 120 mm Hg (systolic)
and 80 mm Hg(diastolic). Both pressures are affected by
cardiac output and peripheral resistance and indicate the
general state of heart and blood vessels. Conditions which proteins denature (50 °C) 50
lead to narrowing and hardening of the arteries (atherosc- body temperature in Man (37 °C) 40
lerosis) or damage to the kidneys may increase the blood
pressure, a condition known as hypertension, and impose a
strain on the heart and blood vessels. This may lead to a water freezes (0 °C) 0
weakening of artery walls and their rupture, or the clogging
of narrowed vessels by blood clots (thrombosis). These are of
life
animal
range
-30
very serious if they affect the brain or the heart and lead to of
life
plant
range
cerebral haemorrhage (stroke), cerebral thrombosis or
coronary thrombosis.
18.1.6 Control of infection

The many mechanisms of defence against
infection are homeostatic in that they attempt to maintain a
constant internal environment. The skin is the major
all molecular motion ceases res
defence against infection, but viruses, bacteria, protozoa,
round worms and flatworms can invade the tissues through
natural openings, cuts and by boring through the skin. The Fig 18.8 Temperature reference points in living organisms
presence of pathogens and their toxic waste products in the and the ranges of temperature they are able to tolerate
body evoke several defence mechanisms including’ the
activity of various white blood cells (phagocytes, mono-
cytes and lymphocytes). The mechanisms of blood clotting, locked up in the chemical bonds of synthesised organic
wound healing and immunity are described in detail in molecules (chapter 9). This provides an endogenous
sections 14.13.5 and 14.14.) (within the body) source of heat energy when released by
the catabolic reactions of respiration (chapter 11).
Temperature indicates the amount of heat energy in a
18.2 Temperature regulation
system and is a major factor determining the rate of
chemical reactions in both living and non-living systems.
Heat is a form of energy with an important
As described in section 6.4.3, heat energy increases the rate
influence upon the maintenance of living systems. All living at which atoms and molecules move and this increases the
systems require a continuous supply of heat in order to probability of reactions occuring between them. The
prevent degradation and cessation of those systems. relationship between temperature and the rate of chemical
The major source of heat for all living organisms is the reactions is expressed as the temperature coefficient, or
Sun. Solar radiation is converted into an exogenous Q,,. In living systems temperature has an effect upon
(outside the body) source of heat energy whenever it enzyme structure which influences the rate of metabolism
strikes and is absorbed by a body. The extent and effect of and exerts the major influence on the distribution and
this solar radiation depends upon geographical location, activity of organisms.
and is a major factor in affecting the climate of a region.
This, in turn, determines the presence and abundance of 18.2.1 The influence of temperature on the
species. For example, organisms inhabit regions where the growth and distribution of plants
normal air temperatures vary from —40 °C, asin the Arctic,
Temperature can act as a limiting factor in
to 50 °C in desert regions. In some of the latter regions the
the growth and development of plants by influencing the
surface temperature may rise as high as 80 °C. The
rates of cell division, cell metabolism and photosynthesis.
majority of living organisms exist within confined limits of
The non-light-requiring reactions of photosynthesis are
temperature, say 10-35 °C, but various organisms show
temperature dependent and control the various metabolic
adaptations enabling them to exploit geographical areas at
pathways described in chapter 9. The rates of photosyn-
both extremes of temperature (fig 18.8). This is not the only
of thesis, and the accumulation of sufficient food materials to
source of heat available to organisms. Solar radiation
enable the plant to complete its life-cycle, are factors
wavelengths within those of the visible spectrum are used
determining the geographical range of plants.
by photosynthetic autotrophs and this energy becomes
671
18.2.2 Adaptations to low temperatures in overheating. It is the aerial parts of plants which are
plants exposed to the heating effect of solar radiation, and the
largest exposed surface is that of the leaves. Leaves
The flora of northern temperate climates and characteristically are thin structures with a large surface
the tundra show many adaptations enabling plants to take area to volume ratio to facilitate gaseous exchange and
maximum advantage of the short, warm summers. For light absorption. This structure is ideal for preventing
example, the only plant species found are mosses, lichens, damage by excessive heating. A thin leaf has arelatively
a few grasses and fast-growing annuals. Plants living in low heat capacity and therefore will usually assume the
extreme northerly or southerly latitudes are subjected to temperature of the surroundings, a phenomenon known as
long periods of adverse conditions, such as low light heterothermism. This effect is offset in hot regions by the
intensity, low temperatures and frozen soil. In order to development of a shiny cuticle secreted by the epidermis
survive in these conditions, plants shown many anatomical,
and which reflects much of the incident light, thus
physiological and behavioural adaptations which may be preventing heat being absorbed and overheating the plant.
related to stages in the life cycle. For example, most The large surface area contains numerous stomatal
temperate woody perennials are deciduous and lose their openings which permit transpiration. As much as
leaves, under the influence of the plant growth regulator of water may be lost from plants by
0.5kgm~*h!
substance abscisic acid (ABA), in order to prevent water transpiration in hot, dry weather, which would account for
loss by transpiration and evaporation during periods when the loss of approximately 350 W m~ in terms of heat
soil water uptake is limited by low temperatures (section energy. This is nearly half the total amount of energy being
15.2.8). Wind and snow damage is also avoided by the absorbed. As a result of these mechanisms plants are able
shedding of leaves during these periods when the rate of to exercise a considerable degree of control over their
photosynthesis would be severely limited by low light temperature. This ability is closely coupled to the relative
intensities, low temperatures and unavailability of water humidity of the air.
and salts. Throughout these periods the regions of next
year’s growth, the buds, are protected by scale leaves and
their metabolic activity is suppressed by the presence of a 18.2 Why do plants suffer permanent
growth regulator substance called dormin. This is so called physiological damage if exposed to temperatures in
because of its effects. Recent investigations, however, have excess of 30 °C when the humidity is high?
shown this to be abscisic acid. Many coniferous species
dominate the vegetation of the more temperate regions,
During hot, sunny days many plants in danger of
particularly in northern latitudes. These species have
overheating show a phenomenon known as ‘photosynthetic
needle-like leaves which reduce the amount of snow which
slump’. This is thought to result from a temporary cessation
can accumulate on them in winter and havea thick cuticle
of metabolic activity as a result of changes in enzyme
to prevent water loss in summer. Many species of annuals
structure, or the closure of stomata as a result of a build-up
have short growing periods and survive the winter by
of respiratory carbon dioxide within the guard cells. It is
producing resistant seeds or organs of perennation. Lower
possible, however, that the reason for closure of the
plants survive by producing resistant cysts which may
stomata is simply one of transient lack of water in the leaf.
remain viable for hundreds of years, as demonstrated by
Wilting is a common response to high temperatures and
cysts which have germinated following their removal from
the permafrost of Siberia. reduced water uptake. An imbalance between the rates of
Low temperatures are required by many plant species in transpiration and water uptake results in an overall loss of
order to break dormancy and initiate vernalisation. For turgidity by those plant cells lacking a thickened cell wall,
example, lilac buds develop more quickly after being such as parenchyma. The adaptive significance of this
response is to reduce the leaf surface area which is exposed
exposed to low temperatures than to high temperatures.
to direct sunlight, thereby preventing overheating. Wilting
Other examples of the effects of low temperature on plant
may be observed in plants growing in greenhouses in
growth are described in chapter 15 and section 21.6.
response to the very high temperatures which develop in
the leaves, even if adequately supplied with water. Once
18.2.3 Adaptations to high temperatures in
the temperature begins to fall the plants recover very
plants
quickly even if the degree of wilting looks severe.
In many regions of the world high tempera- Plants living in dry conditions are called xerophytes and
tures are associated with water shortage, and many of the show many morphological adaptations which enable them
adaptations shown by plants in these regions are related to to survive (section 19.3.2). In most cases these adaptations
their ability to resist desiccation and their need to lose are primarily concerned with regulating water loss,
water in order to cool themselves. although the characteristic needle-shaped leaves permit
Plants are unable to escape high temperatures by moving maximum heat dissipation. The mechanisms for withstand-
to shaded areas and therefore they have to rely on ing high temperatures, on the other hand, are mainly
morphological and physiological adaptations to avoid physiological.
672
18.2.4 The influence of temperature on the 40; cat :
growth and distribution of animals
Man
Temperature influences the metabolic activity
30L platypus
of animals and plants primarily through its influence on
enzymes and the mobility of atoms and molecules. This spiny
directly affects the rate of growth of animals (chapter 21). ant-eater
In addition, temperature may also affect the geographical 20+

distribution and ecological preferences of animals through
its influence on plants as primary producers in the food
chains. The ecological range of most animal species, with
(rectal)/°C
Body
temperature
the exception of some insects, birds and mammals able to
migrate, is determined by local availability of particular
food materials. Restricted powers of locomotion limit
10 20 30 40
other organisms to feeding on readily available food External temperature/°C
sources which are determined by the trophic relationships
existing in the ecosystem. Fig 18.9 The relationships between external and internal
The variety of responses shown by animals to tempera- temperatures in vertebrates kept for 2h at the temperatures
indicated
ture depends upon the degree of thermal stability shown by
the environment and the degree of control the organism is
able to exercise over its own body temperature. Life is
believed to have originated in the marine environment, the
environment that poses fewest problems to living organ-
isms. Temperature fluctuations in aquatic environments
are slight and the biological significance of this is that are unable to maintain their body temperature within
aquatic organisms occupy arelatively stable environment narrow limits. Consequently these animals are described as
with regard to temperature. Most of these organisms, poikilothermic (poikilos, various; therme, heat). Alterna-
including invertebrates and fish, have a body temperature tively, because they rely more on heat derived from the
which varies according to the temperature of the water, environment than metabolic heat in order to raise their
though some very active fish such as the tuna are able to body temperature, they are termed ectothermic (ecto,
maintain body temperatures in excess of that of the water. outside). These animals used to be described as ‘cold-
However, aquatic animals rarely need to exhibit the blooded’ but this is a misleading and inaccurate term.
physiological and behavioural responses shown by terrest- Birds and mammals are able to maintain a fairly constant
rial animals because the physical properties of the water body temperature independently of the environmental
buffer them against sudden and extreme fluctuations. temperature. They are described as homeothermic (or
Water has a maximum density at 4 °C and consequently homoiothermic — homoios, like) or less correctly as
only the surface of the water freezes at 0 °C as ice floats. ‘warm-blooded’. Homeotherms are relatively independent
This enables many aquatic animals to continue to be active of external sources of heat and rely on a high metabolic rate
at times when most terrestrial organisms would be inactive to generate heat which must be conserved. Since these
due to sub-zero conditions. animals rely on internal sources of heat they are described
Air has a relatively low specific heat and therefore air as endothermic (endos, inside). Some poikilotherms may,
temperature can fluctuate widely over a 24 h period. One at times, have temperatures higher than those of
of the problems associated with the colonisation of land by homeotherms and in order to prevent ambiguity, the terms
animals was adapting to, or tolerating, these temperature ectotherm and endotherm are used throughout the text.
fluctuations. This has produced many physiological, beha- Fig 18.9 shows the abilities of several vertebrates to
vioural and ecological responses, as well as being a major regulate their body temperature in various external
determinant of geographical distribution. The nature of temperatures.
these responses and examples are described in subsequent
sections. 18.2.6 Body temperature
Whenever reference is made to body tempera-
18.2.5 Sources of heat for animals ture in animal studies it usually refers to the core
All animals derive heat from two sources, the temperature. This is the temperature of the tissues below a
external environment and from the release of chemical level of 2.5 cm beneath the surface of the skin. This
energy within their cells. The extent to which animals are temperature is normally determined by taking the rectal
able to generate and conserve this heat depends upon temperature. Temperatures in other regions of the body
can vary tremendously depending upon position and the
physiological mechanisms associated with their phylogene-
external temperatures (fig 18.10).
tic position. All invertebrates, fish, amphibia and reptiles
673
38
36
Dale,
Temperature/°C
29.0
external temperature (19 °C)

27.0
| La ! | =|
=
25D 0 10 20 30 40 50 60 70 80 90 100
20.5 Distance from top of pelvis/cm
Fig 18.10 Skin temperatures recorded at various distances from point X at the top of the pelvis. The limb was exposed to an
external temperature of 19°C throughout the period of recording. The low temperature at the knee cap probably reflects the
poorer blood supply to this organ
Whilst the main source of gaining heat differs between Evaporation. Heat is lost from the body
ectotherms and endotherms, the methods of heat transfer surface during the conversion of water to water vapour.
between organism and the environment are the same in The evaporation of 1 cm’ of water (1 g) requires the loss of
both types of organism and these are radiation, convection, 2.45 kJ from the body. Water loss by evaporation takes
conduction and evaporation. Heat can be transferred in place continuously through the skin and from the lungs, as
either direction by the first three methods depending upon expired air, and this insensible water loss cannot be
the thermal gradient (the direction of the temperature controlled. It is a limiting factor in the distribution of many
difference from hot to cold) but can only be Jost from plant and animal species. Regulated evaporative heat loss is
organisms by the latter method. controlled and involves water loss by sweating and panting.
Radiation. Heat is transferred by electro-
magnetic waves which lie in the long-wave, infra-red region
of the electromagnetic spectrum, beyond the visible 18.3 | Ectothermic animals
spectrum. Bodies do not radiate heat to air, since it is
unable to absorb much radiant heat, but transfer it to other The majority of animals are ectothermic, and
bodies at a rate which is proportional to the temperature their activity is determined by the prevailing environmental
difference between the two bodies. Radiation accounts for temperature. The metabolic rate of ectotherms is relatively
about 50% of the total heat loss in Man and provides the low and they lack mechanisms for conserving heat. Aquatic
main route for controlled heat loss in animals. ectotherms live within a restricted temperature range, or
zone of tolerance, determined by the size of the body of
Convection. Heat is transferred between
water in which-they live. For example, the temperature of a
organism and environment by passage through the air. In
pond can vary considerably throughout the year, whereas
the case of endotherms, where the ambient (air) tempera-
that of an ocean may change by only a few degrees. Despite
ture is usually lower than body temperature, the air in
this wide temperature fluctuation in small bodies of water,
contact with the organism rapidly becomes warm, rises, .
many insect species have aquatic larval and pupal, or
and is replaced by cooler air. The rate of heat transfer by
nymphal, stages, since the aquatic temperatures are more
this method is linked to the rate of air movement which
stable and less extreme than terrestrial temperatures
continually brings cooler air into contact with the body. It
during the winter months. Mayflies, dragonflies, caddis-
may be reduced by materials covering the skin such as
flies, midges and mosquitoes all have an aquatic diapause
feathers, fur, hair and clothing.
(section 21.10.1).
Conduction. Heat is transferred by physical Aquatic invertebrates are able to tolerate greater
contact between two bodies, for instance the organism and temperature fluctuations than aquatic vertebrates due to
the ground. Heat exchange by this means is relatively their relatively simple morphology and physiology. Fish
insignificant for most terrestrial organisms, but may be have a higher rate of metabolism than aquatic invertebrates
considerable for aquatic and soil-dwelling organisms. but the majority of this endogenous heat is rapidly
674
dissipated around the body and lost to the environment by
If the temperature becomes too high, they move into the
conduction through the gills and skin. Consequently, fish
water which is relatively cooler. Conversely, at night they
usually have a body temperature which is at thermal
retreat to water in order to avoid the low temperatures
equilibrium with that of the water. Fish cannot maintain a which would be experienced on land. Thermoregulation
temperature below that of the water but may in some cases, mechanisms have been studied most extensively in lizards.
as in the case of tuna fish, retain heat by means of a Many different species of lizard have been studied and
countercurrent heat-exchanger system. This can raise the show a variety of responses to different temperatures.
temperature of the ‘red’ swimming muscle to about 12 °C Lizards are terrestrial reptiles and exhibit many behaviou-
above that of the sea water. ral activities in keeping with other ectotherms; but some
Terrestrial ectotherms have to contend with greater species employ a number of physiological mechanisms
temperature fluctuations than those of aquatic ectotherms, enabling them to raise and maintain their body tempera-
but they have the benefit of living at higher environmental tures above that of the environment, an example is the
temperatures. This allows them to be more active and show Australian monitor (Varanus). Other species are able to
a variety of complex behavioural patterns based upon the keep their body temperature within confined limits by
prevailing temperature conditions. Many species are able varying their activity and taking advantage of shade or
to maintain temperatures slightly above or below air exposure. In both these respects lizards foreshadow many
temperature and thereby avoid extremes. The relatively of the mechanisms of homeothermy shown by birds and
poor thermal conductivity of air reduces the rate of heat mammals.
loss from organisms whilst water loss by evaporation may Surface temperatures in desert regions can rise to
be used to cool the organism. 70-80 °C during the day and fall to 4 °C at dawn. In these
circumstances most lizards seek refuge, during these
extremes, by living in burrows or beneath stones. This
18.3.1 Temperature regulation in terrestrial response and certain physiological responses are shown by
ectotherms the horned lizard (Phrynosoma) which inhabits the
Heat is gained and lost by terrestrial deserts of the south-west of the USA and Mexico. In
ectotherms by behavioural and physiological activities. addition to burrowing, the horned lizard is able to vary its
The main sources of heat gain are the absorption of solar orientation and colour, and as the temperature becomes
radiation and conduction from the air and the ground. The high it can also reduce its body surface area by pulling back
amount of heat absorbed depends upon the colour of the its ribs. Other responses to high temperatures involve
organism, its surface area and position relative to the Sun’s panting, which removes heat by the evaporation of water
rays. For example, a species of Australian grasshopper is from the mouth, pharynx and lungs, eye bulging and the
dark in colour at low temperatures and absorbs solar elimination of urine from the cloaca.
radiation thus heating up rapidly. As the temperature rises The moist skin of amphibia provides an ideal mechan-
above a set point, further absorption is reduced by the ism to enable heat to be lost by evaporation. This water
cuticle lightening in colour. This colour change is believed loss, however, cannot be regulated physiologically as in
to be a direct response shown in pigment cells to body mammals. Amphibia lose water immediately on exposure
temperature. These organisms are known as _ basking to dry conditions and, whilst this aids heat loss, it would
heliotherms (helios, sun). lead to dehydration if the amphibia did not find moist
Changes in orientation of the organism relative to the shaded conditions where the rate of evaporation would be
Sun’s rays vary the surface area exposed to heating. This reduced. The marine iguana (Amblyrhynchus), a reptile,
practice is common in many terrestrial ectotherms includ- normally maintains a temperature of 37 °C as it basks on
ing insects, arachnids, amphibia and reptiles. It is a form of the rocky shores of the Galapagos Islands, but it needs to
behavioural thermoregulation. For example, the desert spend a considerable time in the sea feeding on seaweed at
locust (Schistocerca) is relatively inactive at 17 °C but by a temperature of approximately 25 °C. In order to avoid
aligning itself at right-angles to the Sun’s rays it isable to losing heat rapidly when immersed in water the iguana
absorb heat energy. As the air temperature rises to reduces the blood flow between surface and core tissues by
approximately 40 °C it re-orientates itself parallel to the slowing its heart rate (bradycardia).
Sun’s rays to reduce the exposed surface area. Further
increase in temperature, which may prove fatal, is 18.4 Endothermic animals
prevented by raising the body off the ground or climbing up
vegetation. Air temperature falls off rapidly over short Birds and mammals are endothermic and their
distances above ground level and these movements enable activity is largely independent of prevailing environmental
the locust to secure a more favourable microclimate temperatures. In order to maintain a constant body
(section 12.4.4). temperature, which is normally higher than the ambient
Crocodiles too, regulate their body temperature on land (air) temperature, these organisms need to have a high
by varying their position relative to the Sun’s rays and metabolic rate and an efficient means of controlling heat
opening their mouths to increase heat loss by evaporation. loss from the body surface. The skin is the organ of the
675
body in contact with the environment and therefore dermis, that overly the subcutaneous tissue which contains
monitors the changing temperatures. The actual regulation specialised fat-containing cells known as adipose tissue.
of temperature by various metabolic processes is controlled The thickness of this layer varies according to region of the
by the hypothalamus of the brain. body and from person to person.
18.4.1 Skin Epidermis. The cells of this region are

The term, skin, applies to the outer covering ectodermal in origin and are separated from the dermis by a
of vertebrate animals and should not be used to describe basement membrane. The epidermis is composed of many
the outer coveriug of annelids, molluscs or arthropods. The layers of cells forming a. stratified epithelium (section
skin is the largest organ of the body and is made up of 8.3.2). The cells above the basement membrane are cuboid
connective tissue, blood vessels, sweat glands and sense epithelial cells (section 8.3.1) and form an actively dividing
cells. These enable it to fulfil a great many different region known as the Malpighian layer. By repeated
functions. The structure of the skin varies in different division of its cells, this region gives rise to all of the cells of
vertebrate groups and the major differences will be the epidermis. The Malpighian layer forms the lower
discussed after the basic structure of human skin has been region of the stratum granulosum, which is composed of
described. living cells becoming flatter as they approach the outer
region of the epidermis, the stratum corneum. Cells in this
Human skin structure region become progressively flattened and synthesise
The skin is composed of two main layers, the epidermis and keratin, which is a fibrous sulphur-containing protein
(b)
erector pili muscle stratum stratum
f granulosum corneum
=
ee
Eat
EC
epidermis {
region of
elastic fibres U3 Malpighian
li layer
and collagen
fibres |
hair follicle
dermis
blood vessel ee
sebaceous
glands
part of hair
shaft
secretory
cells
hypodermis
subcutaneous _| sweat duct

fat cells
dermal
papilla
Fig 18.11 (a) VS through human skin. (b) Diagram of skin

structure based on (a)
676
which makes the cells waterproof. As the keratin content of glands by blood capillaries, and the secretory activity of the
the cells increases they become cornified, their nuclei glands is controlled by the activity of the sympathetic nerve
disappear and the cells die. The thickness of the stratum fibres. There are two types of sweat glands, eccrine and
corneum increases in parts of the body where there is apocrine. The former are the most common (approximate-
considerable friction, such as the ball of the foot and the ly 2.5 x 10° in Man), being found in most regions of the
bases of the fingers. The outer covering of the skin forms a body, and the latter are found in the arm pits, around the
semi-transparent, thin, tough, pliant, waterproof covering nipples, the pubic region, hands, feet and anus. These
pierced by pores, which are the openings of the sweat release an odourless fluid, which may later produce an
glands, and by the hair follicles. The outermost squamous unpleasant odour due to bacterial activity in this fluid.
epithelial cells are continually being shed as a result of Certain zinc and aluminium compounds can inhibit the
friction. activity of the glands and destroy bacteria. Most anti-
The stratum corneum has become modified in many perspirants and deodorants contain these compounds.
vertebrates to produce nails, claws, hooves, horns, antlers, Blood capillaries are numerous in the dermis and supply
scales, feathers and hair. Keratin is the main component of the various structures already described. Many of the
all these structures. capillaries form loops and have shunts (fig 18.12) which
enable the body to vary the amount of blood flowing
Dermis. Most of the cells of the dermis are through the cutaneous capillaries. This is one of the many
mesodermal in origin. The dermis is a dense matrix ways of regulating body temperature, as described in
composed of connective tissue rich in elastic fibres and section 18.4.3.
containing blood capillaries, lymph vessels, muscle fibres, Motor neurones innervate the muscles and glands in the
sensory cells, nerve fibres, chromatophores (pigment dermis whilst sensory neurones carry nerve impulses from
cells), sweat glands and hair follicles. the many sense cells situated in the dermis. These sense
Hair follicles are epidermal invaginations with a root cells detect heat, cold, touch, pain and pressure. Some of
hair, or papilla, at the base from which the hair shaft the sense cells are simple and consist of free nerve endings,
develops. Hair is composed of cuboid epithelial cells which whereas others, such as the Pacinian corpuscle, are
become cornified by impregnation with keratin. The outer encapsulated.
cortex of the hair contains varying amounts of the pigment
melanin which determines hair colour. The medulla of the
hair may contain air bubbles, and as the number increases Vertebrate skin structure
with age and melanin production falls off the hair becomes The skin of other vertebrates differs in various ways from
grey. Blood capillaries supply the growing hair with the structure described above. Fish have well-developed
nourishment and remove waste substances. The upper part scales or dermal plates and mucus-secreting glands.
of the hair projects beyond the epidermis and is kept supple Amphibia have a smooth skin which is made slippery by the
and prevented from becoming wetted by sebum, an oily secretion of mucus from the many glands in the dermis.
secretion produced by sebaceous glands which open into This mucus is watery and keeps the skin moist to facilitate
the hair follicle. Sebum contains fatty acids, waxes and gaseous exchange. In some species, such as toads, the
steroids, and spreads along the hair and onto the skin secretion is distasteful to predators. The dermis contains
where it keeps the follicle free from dust and bacteria, as numerous pigment cells enabling amphibia to alter their
well as forming a thin waterproof layer over the skin. This colouration to blend in with the background, and also to
not only prevents water loss from the skin but also prevents absorb or reflect solar radiation. There are many subcu-
water entering the skin. taneous lymph spaces and the skin is only attached to the
At the base of the follicle is a smooth muscle, the erector underlying muscles at intervals which makes the skin
pili muscle which has its origin on the basement membrane extremely loose.
and its insertion on the hair follicle. Contraction of this Reptiles have horny scales covering the body and these
muscle alters the angle between the hair and the skin which form an impermeable layer. The skin is always dry, and the
results in variation of the amount of air trapped above the numerous chromatophores in the epidermis enable reptiles
skin. This is used as a means of thermoregulation and also to alter their colouration. Birds have developed feathers
as a behavioural response to danger in some vertebrates. which are outgrowths from the epidermis. There are
When the hair ‘stands on end’ it increases the apparent size different types of feathers carrying specific functions
of the organism which may be sufficient to frighten off (section 17.7.6) but they all contribute to preventing heat
would-be attackers. The distribution of hair in humans in loss from the body. Birds maintain relatively high body
much more restricted than in other mammals. temperatures, about 40—43 °C, and as they have a large
Sweat glands are coiled tubular exocrine glands situated surface area to volume ratio they need an efficient means of
preventing heat loss.
in the dermis and connected to sweat pores by a duct. They
are found over the entire body in Man, but in some All mammals possess sweat glands and hair, and the
mammals are restricted to the pads of the feet. They are number and distribution of these two structures reflect the
absent in birds. Water, salts and urea are brought to the
efficiency of control of heat balance.
677
Functions of mammalian skin energy comes from active tissues such as the liver and
voluntary (skeletal) muscle. The rate of energy release is
Skin has four main functions. Temperature regulation is
regulated by other factors such as environmental tempera-
described in detail in section 18.4.3. Further functions are
ture and hormones. Thyroxine, released from the thyroid
as follows. rate and therefore heat
gland, increases the metabolic
Protection. The skin protects the internal production. The effects of this hormone are long term,
organs from damage, dehydration and disease. The whereas the effects of adrenaline are mostly short-lived.
cornified epidermis prevents damage by friction, particu- Other sources of metabolic heat energy are initiated by
larly in those regions where pressure and chafing develop. nerve impulses. Repeated stimulation of voluntary muscle
The dermis and subcutaneous tissues prevent mechanical by somatic motor neurones produces the shivering res-
damage from bumps and knocks. Melanin, the dark ponse which can increase heat production by up to five
pigment found in chromatophores, protects the body from times the basal level. During shivering various groups of
excessive harmful ultra-violet light. Variations in hair or muscle fibres within a muscle contract and relax out of
fur pigment, giving patterns, provide camouflage against phase, so that the overall response is an uncoordinated
the background of the natural habitat. In several species, movement. This response may be reinforced by other
such as stoat and the arctic hare, the overall colour changes muscular activity such as rubbing the hands together,
according to the season and thus helps to protect the stamping the feet and limited forms of exercise. In many
organism from attack by predators or recognition by its mammals there are areas near the thoracic blood vessels
prey. The cornified layer and sebum prevent uncontrolled which are rich in brown fat cells; stimulation-of these by
water loss by evaporation. Sebum also prevents the fur of sympathetic neurones causes the rapid metabolism of the
aquatic mammals, such as the otter, from wetting, which numerous fat droplets in the cells. The subsequent release
would destroy the insulatory properties of the air trapped of energy by enzymes in the mitochondria in these cells is
in the hair. The entry of pathogens, including bacteria, particularly important for hibernating animals since it is
viruses, fungi, protozoa, roundworms and flatworms, into instrumental in rapidly raising the core temperature during
the body is greatly hindered by the structure of the skin. arousal from hibernation (section 21.10.3). The hypothala-
Micro-organisms can nevertheless enter through hair mus is the centre initiating heat production for most of the
follicles and sweat glands and through the bites of insect mechanisms described above.
vectors. The larval stages of many parasitic worms, such as
Schistosoma, are capable of boring through the skin. 18.4.3 Loss of heat
Vitamin production. Ultra-violet radiations Heat is lost from endotherms by the four
from the Sun convert a group of steroids, known as sterols, mechanisms described in section 18.2.6, that is conduction,
to vitamin D. This is only a secondary source of the vitamin convection, radiation and evaporation. In all cases, the rate
since most is obtained in the diet (section 10.3.10). of loss depends upon the temperature differences between
Energy storage. Fat is deposited as adipose the body core and the skin, and the skin and the
tissue in the lower regions of the dermis and also in a environment. The rate can be increased or decreased
subcutaneous position. The secondary effect of this is to depending upon the rate of heat production and the
provide thermal insulation as described in section 18.4.3. environmental temperature.
Many mammals build upa store of fat under the skin in There are three factors limiting heat loss, as given below.
preparation for adverse climatic conditions. The rate of blood flow between the body core
and skin. The rate of heat loss from the skin by radiation,
18.4.2 Sources of heat
convection and conduction depends upon the amount of
The major source of heat in endotherms is blood flowing through it. If the blood flow is low the skin
produced from the exothermic biochemical reactions temperature approaches that of the environment, whereas
which occur in living cells. This heat is released by the if the flow is increased the skin temperature then
breakdown of molecules derived from the diet, principally approaches core temperature. The skin of endotherms is
carbohydrates and fats. The amount of heat released in a richly vascularised and blood can flow through it by any of
resting fasting organism is known as the basal metabolic three routes: through capillary networks in the dermis,
rate (BMR) and provides a ‘base-line’ for comparing the through shunt pathways deep in the dermis linking
energy demands during various activities, and in different arterioles and venules, and through subcutaneous, small
organism (see section 11.7.6). The energy content of food connecting veins linking cutaneous arterioles and veins.
required to meet the demands of the basal metabolic rate Arterioles have relatively thick muscular walls which can
for an average-sized male human over a 24 h period is contract or relax altering the diameter of the vessels and the
approximately 8 000 kJ. The exact amount per individual rate of blood flow through them. The degree of contraction
depends upon size, age and sex, being slightly higher in is controlled by sympathetic vasomotor nerves from the
males. vasomotor centre in the brain which receives impulses from
Most of the metabolic energy which appears as heat the thermoregulatory centre in the hypothalamus. The rate
678
skin surface
é very little heat loss 3 large heat loss by
by radiation etc.
arterio-venous anastomosis
(shunt)
connecting vein
small arteryif, \ small vein small arteryip \ small vein
of blood flow through the skin in Man can vary from Fig 18.12 Mechanism of regulation of blood flow through the
less than 1 cm’ min™! 100 g™! in cold conditions to skin. (a) Blood flow through the skin preventing heat loss.
100 cm’ min7' 100 g™! in hot conditions, and this can Constriction of the arteriole reduces blood flow through
capillaries and shunt. Only sufficient blood passes into the
account for an increase in heat loss bya factor of five or six. skin to keep the tissues alive. Most of the blood flowing from
The ‘shunt pathways’ are known as arterio-venous anasto- the body by-passes the skin through the connecting vein and
moses and lie beneath the level of the skin capillary beds. reduces heat loss. (b) Blood flow through the skin increasing
Constriction of these vessels forces blood through the low heat loss. Dilation of the arteriole increases blood flow
through the capillaries and shunt. The capillaries dilate due to
resistance ‘connecting veins’ connecting arteries to veins
the rise in blood pressure within them. Heat is lost from the
and the bulk of the blood by-passes the capillaries and blood by radiation, convection and conduction and blood flow
shunts (fig 18.12). This is a typical response preventing heat is increased to the sweat glands
loss. Dilation of the shunt vessels encourages blood flow
through the capillary beds and shunts and not through the
connecting veins. This increases blood flow through the
skin and therefore heat is lost more rapidly. reduced by low environmental temperatures, high humid-
ity and lack of wind.
The rate of sweat production and evaporation
Many mammals have so much hair on their bodies that
from the skin. Sweat is a watery fluid containing between
sweating is restricted to bare areas, for instance pads of the
0.1 and 0.4% of sodium chloride, sodium lactate and urea.
feet of dogs and cats, and the ears of rats. These mammals
It is hypotonic to blood plasma and is secreted from tissue
increase heat loss by licking their bodies and allowing
fluid by the activity of sweat glands under the control of moisture to evaporate, and by panting and losing heat from
sudomotor neurones. These neurones are part of the the moist nasal and buccal cavities. Man, horse and pig are
sympathetic nervous system and they relay impulses from able to sweat freely over the entire body surface.
the hypothalamus. Sweating begins whenever the body Experiments have now confirmed that sweating only
temperature rises perc its mean value of 36.7 °C. occurs as a result of a rise in core temperature. Experi-
Approximately 900 cm* of sweat are lost per day in a ments on Man and other animals have shown that lowering
temperate climate, but the figure can rise as high as the core temperature, by swallowing ice water or cooling
12 dm’ per day in very hot dry conditions, providing that the carotid blood vessels with an ice pack around the neck,
there is adequate replacement of water and salts. while at the same time exposing the skin to heat, result in a
decrease in the rate of sweating. Opposite effects have
latent heat of evaporation of been recorded by reversing the environmental conditions.
Jalculate the percentage of | | Blood from the carotid vessels flows to the hypothalamus
rom a coalminer who loses and these experiments have indicated its role in thermoreg-
4 dm® per tand has a daily energy ulation. Inserting a thermistor against the ear-drum gives
uptake of 50000 we an acceptable measure of hypothalamic temperature. The
relation between changes in temperature in this region and
the skin, and rate of evaporation of sweat are shown in fig
When sweat evaporates from the skin surface, energy as
18.13. Examine this figure and answer the following
latent heat of evaporation is lost from the body and this
is questions.
reduces body temperature. The rate of evaporation
679
(a) (b) ()) The amount of insulation between the core
and the environment. Insulation for the body is provided
by air trapped outside the skin and by dermal and
subcutaneous fat. Feathers, fur and clothes trap a layer of
air, known as stagnant air, between the skin and the
skin temperature environment, and because it is a poor conductor of heat it
reduces heat loss. The amount of insulation provided by
this means depends upon the thickness of the trapped air.
Temperature/°C Reflex contractions of the erector pili muscles in response
to decreasing temperatures increases the angle between the
feathers or fur and the skin and thus there is more air
hypothalamic temperature
trapped. The response is still present in Man but due to the
minimal amount of body hair it only produces the effect
known as ‘goose-flesh’ or ‘goose-pimples’. Man compen-
os sates for the lack of body hair by taking advantage of the
insulating effects of clothing. The seasonal accumulation of
a
isi
100
Nn o
energy lost by evaporation (sweating) a thick layer of subcutaneous fat is common in mammals
loss/
of
Rate
energy
L 1 = L \ \
70
ae
80
particularly in those species which do not hibernate and
60
manage to withstand cold temperatures. Aquatic mam-
10 20 30 40 50
Time/min
mals, particularly those inhabiting cold waters, such as
Fig 18.13 Graphs showing the relation between skin whale, sea-lion, walrus and seal, have a thick layer of fat
temperature, hypothalamic temperature and rate of
evaporation for a Man in a warm chamber (45 °C). Iced water known as blubber which effectively insulates them against
was swallowed at the points labelled (a), (b) and (c) the cold.
18.4.4 Heat balance and the role of the

hypothalamus
: 18.4 Account for temperature and eva-
poration rate remaining relatively constant during the. The temperature of any body is determined by
the following equation:
rst 20 min. : _
18.5 Describe the relationship between_ heat gained by body = heat lost by body
hypothalamic temperature and rate of sweating. Endothermic animals are able to generate sufficient heat
| 18.6 Suggest why the skin temperature energy and regulate the amount lost so that the two
__ rises shortly after the ingestion of iced water. — expressions above are always in equilibrium and equal a
sky ®
\
30 2€ atmosphere
air temperature
convection
conduction
soil
Fig 18.14 Diagram showing the energy exchanges between a horse, : with a body temperat ure of 38°C,
A i
on a hot sunny day with an air temperature of 30°C. The dotted lines represent hennalvackaien soir sibine
680
constant (fig 18.14). This is known as homeothermy. Any Table 18.1 Functions of the heat loss and heat gain centres
mechanism which has an input and an output and is capable of the hypothalamus. These are situated in the anterior and
of maintaining a constant value must be regulated by a posterior hypothalamus respectively and have antagonistic
control system as described earlier in this chapter. effects
Birds and mammals have a well-developed control
system involving receptors and effectors and an extremely Anterior hypothalamus Posterior hypothalamus
sensitive control centre, the hypothalamus. This organ (heat loss centre) (heat gain centre)
monitors the temperature of the blood, which is in
Activated by increase in the Activated by impulses from
equilibrium with the core temperature, flowing through it. hypothalamic temperature peripheral cold receptors or
If the hypothalamus is to control a constant core tempera- temperature of the
ture, as is the case with endotherms, it is vital that hypothalamus
information regarding changes in the external temperature Increases vasodilation Increases vasoconstriction
is also transmitted to the hypothalamus. Without such Increases heat loss by Decreases heat loss by radiation
information the body would gain or lose a great deal of heat radiation, convection and convection and conduction
before changes in core temperature would activate the conduction
hypothalamus to take corrective measures. This problem is Increases sweating and Inhibits sweating and panting
overcome by having peripheral thermoreceptors, situated panting
in the skin, which detect changes in the environmental Decreases metabolic activity Increases metabolic activity
temperature and initiate impulses to the hypothalamus in through shivering and release
advance of changes in the core temperature. There are two of thyroxine and adrenaline
types of thermoreceptors, hot and cold, which generate Decreases thickness of air Increases thickness of air
impulses in afferent neurones leading from them when layer by flattening hair or layer by action of hair muscles
suitably stimulated. Some pass to the hypothalamus and feathers
others to the sensory areas of the cortex, where the

sensations associated with temperature are registered detected by thermoreceptors in the hypothalamus. In most
according to the intensity of stimulation, the duration and cases the activity of both peripheral and hypothalamic
the numbers of receptors stimulated. There are estimated receptors is instrumental in controlling body temperature.
to be 150 000 cold receptors and 16 000 heat receptors in Investigations into the thermostatic activity of the
Man. This enables the body to make rapid and precise hypothalamus have shown that there are two distinct areas
adjustments to maintain a constant core temperature. In concerned with this type of regulation and the functions of
the context of control systems, the skin receptors act as these areas are summarised in table 18.1. The interrela-
disturbance detectors anticipating changes in body temper- tionships between the cerebral cortex, the hypothalamic
ature. Factors bringing about changes in internal tempera- thermoregulatory centres, core temperature, skin temper-
ture such as metabolic rate or disease will immediately ature and environmental temperatures are shown in fig
affect the core temperature and in these situations be 18.15 and fig 18.16.
Fig 18.15 Summary diagram showing the H

environmental | L
reflex control of body temperature in a mammal & temperature
H =O
involving the environment, hypothalamus and
blood temperature
skin warm skin skin cold

receptors temperature receptors
anterior hypothalamus
|
HEAT LOSS CENTRE inhibition |
|
adrenaline thyroxine
Ss |
|
shivering skin arterioles hair metabolic rate |
sweating _ skin arterioles metabolic rate |
constrict raised increases
dilate decreases |
SS
SS |
eg R
blood |
temperature ee feedback |
Certain diseases produce an increase in core tempera- decreases in size with distance from the Poles. There are
ture as a result of the ‘thermostat’ being set at a higher exceptions to this rule, but the organisms concerned have
temperature. It is believed that certain substances known adaptations favouring survival in these regions. For
as pyrogens, which may be toxins produced by pathogenic example, small mammals in temperate or arctic regions
organisms or substances released by white blood corpuscles have a large appetite enabling them to maintain a high
known as neutrophils, directly affect the hypothalamus and metabolic rate. They have small extremities to reduce heat
increase the set-point. The raised body temperature loss and are forced to hibernate in winter. Large mammals
stimulates the defence responses of the body and aids the living in hot regions, such as the elephant and hippopota-
destruction of pathogens. Antipyretic drugs such as aspirin mus, have the opposite problems. The elephant has
lower the set-point and provide relief from the unpleasant extremely large ears which are well supplied with blood,
symptoms of fever, but probably retard the normal defence and flapping of these ears encourages heat loss by radiation
mechanisms. In cases of extremely high temperature these and convection. The hippopotamus lacks sweat glands and
drugs are valuable in preventing irreversible damage to the adopts a similar behavioural response to temperature as
brain. the crocodile in that it moves between land and water in an
attempt to minimise the effects on its body of changes in
18.7 The onset of fever is often accompa- temperature. The size of the external organs also varies
nied by shivering and a feeling of cold known as chill. according to environmental temperature such that species
Explain these symptoms in terms of mechanism of living in colder climates have smaller extremities than
control of body temperature. related species in warmer climates. This is known as Allen’s
rule and may be seen in closely related species of, for
instance, the fox (fig 18.17).
18.4.5 Adaptations to extreme climates
18.4.6 Adaptations to life at low temperatures
The size of organs and physiological and
behavioural adaptations of organisms vary over their All ectotherms and many endotherms are
geographical range. The total heat production of en- unable to maintain a body temperature which permits
dotherms depends upon the volume of metabolically active normal activity during cold seasons, and they respond by
tissues whilst the rate of heat loss depends upon surface showing some form of dormancy, as described in section
area. The two parameters, surface area and volume, are 21.10. Some of these responses are quite startling; for
inversely proportional to each other. For this reason example, the larva of an insect parasite Bracon, which
animals living in cold regions tend to be large, for example invades the Canadian wheat sawfly, is able to survive
polar bears and whales, whilst animals living in hot climates exposure to temperatures lower than —40.0 °C. This larva
are generally smaller, for instance insectivorous mammals. accumulates glycerol in its haemolymph. The glycerol acts
This phenomenon is known as Bergman’s rule and is as an ‘antifreeze’ and is able to prevent the formation of ice
observed in many species, including the tiger, which crystals by a process known as ‘super-cooling’.
death above 43.0°C
2 a HYPERTHERMIA
|_ body if homeostatic
41 compensates va ol Aconae GA metabolic fate mechanism fails
404 Boel ie vasodilation of arterioles in skin
vs increased blood flow to skin
29 =a a 7. increased radiation, convection and conduction
increase in body sweating and evaporation of water Passe 4
upper ‘normal’ 38b wae flattening of hair (decreased insulation) decrease in body temperature
>| ih
temperature 375°C —— 37L ae fibody ta: area ek eae) aoe
se tads ecreased muscular activity ~~
temperature 36.7 °C . Ne mean body temperature

lower ‘normal’ >| By increase in metabolic rate aa
temperature 35.8°C al \ vasoconstriction of arterioles in skin i sags ae
os NS decreased blood flow to skin ne tease WOT) Ac laptalure
® decrease in body decreased radiation, convection and conduction _— (negative feedback)
2 34- temperature sweating inhibited
aU \ raising of hair (increased insulation)
Be ae i a8 ‘huddling? to decrease surface area
Es \ increased muscular activity
Pe 32s body = =p
5 compensates a
a 31L down to 32.0°C HYPOTHERMIA
if homeostatic
mechanism fails death below 26.0°C
Time
Fig 18.16 Homeostatic control of body temperature
682
Arctic fox mammals and in the blood supply to the testes in mammals
body temperature 37°C
(fig 18.18).
average environmental temperature
The countercurrent exchange principle is used for the
rE transfer of materials other than heat, such as respiratory
gases and ions (sections 11.7 and 19.5.7).
The induction of a state of hypothermia is being used
increasingly in heart surgery since it allows the surgeon to
(| European fox carry out repairs to the heart without the risk of brain
damage to the patient. By reducing the body temperature
body temperature 37°C
to 15 °C, the metabolic demands of the brain cells are so
average environmental temperature
IDG reduced that blood flow to the brain can be stopped,
without any adverse effects, for up to one hour. For
operations requiring a longer time than this a heart-lung
machine is used, in addition to hypothermia, to maintain
blood circulation in the tissues.
African bat-eared fox
body temperature 37°C 18.4.7 Adaptations to life at high tempera-

average environmental temperature tures
25°C
Animals living in conditions where the air
temperatures exceed skin temperature gain heat, and the
Fig 18.17 Variation in ear length shown by three species of
fox (Lycaon) which each occupies a different geographical
only means of reducing body temperature is by evaporation
region. This is an example of Allen’s rule of water from the body surface. For climatic reasons hot
regions may be, in addition, either dry or humid, and this
poses an additional problem. In hot dry regions heat can be
Excessive heat loss to the environment from appendages lost by the free evaporation of water, but animals in these
is prevented in many organisms by the arrangement of regions have the problem of finding adequate supplies of
blood vessels within the appendages. The arteries carrying water to satisfy the demands of evaporative cooling. In hot
blood towards the appendage are surrounded by veins humid regions water is freely available to organisms but the
carrying blood back to the body. The warm arterial blood humidity gradient between organisms and the environment
from the body is cooled by the cold venous blood flowing often prevents evaporation. In the latter case physiological
towards the body. Similarly the cold venous blood from the mechanisms of temperature control are often sup-
appendage is warmed by the warm arterial blood flowing plemented and there may be behavioural activities which
towards the appendage. Because the blood reaching the take advantage of the shade and breezes associated with
appendage is already cooled the amount of heat lost is humid forest and jungle habitats.
considerably reduced. This arrangement is known as a The heat load of an organism is the amount of heat
countercurrent heat exchanger and is found in the flippers gained by metabolic activities and from the environment,
and flukes of seals and whales, in the limbs of birds and and the latter is approximately proportional to the body
environmental
temperature 4 °C 4
ies
cate 2
arterial blood ———————_>
capillary ‘
network ,
venous blood <———____-
34 29 24 19 14 9 4
LeeGee
i } ing the blood supply between the body of an endotherm with a stable temperature of 35°C and an
pee Pe ret at 4°C. fectHews from a warm body to a cool body and the rate of heat loss between the two
blood to the
bodies is proportional to the temperature difference between them. The countercurrent flow shown delivers
is therefore proportional
capillary network at 5 °C and collects blood from it at 4 °C. The amount of heat lost to the environment
than the blood leaving
to the temperature difference of 1°C. Likewise blood returning to the core of the body is only 1 C cooler
core temperature at 35 °C
the core. This mechanism prevents the excessive loss of metabolic energy and helps maintain the
683
surface area. The majority of animals living in deserts, steady state. The liver is derived from an endodermal
therefore, are small, such as the kangeroo rat (Dipo- outpushing of the alimentary canal and many of its
domys), and have fewer problems than large animals, such functions are associated with the preparation, production
as camels. In addition they are able to live in burrows in and control of substances derived from absorbed food
sand and soil where the microclimate poses fewer problems materials. There is a unique dual blood supply to the liver
to life. and because of its rich vascularisation it regulates many
A camel in hot dry conditions, with free access to water, activities associated with blood and the circulatory system
is able to regulate its body temperature between about 36 (fig 18.19). Paradoxically, despite the enormous variety of
and 38 °C. It is able to do this by losing heat through the metabolic activities carried out by the liver its histological
evaporation of water from the body surface. If the camel is structure is relatively uniform and simple.
deprived of water, as say during a journey across the desert
lasting several days, the difference in the body tempera-
tures at morning and evening steadily increase according to hepatic vein inferior
the degree of dehydration. This diurnal fluctuation can be vena cava
from 34°C in the early morning to 41 °C in the late

afternoon. By effectively storing up heat during the day the
camel does not need to lose this heat by the evaporation of
water. It functions in fact, like a storage radiator. In a series gall bladder
of investigations carried out by Schmidt-Nielsen it was bile duct
found that a 500 kg camel tolerating a 7 °C temperature
rise stored approximately 12 000 kJ of heat energy. If this duodenum
amount of heat were lost by evaporative cooling, in order
to maintain a constant body temperature, it would require
the loss of 5 dm® of water. Instead this heat is lost by splenic vein
radiation, conduction and convection during the night. A (spleen, stomach,
pancreas)
second advantage of becoming ‘partially ectothermic’
during the day is that this reduces the temperature anterior
mesenteric
difference between the hot desert air and the camel, and
therefore reduces the rate of heat gain. The fur of the camel
acts as an efficient insulating barrier by reducing heat gain
and water loss. In an experiment, in which a camel was
shorn, the water loss increased by 50% over that of a intestine
control camel. The final significant advantage shown by the
camel is its ability to tolerate dehydration. Most mammals posterior mesenteric vein
(colon and rectum)
cannot tolerate dehydration beyond aloss of body mass of
10-14%, but the camel can survive losses up to 30%, Fig 18.19 Diagram showing the blood supply to and from
because it is able to maintain its plasma volume even when the liver and the relative position of the bile duct
dehydrated. Heat death as a result of dehydration is due to
the inability of the circulatory system to transfer heat from
the body core to the surface quickly enough to prevent
18.5.1 The position and structure of the liver
overheating as a result of a fall in volume of the plasma.
Contrary to popular belief, the camel is unable to store The human liver is a large organ making up
water in advance of conditions of water shortage, and there 3-5% of the body weight and lies immediately beneath the
is doubt whether it can obtain water from the metabolism of diaphragm to which it is attached by the falciform ligament.
fatstoredin the hump. Thecamelis, however, able to drinka It is made up of several lobes and has a variable shape
vast volume of water in a short space of time to rehydrate depending upon the amount of blood present within it. The
the body tissues after a period of severe dehydration. For liver is surrounded by a capsule made up of two layers, the
example, a 325 kg camel is known to have drunk 30 dm’ of outer layer being asmooth, moist peritoneum and the inner
water in less than ten minutes. This is roughly equivalent to a fibrous covering known as Glisson’s capsule which
a Man of average build and weight drinking about 7 dm surrounds all structures entering and leaving the liver. The
(12 pints) of water! fibres of Glisson’s capsule form an ‘internal skeleton’ which
supports the rest of the liver.
18.5 The mammalian liver The cells of the liver are called hepatocytes and show no
structural or functional differentiation. The only other cells
The liver is the largest visceral organ of found in the liver are nerve cells and cells associated with
homeostasis and controls many metabolic activities essen- blood and lymph vessels. Hepatocytes have prominent
tial for the maintenance of the composition of blood at a nuclei and Golgi apparatus, many mitochondria and
684
po es ee ee ee
a portal tract
/ lymphatic branch of hepatic

L vessel portal vein
se
| oe
ao
| <—_—_— ————
| X \ : cs
|
r a bile canaliculus
\ fibrous branch of branch of bile ductule

\ capsule hepatic nerve hepatic artery eo branch of
Seo SS | ee LAVERACINUS hepatic vein
Fig 18.20 A simplified diagram of a liver acinus showing a transverse section of a portal tract and a longitudinal section
through a branch of the hepatic vein
lysosomes, and are rich in glycogen granules and fat culi are lined with microvilli and take up bile from the
droplets. They are tightly packed together, and where their hepatocytes by some form of active transport. The
surface is in contact with blood vessels there are microvilli canaliculi form a branching network which unites to form
which are used for the exchange of materials between the bile ductules. These join together in the portal tract to form
two. bile ducts and these eventually fuse before leaving the liver
The whole internal structure of the liver is complex and as the common hepatic duct.
not fully understood. It is based upon an arrangement of The structure of the liver, as described above, shows it to
hepatocytes and two systems of blood vessels which be a vast network of hepatocytes, blood, blood spaces and
interdigitate with channels called bile canaliculi. The bile canaliculi. This produces a structure with an immense
hepatic portal vein forms many branches inside the liver surface area where each cell is in direct contact with blood
and they carry alongside them branches of the hepatic thus facilitating maximum exchange between cell and
artery, bile duct, nerves, lymphatic vessels and fibrous blood and control of substances in the blood.
tissue of Glisson’s capsule. This arrangement forms a The structure of the liver in the pig is much simpler than
structure known as a portal tract(canal) (fig 18.20). Blood the human liver but is atypical of other mammals. It is
vessels from both the hepatic portal vein, carrying described because its simplicity highlights many of the
absorbed materials from the alimentary canal, and the structural and functional relationships which may not be
hepatic artery, carrying oxygenated blood, join together to readily evident from the account of the typical mammalian
form ‘venules’ which carry blood to the hepatocytes. This situation described above. The pig’s liver is composed of a
arrangement of ‘venules’ and hepatocytes is believed to large number of discrete units, called lobules which are
form the functional unit of the liver and is known as an polygonal in transverse section and have a diameter of
acinus. Smaller blood vessels called sinusoids arise from the approximately 1mm. They form thimble-shaped units
‘venules’ and form a vast network of capillaries before about 2-3 mm long and are enclosed in a connective tissue
uniting to carry blood to branches of the hepatic vein. sheath continuous with Glisson’s capsule. Between the
Adjacent sinusoids are separated from each other by lobules are the interlobular blood vessels consisting of an
‘plates’ of hepatocytes which are often only one cell thick. arteriole of the hepatic artery and a branch of the hepatic
As blood flows along the sinusoids, materials are ex- portal vein anda bile ductule (as found in the portal tract of
changed between the blood and the hepatocytes. The other mammals). An intralobular vein is situated in the
presence of pores, having a diameter up to 10 nm, in the centre of the lobule and receives blood which flows along
endothelial lining of the sinusoids, and the microvilli on the the sinusoids from the portal vein and artery. Blood from
hepatocytes where they touch the sinusoids, facilitates the the intralobular veins passes into the hepatic vein. The bulk
exchange of materials. Bile produced in the hepatocytes of the lobule is composed of strands of liver cells running
does not enter the sinusoids but is secreted into minute bile from the periphery of the lobule and converging on the
canaliculi which replace the sinusoids at various points and intralobular vein. Exchange of materials between hepato-
run between adjacent plates of hepatocytes. These canali- cytes and blood occurs as described previously. The
685
One other type of cell is found in the liver and it forms
part of a more extensive system known as the reticulo-
endothelial system (see section 14.14). The cells are called
Kupffer cells and are found attached to the walls of the
sinusoids by cytoplasmic projections. They are phagocytic
and are involved in the breakdown of old(effete) erythro-
cytes and the ingestion of pathogenic organisms.
18.5.2 Functions of the liver

It has been estimated that the liver carries out
several hundred separate functions involving thousands of
different chemical reactions. These functions are related to
the position of the liver in the circulatory system and the
vast amount of blood which flows through it at any given
time (approximately 20% of the total blood volume). The
liver and the kidney between them are the major organs
responsible for regulating the steady state of blood
metabolites and the composition of the blood tissues. All
food materials absorbed from the alimentary canal pass
Fig 18.21 TS pig liver showing lobules
directly to the liver where they are stored or converted into
some other form as required by the body at that time.
The functions of the liver therefore fall into two main
sinusoids alternate with bile canaliculi which carry bile to categories: the storage of food materials and synthesis of
the intralobular bile ductule. This relatively simple and their derivatives, and the breakdown of substances not
regular pattern is shown in figs 18.21 and 18.22. required by the body prior to their excretion. Finally, as a
The functional unit of pig liver is the lobule and its result of the number of metabolic activities occurring
regular structure arises because the intralobular and within the liver, it may be a source of heat production for
interlobular blood vessels run parallel to each other. This animals living in cold climates.
situation is not generally found in other mammals where
lobulation is absent or less definite. Carbohydrate metabolism
The lymph found in the lymphatic vessels of the portal Hexose sugars enter the liver from the gut by the hepatic
tract is surprisingly rich in protein and is produced from portal vein, which is the only blood vessel in the body
plasma which escapes from the endothelial pores in the having an extremely variable sugar content. This gives a
‘venules’ and sinusoids. clue to the role of the liver in carbohydrate metabolism as
liver cords (sinusoids

intralobular vein and canalicul1)
branch of hepatic portal vein bile ductule
interlobular blood vessels
bile canaliculi
branch of hepatic artery
hepatocytes
sinusoid
Fig 18.22 TS of pig liver showing the regular arrangement of fibrous capsule of
lobules due to the parallel flow of interlobular and intralobular lobule
blood vessels. (Arrows indicate fluid flow in sinusoids and
canaliculi.)
686
amino acids glycerol, fatty acids
gluconeogenesis
(glucocorticoids) glycogenolysis glycogen (liver)
(glucagon, adrenaline,
glucocorticoids) glycogenesis
carbohydrate digestion ;
blood glucose<———_————— glucose-6-phosphate—__—
(insulin) (tissues + liver) erate
Set,
eee
e
a
e
pyruvic acid
ace
aerobic anaerobic
ak
CO, + H,O+ energy lactic acid + energy
Fig 18.23 Summary diagram of carbohydrate metabolism
the organ which maintains the blood glucose level at When the demand for glucose has exhausted the
approximately 90 mg glucose 100 cm blood. The liver glycogen store in the liver, glucose can be synthesised from
prevents blood glucose levels from fluctuating according to non-carbohydrate sources. This is called gluconeogenesis.
feeding patterns thus preventing damage to tissues which Low blood glucose levels (hypoglycaemia) stimulate the
cannot store glucose, such as the brain. All hexose sugars, sympathetic nervous system to release adrenaline which
including galactose and fructose, are converted to glucose helps satisfy immediate demand as described above. Low
by the liver and stored as the insoluble polysaccharide, blood glucose levels also stimulate the hypothalamus to
glycogen. Up to 100 g of glycogen are stored here but more release CRF (section 16.6.2) which in turn releases
is stored in muscle. The conversion of glucose to glycogen adrenocorticotrophic hormone (ACTH) from the anterior
is known as glycogenesis and is stimulated by the presence pituitary gland. This leads to the synthesis and release of
of insulin: increasing amounts of the glucocorticoid hormones,
cortisone and hydrocortisone. These stimulate the release
insulin of amino acids, glycerol and fatty acids, present in the
glucose <> glucose-6-phosphate + glucose-1-phosphate > glycogen tissues, into the blood and increase the rate of synthesis of
(phosphorylation) (condensation)
enzymes in the liver which convert amino acids and glycerol
Glycogen is broken down to glucose to prevent the blood into glucose. (Fatty acids are converted into acetyl
glucose level falling below 60 mg 100 cm blood. This coenzyme A and used directly in the Krebs cycle.)
process is called glycogenolysis and involves the activation Carbohydrate in the body which cannot be utilised or
stored as glycogen is converted into fats and stored. A
of a phosphorylase enzyme by the pancreatic hormone,
glucagon. In times of danger, stress or cold this activity is summary of carbohydrate metabolism involving the liver,
muscles and tissues is shown in fig 18.23.
also stimulated by adrenaline, released by the adrenal
medulla, and noradrenaline released by the endings of the Protein metabolism
sympathetic neurones (section 16.2.3).
The liver plays an important role in protein metabolism
phosphorylase phosphoglucomutase glucose-6-phosphatase which may be considered under the headings of deamina-
glycogen = glucose-1-phosphate = glucose-6-phosphate <> glucose tion, urea formation, transamination and plasma protein
(store) (free)
synthesis.
Muscle lacks the enzyme glucose-6-phosphatase and
Deamination. The body is unable to store
cannot convert glycogen directly to glucose via glucose-6-
absorbed amino acids, and those not immediately required
phosphate as shown above. Instead, glucose-6-phosphate
for protein synthesis or gluconeogenesis are deaminated in
is converted into pyruvic acid which is used to produce ATP
the liver. This involves the enzymic removal of the amino
during aerobic or anaerobic respiration. Lactic acid
group (-NH,) from the amino acid with the simultaneous
produced by anaerobic respiration in skeletal muscle can
oxidation of the remainer of the molecule to form a
be converted later into glucose and hence glycogen in the
carbohydrate which is utilised in respiration. The amino
liver by a biochemical pathway known as the Cori cycle:
group is removed along with a hydrogen atom so that the
lactic acid—>pyruvic acid— glucose glycogen nitrogenous product of deamination is ammonia (NH3).
687
For example, Transamination. This is the synthesis of
amino acids by the enzymic transfer of the amino group
from an amino acid to a carbohydrate in the form of a keto
R R
INE ——E——COOs! ar 0, > 2 C—COOH + 2NH, acid (chapter 5). For example the amino acid, glutamic
acid, could be synthesised by the following reactions:
H ©)
amino acid oxygen keto acid ammonia CH,COOH
or specifically, CH. CH,
CH, CH, NH,—C—COOH_ + O0==C— COOH aan
2NH,—C—COOH + 0O,—————» 2C—COOH + 2NH, H a-oxoglutaric acid
alanine
H O
pyruvic acid ammonia CH,COOH
alanine oxygen
GH
This ammonia may be used for the synthesis of certain CEH
amino acids or nitrogenous bases, such as adenine and NH,—-C—-COOH + O==C——COOH

guanine (section 5.6), or excreted.
H _ pyruvic acid
Urea formation. Ammonia produced by
glutamic acid
deamination is converted in the liver into the soluble
excretory product urea:
The general principle underlying these reactions is the
~~ mutual exchange of characteristic radicals between the
Nee oo CO) Q==30) S5 8b© amino acid and the keto acid:
;
NH,
ammonia carbon urea water

dioxide
NH,—C—COOH + O==C——-COOH————_>
This occurs by a cyclic reaction known as the ornithine cycle
H keto acid B
which is summarised in fig 18.24.
amino acid A
re
deamination
H——C—_CooH +; O=—C—_ COO

H,
H ‘ keto acid A
amino acid B
Ss
Transamination is the means of producing amino acids
which are deficient from the diet, and this is yet another of
V- aa
the liver’s homeostatic mechanisms. The ‘essential’ amino
arginine ornithine acids, described in section 10.3, cannot be synthesised by
transamination in the liver and must be obtained from the
diet.
Plasma protein production. Plasma proteins
are vital components of plasma and the majority of them
are synthesised from amino acids in the liver. Plasma
albumin is the commonest protein and about 4 g 100cm° is
normally present in the blood. It plays an important part in
exerting a colloid osmotic pressure which opposes the
NH, ee hydrostatic pressure developed in blood vessels. The
urea antagonistic effects of these two factors maintains the
balance of fluids inside and outside of blood vessels (section
Fig 18.24 Summary diagram of the
ornithine cycle in mammalian liver (ornithine 14.12.1). Albumins also act as transport molecules within
and citrulline are amino acids but are not the circulatory system, carrying substances such as calcium,
obtained from the diet) tryptophan, bilirubin, bile salts, aspirin and some steroid
688
hormones. Plasma globulins are very large molecules and Once this process is established the liver takes an opposite
blood carries about 34 g dm~*. a- and -globulins transport role and assists in breaking down erythrocytes and
hormones, including thyroxine and insulin, cholesterol, haemoglobin.
lipids, iron and the vitamins B,,, A, D and K. y-globulins
are produced by lymphocytes and the other cells of the Breakdown of haemoglobin
reticulo-endothelial tissues, and are involved in the Erythrocytes have a life-span of about 120 days. By this
immune response (section 14.14.3). The other main plasma stage they are effete and are broken down by the activity of
proteins are the blood-clotting factors, prothrombin and phagocytic macrophage cells of the reticulo-endothelial
fibrinogen, and their functions are described in section system of the liver, spleen and bone marrow. Haemoglobin
14.13.5. is broken down into haem and globin. The globin is reduced
to its constituent amino acids and enters the liver’s amino
Fat metabolism
acid pool to be used according to demand. The iron is
The liver is involved in the processing and transport of fats removed from haem and the remaining pyrrole rings form a
rather than their storage. Liver cells carry out the following green pigment biliverdin. This is converted to bilirubin,
functions: converting excess carbohydrates to fat; remov- which is yellow and a component of bile. The accumulation
ing cholesterol and phospholipids from the blood and of bilirubin in the blood is symptomatic of liver disease and
breaking them down or, when necessary, synthesising them produces a yellowing of the skin, a condition known as
and producing globulins to transport lipids. jaundice.
Vitamin storage Bile production
The liver stores some of the water-soluble Vitamins B and Bile is a viscous, greenish yellow fluid secreted by
C, especially those of the B group such as nicotinic acid, hepatocytes. Between 500-1 000 cm’ of bile are produced
vitamin B,, and folic acid. Vitamin B,, (cyanocobalamin) each day and it is composed of 98% water, 0.8% bile salts,
and folic acid are required by the bone marrow for the 0.2% bile pigments, 0.7% inorganic salts and 0.6%
formation of erythrocytes and deficiency of these vitamins cholesterol. It is involved in digestion, the absorption of
leads to various degrees of anaemia. The main vitamins fats and is a means of excretion of bile pigments.
stored in the liver, however, are the fat-soluble vitamins A, Bile salts are derivatives of the steroid cholesterol which
D, E and K. The liver of certain fish contains high is synthesised in hepatocytes. The commonest bile salts are
concentrations of vitamins A and D, for example cod and sodium glycocholate and sodium taurocholate. They are
halibut. Vitamin K is a vital factor in blood clotting. secreted with cholesterol and phospholipids as large
Mineral storage particles called micelles. The cholesterol and phospholipids
hold the polar bile salt molecules together so that all the
Those elements required in small amounts such as copper,
hydrophobic ends of the molecules are orientated the same
zinc, cobalt and molybdenum (trace elements) are stored way. The hydrophobic ends attach to lipid droplets whilst
in the liver along with iron and potassium. Iron is stored the other ends are attached to water. This decreases the
primarily as a compound called ferritin which is a complex surface tension of the droplets and enables the lipids to
of iron and 6-globulin. Approximately 1 mg g' dry mass separate, forming an emulsion. These smaller droplets
of liver tissue in Man is iron. Most of this iron in the liver is have an increased surface area for attack by pancreatic
temporary and comes from the breakdown of old erythro- lipase which converts the lipids into glycerol and fatty acids
cytes and is stored here for later use in the manufacture of so that they can then be absorbed from the gut. Bile salts
new erythrocytes in the bone marrow. also activate the enzyme lipase, but their action, in all
Storage of blood cases, is purely physical. Too little bile salt in bile increases
the concentration of cholesterol which may precipitate out
The blood vessels leaving the spleen and gut join to form in the gall bladder or bile duct as cholesterol gall stones.
the hepatic portal vein and, together with the blood vessels These can block the bile duct and cause severe discomfort.
of the liver, contain a large volume of blood which acts as a Bile pigments have no function and their presence is
reservoir, though this is not a static store. Sympathetic purely excretory.
neurones and adrenaline from the adrenal medulla can
Cholesterol is produced by the liver and is the precursor
constrict many of these hepatic vessels and make more molecule in the synthesis of other steroid molecules. The
blood available to the general circulation. Likewise, if the major source of cholesterol is the diet, and many dairy
blood volume increases, as for example during a blood products are rich in cholesterol or fatty acids from which
transfusion, the hepatic veins along with other veins can cholesterol can be synthesised. Thyroxine both stimulates
dilate to accommodate the excess volume. cholesterol formation in the liver and increases its rate of
Formation of erythrocytes excretion in the bile. Excessive amounts of cholesterol in
the blood can lead to its deposition in the walls of arteries
The liver of the foetus is responsible for erythrocyte
leading to atherosclerosis (narrowing of the arteries) and
production (erythropoiesis) but this function is gradually the increased risk of the formation of a blood clot which
taken over by cells of the bone marrow (section 14.11.2).
689
may block blood vessels, a condition known as arterial Detoxification
thrombosis. This is often fatal if it occurs in the heart or This term covers a range of homeostatic activities carried
brain. Cholesterol is often cited as a major cause of out by the liver so as to maintain the composition of blood
cardiovascular disease but, as yet, much of the evidence is at a steady state. Bacteria and other pathogens are
contradictory. removed from the blood in the sinusoids by Kupffer cells
Bile is stored and concentrated in the gall bladder by but the toxins they produce are dealt with by biochemical
absorption of sodium ions (and water) into surrounding reactions in the hepatocytes. Toxins are rendered harmless
blood capillaries. The stimulus for the release of bile into by one or more of the following reactions: oxidation,
the duodenum is the presence of the hormone cholecysto- reduction, methylation (the addition of a -CH; group) or
kinin-pancreozymin (CCK-PZ) as described in section combination with another organic or inorganic molecule.
1033: Following detoxification these substances, now harmless,
are excreted by the kidney. The major toxic substance in
Hormone production and breakdown the blood though is ammonia, whose fate is described
Whilst the liver is not generally considered as an endocrine above in this chapter. The detoxification process also
gland, it synthesises and releases growth-promoting factors includes harmful substances taken in to the body such as
called somatomedins under the influence of the hormone alcohol and nicotine. Alcohol taken in excess (in gradually
somatotrophin, released from the pituitary gland. This increasing dosage) can result in liver breakdown, such as
control of growth is described in more detail in section cirrhosis of the liver in alcoholics.
21.8.1. The liver destroys almost all hormones to various Some of the metabolic activities of the liver may be
extents. Testosterone and aldosterone are rapidly des- potentially harmful and evidence is growing that certain
troyed, whereas insulin, glucagon and gut hormones, food additives may be converted into poisonous or
female sex hormones, adrenal hormones, ADH and carcinogenic substances by liver activity. Even the pain
thyroxine are destroyed less rapidly. In this way the liver killer paracetamol, if taken in excess, is changed into a
has a homeostatic effect on the activities of these substance which affects enzyme systems and can cause
hormones. liver, and other tissue, damage.
INTESTINE LIVER BLOOD
monosaccharides glycogen & glucose

a ae ae
stored ee ee |
lipids =——fats pe fatty acidsand glycerol
amino acids — e aminoacids

a transamination ps! Lamninercds
ee ae
keto acids NH,
carbohydrates urea > urea

fatty acids
protein synthesis ’
ar & plasma proteins
vitamins > BandC
fat-soluble A, D, E and K stored
mineral salts po Gus, Za |Cou
2+ Be2
—a ‘trace elements’and Fe?* and Kt stored SS Mer Hee
blood in hepatic i & blood volume
portal vein [stored}
formation of erythrocytes (foetus) ———__———_|_ erythrocytes
haemoglobin breakdown——-——_|_ Fe?+
biliverdin
EA bilirubin
terol
eee & cholesterol
converted to bile salts
bile rt broken down
detoxification 4 hormones
toxins
(harmless compounds)
Fig 18.25 Summary diagram of the functions of the liver
690
Heat production
Evidence is accumulating to show that the widespread
belief that the metabolic activity of the liver results in it
being a major source of heat production in the body of
mammals may be false. Many of the liver’s metabolic
activities are endothermic and therefore require heat
energy rather than release it. Under conditions of extreme
cold the hypothalamus will increase the ectothermic
activity of the liver by its influence on the release of
adrenaline by the sympathetic nervous system and the
release of thyroxine. In ‘normal’ temperatures, however,
the liver has been shown to be ‘thermally neutral’ but is
usually 1-2 °C hotter than the rest of the body core.
The liver provides yet another example of the intimate
relationship which exists in biological systems between
structure and function. What is remarkable about the liver
is the diversity of function achieved by a highly complex
morphological structure which has such a simple, undiffer-
entiated histological structure.
691
‘St eee -
ee
<a
.
Chapter Nineteen
Excretion and osmoregulation
Excretion and osmoregulation are two important homeo- (2) The removal of metabolic waste substances which, if
static processes occurring in living organisms. Each process they accumulated, would affect the metabolic activity of
enables organisms to maintain, to varying degrees, the the organism. Many of these substances are toxic acting as
internal environment at a steady state despite changes in inhibitors of enzymes involved in metabolic pathways.
the external environment. (3) The regulation of the ionic content of body fluids. Salts
Excretion is the elimination from the body of waste behave as electrolytes and undergo dissociation in the
metabolic substances which if permitted to accumulate aqueous media of living organisms. For example,
would prevent the maintenance of a steady state. Many sodium chloride, taken in as part of the diet, exists in
substances which are not metabolic, that is those which body fluids as sodium ions (Na) and chloride ions
have not been synthesised by the organism, are eliminated. (Cl-). If the balance of these and other ions is not
To distinguish excretion from these latter functions it is carefully regulated within narrow limits, the efficiency
necessary to define them. Secretion is the passive or active of many physiological and biochemical activities is
removal of molecules from cells into the extracellular impaired; for instance a reduction in Na* concentration
environment, such as the bloodstream, gastro-intestinal leads to a decrease in nervous coordination. Other
tract or external environment. These molecules have been important ions whose concentrations must be carefully
synthesised in vivo (in life), for example hormones and regulated are K*, Mg’*, Ca’*, Fe’*, H*, Cl’, I", PO,”
enzymes, and are therefore considered to have been and HCO,, as they are vital for many metabolic
metabolised, but are not regarded as waste substances. activities including enzyme activity, protein synthesis,
Secretion may form part of the process of excretion as is production of hormones and respiratory pigments,
described later in the chapter. Egestion is the elimination of membrane permeability, electrical activity and muscle
waste substances, mainly undigested food, which have contraction. Their effects on water content, osmotic
never been involved in the metabolic activities of cells. pressure and pH of body fluids are described below.
(4) The regulation of water content of body fluids. The
amount of water within the body fluids and its
19.1 The significance of excretion and regulation is one of the major physiological problems
osmoregulation faced by organisms in the colonisation of many of the
available ecological niches on this planet. The solutions
Excretion and osmoregulation have a number to this problem have produced some of the most
of functions which may be listed and summarised as important structural and functional adaptations shown
follows. by organisms. The mechanisms of obtaining water,
(1) The removal of metabolic waste substances which are preventing water loss and eliminating water are
often by-products of major metabolic pathways. This is diverse, but they are of great importance in maintaining
necessary in order to prevent unbalancing the chemical the osmotic pressure and volume of body fluids at a
equilibria of reactions. Many metabolic reactions are steady state, as described later in this chapter. Before
reversible and the direction of the reaction is deter- describing these it is important to emphasise that the
mined solely by the relative concentrations of reactants osmotic pressure of body fluids depends upon the
and products in accordance with the law of mass action. relative amounts of solute and solvent, that is water,
For example, in the enzyme-catalysed reaction: present. The mechanisms of regulation of solutes and
water are known as osmoregulation.
ee eae (5) The regulation of hydrogen ion concentration (pH) of
(reactants) (products) body fluids. The nature of pH and methods of its
measurement are described in appendix A1.1.5 but the
the continued production of C, a vital requirement of mechanisms of excreting those ions which have a major
metabolism, is ensured by the removal of D, a waste influence on pH, such as H* and HCO,, are
product. This will ensure that the equilibrium of
the considered in this chapter. For example, the pH of
reaction favours the reaction to proceed from reacta nts urine may vary between 4.5 and 8 in order to maintain
to products. the pH of the body fluids at a fairly constant level.
693
Table 19.1 Summary of the products, sources, functions demand for oxygen in respiration and this escapes from
and fates of the major excretory products plants into the environment by diffusion.
eS Saw>_<—wmaom> Many organic waste products of plants are stored within
Product Source Function! Fate dead permanent tissues such as the ‘heart-wood’ or within
E
Preiig ieee Siti e Se leaves or bark which are periodically removed. The bulk of
Oxygen Photosynthesis in Reactant in aerobic most perennial plants is composed of dead tissues into
autotrophic organisms respiration
which excretory materials are passed. In this state they
Carbon Aerobic respiration in Reactant in — have no adverse effects upon the activities of the living
dioxide all organisms photosynthesis
Decreases pH of tissues. Similarly, many mineral salts, taken up as ions,
Breakdown of urea
body fluids may accumulate due to the differential use of cations and
Aerobic respiration in Solvent in all anions. Organic acids, which might prove harmful to
Water
all organisms metabolic activities plants, often combine with excess cations and precipitate
Condensation reactions Reactant in out as insoluble crystals which can be safely stored in plant
photosynthesis, etc. cells. For example, calcium ions and sulphate ions are
Ions (salts) Nutrient metabolism Maintenance of taken up together, but sulphate is used up immediately in
osmotic pressures amino acid synthesis leaving an excess of calcium ions.
Recycled through
ecosystem These combine freely with oxalic and pectic acids to form
harmless insoluble products such as calcium oxalate and
Bile salts Lipid metabolism in Emulsification of
liver fats calcium pectate. Other ions, such as iron and manganese,
None
and organic acids, such as tannic and nicotinic acids, pass
Bile pigments Breakdown of haem
in liver into leaves where they accumulate and contribute to the
Tannins and Nitrogen and Bitter substances characteristic autumn tints of leaves prior to their loss
other organic carbohydrate metabolism deter ingestion during leaf abscission. Substances are not only eliminated
acids in certain plant species by animals through leaf loss but also through petals, fruits and seeds,
Nitrogenous _ Protein and nucleic Decompose and although this excretory function is not the primary function
substances acid metabolism recycled through of their dispersal. Aquatic plants lose most of their
ecosystem
metabolic wastes by diffusion directly into the water
surrounding them.
19.1.1 Excretory products 19.1.3 Excretion in animals

The major excretory products of animals and Any permeable surface which directly con-
plants and their sources are summarised in table 19.1. Not nects a region containing excretory products to the external
all excretory products are waste in the sense of serving no environment is a potential area of excretion. These include
further useful purpose to the body. Indeed, many serve the cell membrane of unicellular organisms, the epidermis
useful purposes prior to their elimination and afterwards, of lower invertebrates, trachea of arthropods, gills and skin
for reasons described in table 19.1. of fish and amphibia and the lungs and skin of higher
vertebrates. The cells of organisms having a relatively
19.1.2 Excretion in plants simple structure are usually in direct contact with the
environment and their excretory products are immediately
Plants do not have as many problems regard-
removed by diffusion. As organisms increase in complex-
ing excretion as do animals. This is because of fundamental
ity, excretory organs develop to convey excretory products
differences in the physiology and mode of life between
from the body directly or indirectly to the external
animals and plants. Plants are primary producers and they
environment through ducts and pores. In the case of the
synthesise all their organic requirements according to the
higher vertebrates specialised excretory structures are
demand for them. For example, plants manufacture only the
present to augment the activity of the vascular system
amount of protein necessary to satisfy immediate demand.
which removes metabolic wastes from cells and transfers
There is never an excess of protein and therefore very little
them to the excretory organs. The most important
excretion of nitrogenous waste substances, produced by the
excretory organs in these organisms are the skin, lungs,
catabolism (breakdown) of proteins, occurs. If proteins are
liver and kidney. The roles of the first three only will be
broken down into amino acids, the latter can be recycled into
described at this stage.
new proteins. Three of the waste substances produced by
certain metabolic activities in plants, that is oxygen, carbon Skin. Water, urea and salts are actively
dioxide and water, are raw materials (reactants) for other secreted from capillaries in the skin by the tubules of the
reactions, and excesses of carbon dioxide and water are sweat glands. Sweat is secreted onto the skin where the
used up in this way. The only major gaseous excretory water evaporates using latent heat of evaporation. In this
product of plants is oxygen. During light periods the rate way heat is lost from the body and this helps to regulate the
of production of oxygen is far greater than the plant’s body temperature.
694
Lungs. Carbon dioxide and water vapour Fig 19.1 Molecular structure of the O
diffuse from the moist alveolar surfaces of the lungs, which three main nitrogenous excretory |
in mammals are the sole excretory organs for carbon products a
H
dioxide. Some of the water released at the lung surface is Van NH,iN HN Eg= BS
metabolic, that is, produced as a waste product of aa ere | | —=0
respiration and therefore excretory, but its exact origin is
not really important in view of the large volume of water
Pe Sth H
contained within the body. ammonia urea uric acid
Liver. Considering the many homeostatic excretory product is determined by the metabolic capabil-
roles of the liver described in section 18.5.2 it is not
ity of the organism (that is, which enzymes are present), the
surprising that these include excretion. Bile pigments are
availability of water to the organism (that is, its habitat),
excretory products from the breakdown of the haemoglo- and the extent to which water loss is controlled by the
bin of effete (old) red blood cells. They pass to the organism.
duodenum as a constituent of bile for removal from the Animals may be classified according to the nature of
body along with the faeces, to which they impart a their major nitrogenous excretory product and, as shown in
characteristic colour. The most important excretory role of table 19.2, there is a degree of correlation between
the liver is the formation of nitrogenous waste products by excretory product, embryonic environment and adult
the deamination of excess amino acids (section 18.5.2). habitat. The classification of organisms and the correlation
with habitats for the majority of organisms may be
19.2 Nitrogenous excretory products summarised thus:
ammonia ammoniotelic aquatic
Nitrogenous waste products are produced by urea ureotelic aquatic/terrestrial
the catabolism of proteins and nucleic acids. The immedi- uric acid __uricotelic terrestrial
ate nitrogenous waste product of the deamination of
19.2.1 Ammonia
proteins is ammonia, and the basis of this reaction is
described in section 18.5.2. Ammonia may be excreted Ammonia is an extremely soluble molecule
immediately or converted into the major nitrogenous with a low relative molecular mass (17) and diffuses rapidly
compounds, urea and uric acid, which differ in their through water. It is toxic to animals and cannot be stored
solubility and toxicity (fig 19.1). The exact nature of the in the body. Mammals are very sensitive to ammonia
and cannot tolerate concentrations in excess of
Table 19.2. Summary of the relationships between excretory 0.02 mg 100 cm™ blood. The high solubility of ammonia
products and stages in the life cycle of various animal groups aids its rapid excretion as ammonium ions (NH,’) in most
aquatic organisms from protozoa to amphibia before it
Animal Excretory Embryonic Adult habitat reaches concentrations which are toxic to the organisms.
product environment
19.2.2 Urea
protozoan ammonia aquatic aquatic
Urea is formed in the liver of vertebrates by
poriferan ammonia aquatic aquatic
the interaction of ammonia, produced by deamination, and
coelenterate ammonia aquauc aquatic
carbon dioxide, produced by respiration, in a cyclical
platyhelminth ammonia aquatic aquatic
reaction known as the ornithine cycle. It is less soluble and
ammonia aquatic aquatic
aquatic crustacean less toxic than ammonia and is the main nitrogenous
terrestrial insect uric acid cleidoic egg terrestrial excretory substance in elasmobranchs, certain teleost fish,
gastropod mollusc uric acid cleidoic egg terrestrial adult amphibia and mammals. The normal level of urea in
echinoderm ammonia aquatic aquatic mammalian blood is 2.5-6.0 mmol dm~ blood. The
elasmobranch urea aquatic aquatic metamorphosis of the tadpole stage to the adult form in the
freshwater teleost ammonia aquatic aquatic frog is marked by the change from ammonia excretion to
urea, aquatic aquatic urea excretion.
marine teleost
trimethyl- 19.2.3 Uric acid
amine
oxide Uric acid and its salts are ideal excretory
larval amphibian ammonia aquatic aquatic products for terrestrial organisms and a pre-requisite for
adult amphibian urea — semi-terrestrial organisms producing a cleidoic egg (shelled egg) (section
reptile uric acid cleidoic egg terrestrial 19.4.8) since they combine a high nitrogen content with low
bird uric acid cleidoic egg terrestrial toxicity and low solubility. They can be stored in cells,
urea aquatic terrestrial tissues and organs without producing any toxic or adverse
mammal
osmoregulatory effects, and they require a minimal
Se
695
amount of water for their excretion. As the concentration store for the regeneration of muscle ADP to ATP (section
of uric acid in the tissue rises it settles out as a solid 17.4.8).
precipitate. The biochemical details of uric acid formation A final group of nitrogenous waste compounds results
and excretion are described in section 19.4.5. Uric acid and from detoxification processes occurring in the liver. The
ammonium urate are the forms in which nitrogenous commonest product is hippuric acid (hippos, horse) which
excretion occurs in insects, lizards, snakes and birds. Man, was discovered in horse urine and is formed by the
apes and, because of a kidney defect, the Dalmatian dog conjugation of benzoic acid (from plant foods) and the
excrete small quantities of uric acid but this is produced amino acid glycine. Many other phenolic compounds such
from the breakdown of nucleic acids and not from the as benzoic acid are rendered harmless by similar detoxifica-
breakdown of proteins. The normal level of uric acid in the tion reactions.
blood of Man is 3 mg 100 cm~™* blood and approximately
1 g of uric acid is excreted in urine per day.
19.3 Nitrogenous excretion and
19.2.4 Other nitrogenous excretory com- osmoregulation
pounds
The major source of waste nitrogenous sub-
In addition to the excretory products previous-
stances is the deamination of excess amino acids. This
ly described, excess dietary protein is converted in some
produces ammonia which is extremely toxic and must be
marine fishes into another excretory substance, trimethyla-
eliminated. Being soluble, ammonia can be eliminated
mine oxide.
from the body rapidly and safely if diluted in a sufficient
volume of water. This presents no real problems to
organisms which have ready access to water but this applies
only to those organisms living in the freshwater environ-
trimethylamine oxide
ment. Marine and terrestrial organisms have an acute
This is produced by the addition of methyl groups to problem of gaining or conserving water respectively,
ammonia, formed by deamination, and the subsequent therefore very little is available for the elimination of
oxidation of intermediate molecules. Trimethylamine nitrogenous waste. Table 19.2 reveals that organisms living
oxide gives fish its characteristic smell. in these environments have developed alternative means of
The only other source of nitrogenous waste substances is nitrogen excretion. These involve the development of
nucleic acid. Foods such as yeast, liver and kidney are rich many anatomical, biochemical, physiological and behav-
in small cells and have abundant nuclei. The breakdown of ioural mechanisms involving the elimination of nitro-
these foodstuffs releases significant amounts of the nucleic genous waste whilst maintaining the composition of the
acid bases, purines and pyrimidines. Spiders and some body fluids at a steady state. Since these may involve
mammals excrete the purines adenine and guanine direct- excretion and osmoregulation the two processes will be
ly, but in other organisms these are broken down into uric considered together.
acid as described later. Osmoregulation is a homeostatic process by which
Adenine and guanine have a similar structure to uric acid animals and plants maintain the concentration of their
whereas the pyrimidine bases cytosine, thymine and uracil body fluids at a steady state. Body fluids are found within
have a structure which enables them to be broken down cells (intracellular) and outside cells (extracellular). For
into a molecule of ammonia and a molecule of amino acid. example, the fluid within plant cell vacuoles (cell sap) is
The amino acid then undergoes deamination and the intracellular whereas the fluid surrounding the cells of the
nitrogenous waste is excreted in the form which is typical of cortex of a plant stem or root is extracellular. In
that organism. multicellular animals intracellular fluid is dispersed fairly
Insects, terrestrial reptiles, birds, Man, apes and the evenly throughout the cell whereas extracellular fluid exists
Dalmatian dog excrete purines as uric acid. The majority of as plasma and interstitial fluid. The latter is further
mammals, however, possess the enzyme uricase in the subdivided in vertebrates into tissue fluid and lymph. It is
liver, which converts uric acid directly to the excretory vital that the composition of these fluids should remain at a
product allantoin. Dipteran insects also excrete allantoin. steady state in order for the metabolic activities of the cells
Some teleost fish produce allantoin which oxidises to the to work efficiently. The nature of the intra- and extracellu-
excretory product allantoic acid. lar fluids and their regulation in plants is described in
Creatine and its derivative creatinine are other nitro- section 19.3.2.
genous waste products. Creatine is formed in the liver of Osmoregulation is not a term used simply to describe the
vertebrates from the amino acids, arginine, methionine control of water balance within an organism. It refers to the
and glycine. Approximately 2% of the total amount of control of the composition of body fluids, which in all cases
creatine in the body is lost each day as creatinine. Some are solutions of varying complexity. Details of the physical
creatine is phosphorylated in the muscles to form creati- and chemical properties of solutions are described in
nine phosphate (phosphagen) where it acts as an energy appendix A1.4.
696
In living systems, even if the osmotic pressures of two 19.3.2 Osmoregulation in plants
solutions are the same (that is they are isotonic), solutes
will move if their relative concentrations are different. The Plant tissue contains a higher proportion of
movement of water molecules between two solutions, by water than animal tissue, and the effective and efficient
Osmosis, Occurs in response to the relative osmotic functioning of the plant cell and the whole plant depends
pressures of the two solutions. Solute molecules move upon maintaining the water content at asteady state. Plants
across differentially permeable membranes in a direction do not have the same problems of osmoregulation as
determined by their relative concentrations on either side animals and they can be considered simply in relation to
of the membrane and their size in relation to the pores of their environment. On this basis plants are classified as
the membrane. This movement may be passive and outlined below.
molecules move down a concentration gradient from a Hydrophytes
region of their high concentration by diffusion, or they may Freshwater aquatic plants such as Canadian pondweed
move against the concentration gradient asa result of active (Elodea canadensis), water milfoil (Myriophyllum) and the
transport by carrier mechanisms located in the membrane. water lily (Nymphaea) are classed as hydrophytes and have
Membranes, including the plasmalemma (cell or plasma fewer osmoregulatory problems than any of the other plant
membrane), cytoplasmic tissue layers, body surfaces and types. Plant cells in fresh water are surrounded by a
gills, can all act as differentially (or selectively) permeable hypotonic environment and water enters the vacuolar sap
membranes through which water and solutes can pass. by osmosis. The water passes through the freely permeable
The osmotic concentration of solutions is described in cell wall and the differentially permeable plasma and
this chapter in terms of osmotic potential and expressed tonoplast membranes. As the volume of the vacuole
either in milli-osmoles per litre (mOsm dm~) or milli- increases due to water uptake, it generates a turgor
osmoles per kilogram of water (mOsm kg’). pressure. The cell becomes turgid and a point is reached
For biological purposes involving osmoregulation the when the water potential has increased to equal that of the
concentration of a solution may also be described in terms surrounding water (about zero), and no further water
of the freezing point depression of the solution. Pure enters (see section 14.1.5). This is termed mechanical
water freezes at 0 °C, but as solutes are added the freezing osmoregulation. Nitella clavata is a freshwater alga with an
point falls below 0 °C and the new freezing point indicates extremely hypertonic sap, whose osmoregulatory prob-
the concentration of solutes in the solution. For example, lems only concern the maintenance of the ionic contents of
sea water has an osmotic potential of —1 000 mOsm dm~ the sap. This is carried out by active uptake from the
and a freezing point depression (A) of —1.7 °C. surrounding water.
Halophytes
19.3.1 Osmoregulatory mechanisms The only plants able to live immersed in sea water are algae
The osmotic relationships between two solu- and they form the major source of vegetation on the
tions separated by a differentially permeable membrane seashore. The distribution of algal species down the shore
can be described in terms of tonicity or osmoticity. There is determined by many factors, including tolerance to wave
are important differences between the two terms regarding
the nature of solutions and the circumstances in which they 100 Fig 19.2 Graph showing the comparative
are used; however, for simplicity, in this text the relation- rates of water loss upon exposure to air for
ships between solutions are described in terms of tonicity, 90 four species of algae found on the sea-shore.
that is solutions are either hypotonic fo or hypertonic (From J. Zaneveld (1937) J. Ecol., 25, 431-68
to another solution or isotonic with another solution.
80
The body fluids of freshwater organisms are usually
hypertonic to their aquatic environment whilst those of
many marine organisms, particularly vertebrates, are Seo
hypotonic to sea water. Many marine invertebrates, on
ie \
a VV
5 oot
Ga \
the other hand, are isotonic with the marine environment.
ob \\\, ceteeoe Fucus spiralis
If the concentration of solutes in an aquatic environment 5 Pad
animals N pata ee Ascophyllum nodosum
increases, or the volume of water decreases,
respond in either of two ways. An osmoconformer would a x — — — Fucus vesiculosus
fluids to equal those of \r\\
alter the concentrations of its body 40 - Fucus serratus
the new surroundings, whilst an osmoregulator would

maintain its osmotic concentration despite changes in the 30 -
external environment. In homeostatic terms, osmoconfor-
mers are described increasingly as poikilosmotic and 20 | l It i a eee | | | ! | | |
osmoregulators as homeosmotic, in line with the prefixes 6 12 18 24 30 36 42 48 54

Time/h
60 66 72 78 84 90 96
used in temperature regulation.

action, desiccation when exposed by tides and the nature of grow, flower and complete seed formation in four weeks,
their photosynthetic pigments. In all cases these species can for example the Californian poppy (Escholtzia). The
tolerate increases in salinity and their main osmoregulatory seeds produced lie dormant until the next rainy spell.
problem is the prevention of water loss by evaporation. Drought endurers, on the other hand, show many
Channel wrack (Pelvetia canaliculata) occupies the highest structural (xeromorphic) and physiological adaptations
algal zone on sheltered rocky shores surrounding the enabling them to survive in extremely dry conditions. Most
British Isles, and its habitat tolerance is aided by thick cell of the xerophytic species of the British Isles are associated
walls, a thick covering of mucilage and a stipe shaped as a with the strand line and sand dunes, such as saltwort
channel. Fig 19.2 shows the rate of water loss and degree of (Salsola) and sea sandwort (Honkenya) found growing in
tolerance of four common British species of seaweed which small mounds of sand on the shore. Sand couch grass
are zoned according to their ability to retain water when (Agropyron) and marram grass (Ammophila) are domi-
exposed to air. nant species of embryo dunes and have extensive rhizome
Halophytes, however, are defined as plants inhabiting systems with adventitious roots for obtaining water from
areas of high salinity such as those encountered in estuaries well below sand level. Agropyron is able to tolerate salt
and salt marshes where salinity is constantly changing and concentrations in the sand up to 20 times that of sea water.
may exceed that of sea water. Whilst the shoot system is not Both Ammophila and Agropyron are important pioneer
regularly exposed to high salinities, the root system must plants in the development of sand-dune systems.
tolerate the increased salinities of the sand and mud which Xerophytic plant species of desert regions show several
accompany hot windy periods when the tide is out. It was adaptations to reducing water loss and obtaining and
thought that these plants must tolerate periods of ‘physi- storing water. Some of these are summarised in table 19.3.
ological drought’ when water is unavailable to the tissues
Table 19.3. Summary of methods of conserving water
due to the hypertonic nature of the environment of the
shown by various plant species (see chapters 14 and 16)
roots. However, this does not seem to be the case and high
transpiration rates and high osmotic pressures in root cells
Mechanism of
enable water to be taken up. Cord grass (Spartina) is a water
common halophyte found low down on estuaries and salt conservation Adaptation Example
marshes; it has an extensive system of rhizomes for
propagation, bearing adventitious roots for anchorage and waxy cuticle \ prickly pear
purposes of water and ion uptake. Other halophytes of few stomata (Opuntia),
aoe sunken stomata pine (Pinus),
estuaries and salt marshes include smaller plants which reduction in
franshiranan
:
stomata open at night
:
ice plant
store water when it is freely available. Common examples ae p and closed by day (Mesembryanthemum)
of these species are glasswort (Salicornia), seablite surface covered with
(Suaeda maritima) and sea purslane (Halimione). Some fine hairs
species, such as sea milkwort (Glaux) and Spartina are able curled leaves marram grass
(Ammophila)
to regulate their salt content by excreting salt from glands
at the margins of the leaf. fleshy succulent leaves Bryophyllum
storage of fleshy succulent stems Pak aa
Mesophytes
water
The majority of angiosperm plant species are mesophytes, fleshy underground Raphionacme
and they occupy habitats with adequate water supplies. tuber
They are faced with the problem of water loss by
evaporation from all aerial parts. Features which help to deep root system acacia
water uptake below water table oleander
reduce water loss are both structural (xeromorphic) and
shallow root system cactus
physiological, and include the presence of a cuticle, absorbing surface moisture
protected stomata whose diameters can be regulated, a
variable leaf shape, abscission and an ecological distribu-
tion based upon tolerance to dehydration. Further details 19.3.3 Processes associated with excretion
and examples of these mechanisms are found in section and osmoregulation
14.3.
Ultrafiltration is the process by which solvent
Xerophytes and solute molecules separate from a solution according to
Plants adapted to life in dry regions and able to survive their differential abilities to pass through the pores in a
long periods of drought are called xerophytes. These form filter. The filter in most animals is the layer separating the
the typical flora of desert and semi-desert regions and are circulatory system and the osmoregulatory or excretory
common along the strand line of the seashore and in sand organ. The force required to produce filtration is a
dunes. Some plants respond to extreme conditions by hydrostatic pressure and comes from the blood pressure.
surviving in the seed or spore stage. These are known as The filtered solution is known as filtrate. Most of the
drought evaders and can germinate following rainfall and contents of blood are removed by ultrafiltration, the
698
exceptions being really large molecules, such as proteins, fluid by specialised osmoregulatory/excretory organs, such
and cells, such as red blood cells. as Malpighian tubules and kidneys. A balance must be
Selective reabsorption involves the selective uptake of achieved between the amount of water and ions lost and
solute molecules and water in amounts which are useful to gained. The problems of water balance are described in
the body. Those substances which are metabolic wastes are detail in section 19.4.
not reabsorbed, nor are solute and water molecules if their
reabsorption would result in their exceeding the normal Adaptations to severe drought
steady-state composition of the body fluids. Reabsorption The kangaroo rat (Dipodomys) is quite remarkable among
occurs initially by passive diffusion until the diffusion mammals in being able to tolerate drought conditions in the
gradient levels out, after which further reabsorption occurs deserts of North America. It flourishes in these conditions
by active transport. As solutes are reabsorbed, the filtrate by possessing a unique combination of structural, physio-
becomes progressively more dilute or hypotonic to the logical and behavioural adaptations. Water loss by eva-
body fluids so that water molecules follow the movement of poration from the lungs is reduced by exhaling air at a
ions by osmosis to produce a filtrate which is isotonic to the temperature below core temperature. As air is inhaled it
body fluids. A hypotonic filtrate is produced by the further gains heat from the nasal passages which assume a lower
uptake of ions from the filtrate in a region of the temperature. During exhalation water vapour in the warm
osmoregulatory/excretory organ which is impermeable to air condenses on the nasal passages and is conserved. The
water. kangaroo rat feeds on dry seeds and other dry plant
Secretion occurs by active transport and removes solutes material and does not drink. Water produced by the
from the body fluids to the filtrate or directly to the chemical reactions of respiration, and that present in
environment. It therefore operates in the opposite direc- minute amounts in its food, are its only sources of water.
tion to reabsorption. This mechanism further increases the The classic investigations by Knut Schmidt-Nielsen
osmotic pressure of the filtrate and decreases the osmotic (summarised in table 19.4) revealed the overall water
pressure of the body fluids. metabolism balance for a kangaroo rat weighing 35 g
The net effect of these three mechanisms of ultrafiltra- metabolising 100 g of barley in experimental surroundings
tion, selective reabsorption and secretion is homeostatic, in at 25 °C and a relative humidity of 20%. Throughout this
that it maintains the composition of the body fluids at a period the only source of water was the barley grain.
steady state. Table 19.4 Water metabolism for a kangaroo rat under
There is a range of osmoregulatory or excretory organs experimental conditions. The absorbed water was the
and organelles which show variety in morphological water present in the food
structure and anatomical location, yet they all rely for their
function on one or more of the mechanisms described Water gains cm
3
Water losses cm
3
above. Some of the structures are relatively non-

specialised and share common characteristics within a oxidation water 54.0 urine 13.5
range of organisms, for example contractile vacuoles, absorbed water 6.0 faeces 2.6
nephridia and kidneys, whereas others are relatively evaporation 43.9
specialised, such as gills, rectal glands and salt glands. Total water gain 60.0 Total water loss 60.0
19.3.4 The effect of environment on Finally the kangaroo rat avoids excessive evaporative
excretion and osmoregulation water losses in the wild by spending much of its time in the
The nature of the environment produces relatively humid atmosphere of its underground burrow.
certain osmoregulatory problems in different groups of The other spectacular example of water conservation is
organisms. Many aquatic organisms living in a hypertonic the camel, whose physiological adaptations are described
environment Jose water by osmosis and gain solutes by in section 18.4.7.
diffusion. The water loss is replaced in various ways,
including drinking and eating, but this increases the solute 19.4 A phylogenetic review of organs
concentrations of the body fluids and necessitates the and processes of nitrogenous
removal of excess solute molecules by active transport. excretion and osmoregulation
Organisms living in a hypotonic environment gainwater by
osmosis and lose solutes by diffusion. In order to minimise Throughout this review the following points
these exchanges the organisms often have an impermeable should be considered:
outer covering and take up ions from the environment by (1) the environment influences the nature of the excretory
active transport. product and the process of osmoregulation,
All terrestrial organisms face the problem of water and (2) some groups of organisms have species adapted to life
in more than one environment,
solute loss from their body fluids to the environment. The
intracellular body fluid of these organisms is maintained at (3) some organisms are able to withstand considerable
a steady state by the regulation of the extracellular body changes in the environment.
699
; a : contractile
- 4 vacuole
vacuole
membrane
vacuole
membrane
contractile
vacuole
Fig 19.3 Electron micrograph of contractile vacuole of Amoeba. Water is secreted into
tiny vesicles which fuse with the membrane of the contractile vacuole discharging water
into the vacuole
Investigations carried out into the function of the

19.4.1 Protozoa
contractile vacuole in the giant amoeba (Chaos chaos)
Protozoans are found in freshwater and show that the calculated osmotic influx of water based on
marine habitats and the body fluids of other organisms. The the osmotic pressure of the intracellular fluid agrees with
intracellular fluid of a protozoan is separated from the observed estimates of the volume eliminated by the
external environment only bya differentially permeable contractile vacuole. The contents of the contractile vacuole
plasma membrane. Excretion of carbon dioxide and are hypotonic to the intracellular fluid yet hypertonic to the
ammonia occurs by diffusion over the entire surface of the external medium. Several hypotheses have been put
cell. This has a relatively large surface area to volume ratio forward to account for formation of vacuolar fluid. A
which assists the removal of waste substances. probable explanation is shown in fig 19.5.
All freshwater species of protozoans are hypertonic to Many of the marine rhizopod protozoa do not have
their surroundings and have osmoregulatory organelles functional contractile vacuoles because their intracellular
known as contractile vacuoles. These are necessary to fluid composition is isotonic with sea water. This evidence
remove water which enters the cell by osmosis from the suggests that the primary role of the contractile vacuole is
hypotonic medium through the plasma membrane, and to osmoregulation.
regulate the volume of the cell and prevent it increasing in
size. The exact location and structure of the contractile 19.4.2 Coelenterata
vacuole is extremely variable. In Amoeba proteus a
contractile vacuole can form anywhere within the cell and Coelenterates do not appear to possess any
release its fluid into the external environment at any point excretory or osmoregulatory organs or organelles and the
on its outer surface (fig 19.3). In Paramecium aurelia there mechanism of osmoregulation is unknown. Carbon dioxide
and ammonia are the principal toxic metabolic waste
are two contractile vacuoles with fixed positions (fig 19.4).
substances and they are removed by diffusion from the cells
The method of functioning, however, appears to be similar
in all species and involves the movement of water from the directly into the water of the extracellular environment.
cytoplasm into small vesicles which fuse with, and empty
their water into, the contractile vacuole. Mitochondria
19.4.3 Platyhelminthes
collect around contractile vacuoles, and presumably supply Most of the metabolic waste products of
the energy for the ‘osmotic’ work of filling them. platyhelminths pass into the much-branched gut and are
700
eliminated via the mouth. Some, however, pass into a
series of tubules which form a joint excretory and
osmoregulatory system. This is a primitive type of
nephridium, known as a protonephridium, whose principal
function is osmoregulation. Protonephridia are found
mainly in animals that lack a true body cavity (coelom) such
ontractile
vacuole as the platyhelminths and rotiferans. Planaria hasa pair of
with canals protonephridia which run the entire length of the body and
open to the exterior via numerous excretory pores (fig
19.6a). Each protonephridium is made up of many tubules
which branch and end in enlarged hollow cells from which
cilia project into the tubule. If only one cilium is present the
cell is known as a solenocyte and if several are present it is
known as aflame cell (fig 19.6). The cilia of the flame cells
undulate, and this movement resembles the flickering of a
flame, hence the name. This ‘flickering’ appears to agitate
the fluid in the tubule and propel it along the nephridial
ducts towards the excretory pores. The fluid in the flame
cell is composed of water and waste substances produced
by the tissues. It is thought that some of these wastes are
secreted into the tubule by active transport and some by
ultrafiltration through the cytoplasm of the flame cell.
Water enters the lumen of the flame cell by osmosis. Flame
cells are found in some annelids and solenocytes are found
principally in the cephalochordate, Amphioxus.
19.4.4 Annelida
Annelids have a combined excretory and
osmoregulatory organ known as a metanephridium, or
simply a nephridium, which regulates the chemical com-
position of the body fluids. The exact structure and
distribution of nephridia varies in each of the three orders,
the Polychaeta, Oligochaeta and Hirudinea, but the basic
structure and function is similar in all. Nephridia are
unbranched tubules which link the coelom to the exterior.
In some species, such as the lugworm (Arenicola), the
nephridium is formed by the fusion of an ectodermal tubule
which opens to the exterior by a pore and a mesodermal
tubule or coelomoduct which opens into the coelom. The
earthworm (Lumbricus) has a pair of nephridia in each
segment apart from the first three and the last, but
polychaetes and leeches have fewer.
A nephridium consists of a ciliated funnel, the nephro-
stome, which leads via a long ciliated and muscular tubule to
Cae 0g a bladder where fluid is stored prior to its release through
contractile an external opening, the nephridiopore. The waste fluid is
vacuole called urine and is formed by the processes of ultrafiltra-
tion, selective reabsorption and active secretion.
Coelomic fluid containing useful and waste substances
passes into the nephrostome by pressure created by the
beating of cilia (fig 19.7). Fluid passes along the long
Fig. 19.4 Photomicrograph of the fixed contractile vacuoles narrow tube by the action of cilia and muscles but no
of Paramecium reabsorption of useful substances occurs here. The cells
lining the short middle tube and the longer wide tube
reabsorb useful substances into the blood capillaries in
their walls, whilst further waste is actively secreted into the
701
H,O
(a) f
(o)vesicle
is now hypotonic
|
vesicles contain water and
due to loss of K* ions
are isotonic with
vesicle and reduced uptake of Na*
cytoplasm
ions, but water is
retained due to the
impermeability of the
vesicle membrane to water
H,O
Nat Kt contractile
(d) vacuole
(6) vesicle fuses with
Na? ions are pumped into
contractile vacuole and Nate
the vesicles in exchange
Ke the contents are
for Kt ions so that Na* discharged. Nat ions lost ieee ae
influx is less than Kt ES
from the cell in the
efflux (see Na*/ K* pump,
vacuolar fluid are replaced
section 7.2.2)
by active transport from
the external medium
(a) (b) nucleus Fig 19.5 (above) Diagrammatic explanation of a possible

mechanism of water uptake by a contractile vacuole
flame cell
cytoplasm
Fig 19.6 (left) Features of the protonephridial excretory

system of platyhelminths. (a) Gross structure of the system in
Planaria, (b) single flame-cell
nephridial
excretory Aut
pore
excretory
Fig 19.7 (below) Stages in the formation of urine in the
canal earthworm. Heavy arrows show regions of active secretion of
substances. Protein is known to be present in the
nephrostome but not in the urine excreted by the
nephridiopore. At some stage it is removed from the
nephridium but as yet no mechanism is known to account for
this uptake. (Graph after Ramsay.)
water glucose
glucose _ amino acids
amino acids ammonia proteins ammonia
protein urea salts some salts urea water
salts
ammonia
urea | water
Oe ae | ammonia
ma Lo | urea
2 a SS
|
os? narrow tube | :
middle tube :
nephrostome wide tube
muscular tube |
100 bladder
nephridiopore
75
Urine
concentration as
percentage 50
of blood
concentration
DS
702
tubule from the capillaries. As substances are reabsorbed
the concentration of waste solutes in the urine rises and the
urine becomes more dilute. This urine, which is hypotonic
250
to coelomic fluid, is excreted through the nephridiopore.
The ability to produce a dilute urine Suggests that the
nephridium has an osmoregulatory function. Investiga-
tions have revealed that Lumbricus behaves as a freshwater
osmoregulator since, when placed in salt solutions of
various concentrations, it remains hypertonic to its en- 24 T2028)
| | al a
30) G2 aGNTISSs T40MNT 4? 544 ead
vironment yet produces a hypotonic urine. Although the Temperature
/°C
natural habitat of the earthworm appears terrestrial, it
actually lives in direct contact with the water films that Fig 19.8 (above) Graph showing the water loss from the
cuticle of a cockroach at various air temperatures (triangles).
surround the soil particles of the walls of its burrow. Hence The circles indicate water loss plotted against the surface
it may be considered to be a freshwater organism. The temperature of the cuticle. This shows the dramatic increase
osmoregulatory activities of typically marine polychaete in water loss at about 29.5 °C, the transition temperature
species such as Nereis diversicolor are described in section
19.4.6.
Fig 19.9 (below) Diagram showing the position of
19.4.5 Arthropoda Malpighian tubules in relation to the alimentary canal of
Rhodnius prolixus
Arthropods are adapted to conditions in a vast
anterior
range of habitats from marine to fully terrestrial. It is not posterior
surprising, therefore, that as a phylum they display a range

_--—~_
of excretory and osmoregulatory mechanisms. Adapta- eases a
tions shown by insects to terrestrial life and crustaceans to

marine, estuarine and freshwater life have been selected as —_——~
——S
~
representative of the range of features shown by arthro-
pods and are described in this section.
One of the major problems of life on land is the
prevention of water loss. Insects have an almost imper- Malpighian
tubule
meable cuticle to reduce water loss from the body surface
and spiracles to reduce water loss from the gaseous ex-
change system of tracheae and tracheoles.
The strong cuticle is composed of a chitinous exo- and
endocuticle covered by a thin waterproof layer, the The problem of preventing water loss by excretion is
epicuticle (0.3 um thick) as described in section 4.10.1. overcome by specialised excretory organs called Mal-
Water loss by evaporation is prevented by the impermeable pighian tubules which produce and excrete the almost
properties of the epicuticle produced by a highly organised insoluble waste substance uric acid.
monolayer of lipid molecules covered by several layers of Malpighian tubules are blind-ending tubules which lie in
irregularly orientated lipid molecules. If these wax or the intercellular space of the abdomen and are bathed in
grease lipid layers are abraded by sharp particles, such as haemolymph. The number of tubules is variable in insects,
sand or alumina, the evaporation rate increases and the some have a pair and others may have several hundred.
insect risks dehydration. Interestingly, as the air tempera- Rhodnius, a blood-sucking hemipteran, has four tubules;
ture surrounding an insect is increased steadily there is a in all cases they open into the hindgut at its junction with
gradual increase in the rate of evaporation until a particular the midgut and may be long and slender or short and
temperature is exceeded after which the evaporation rate compact (fig 19.9).
increases rapidly. This point is known as the transition Wigglesworth investigated the function of the tubules in
temperature. If the water loss from the insect is plotted uric acid formation in Rhodnius and the mechanism
against the insect’s surface temperature this transition appears to be as follows.
point can be seen more clearly and marks the temperature The tubule has two histologically distinct regions, an
at which the ordered orientation of the wax monolayer upper segment (distal to the gut), composed of a single
breaks down, as shown in fig 19.8. layer of cells and containing a clear solution, and a lower
Some insects living on dry food in very dry habitats are segment. The cells of the lower segment have microvilli on
able to take up water from the air providing that the their inner surface and it is here that crystals of uric acid
relative humidity of the air is above a certain value, such as precipitate out of solution (fig 19.10). The contents of the
90% for the mealworm (Tenebrio) and 70% for the house tubule pass into the hindgut or rectum where they mix with
mite (Dermatophagoides). waste materials from digestive processes. Rectal glands in
703
the wall of the rectum absorb water from the faeces and Water and solutes are filtered from the haemolymph into
uric acid suspension until the waste is dry enough for it to the end sac by the hydrostatic pressure of the blood. As this
be eliminated from the body as pellets. filtrate passes through the spaces in the glandular lining of
Fully terrestrial organisms do not have the same the labyrinth and along the nephridial canal, useful
osmoregulatory problems as aquatic or semi-terrestrial materials are selectively reabsorbed into the haemolymph
organisms. Insects, however, do have to regulate the ionic and further waste substances, including nitrogenous waste,
contents of their haemolymph and this is achieved by are secreted into the filtrate.
maintaining a balance between ions taken up in the diet and Astacus is hypertonic to its environment and takes in
those lost through synthesis, egestion and excretion. It is water through any permeable surface especially the gills.
aquatic arthropods such as the freshwater crustacean Large volumes of urine, which is hypotonic to
Astacus, the crayfish, and the marine crustacean Carcinus haemolymph, are produced to counteract this osmotic
maenas, the shore crab, that show most adaptations of their influx. Most of the nitrogenous waste is lost as ammonia but
excretory and osmoregulatory organs to their habitats and some urea is also produced.
modes of life. Carcinus lives in the intertidal zone of the sea-shore,
Astacus lives in freshwater streams which provide a surrounded for most of the time by sea water. It has
hypotonic environment. Some waste nitrogenous sub- antennal glands similar to those of Astacus which it uses to
stances and carbon dioxide are deposited in the cuticle of eliminate waste nitrogenous material, especially ammonia.
the nymphal stages and shed during moulting, but in the In common with many other marine species, parts of its
adult, nitrogenous waste is removed as ammonia through surface are permeable to salts and water and the body fluids
specialised excretory/osmoregulatory organs known as are isotonic with the sea water. This is an economical
antennal or green glands. measure for marine organisms as they do not need to
The antennal glands are blind-ending mesodermal expend energy in maintaining their body fluids at a higher
structures which lie in the haemocoel just in front of the or lower osmotic potential than their environment. How-
mouth region and open to the exterior by a pore situated ever, even though they are isotonic, the ionic concen-
underneath the base of the antennae. Each antennal gland tration of the body fluids may be maintained ata different
is composed of four regions, a blind-ending sac, a green level from that of the sea, and energy is required for ionic
tube known as the labyrinth, a long white nephridial canal regulation.
and a thin-walled bladder which opens to the exterior (fig Carcinus is found in a wide range of habitats. It can
19.11). tolerate the hypertonic conditions which may be encoun-
tered in rock pools on sea-shores and salt pans in salt
H,U marshes as water evaporates from them on hot days. In
uric acid these conditions it acts as an osmoconformer and decreases
haemolymph @
the osmotic pressure of its body fluids by retaining salts
KHU+ CO,+2H,0 KHCO, + H,O which the tissues tolerate. In these conditions the urine is
hypertonic to the body fluids. Carcinus is able to tolerate
the changing conditions found in estuaries and, in keeping
with many species found in estuaries, it is euryhaline. As
the salt concentration of the water decreases due to the
KHU + CO, 4 1H,0 DL» KHCO, + H,0+(H,U> diluting effects of river water or rainfall the body fluids of
uric acid Carcinus become hypertonic to the surrounding medium
(fig 19.12). Water tends to enter by osmosis and solutes
Malpighian leave by diffusion. Under these conditions Carcinus
tubule upper segment lower segment becomes an osmoregulator and maintains the body fluid
composition at a steady state by actively secreting sodium
from the urine back into the haemolymph through the cells
Fig 19.10 Summary of the suggested mechanism of uric of the antennal gland. Some water will be retained as the
acid excretion by the Malpighian tubule. (1) Uric acid (H2U) sodium is reabsorbed but the overall volume of the
produced by the cells of the body is secreted into the organism is prevented from increasing due to the tough
haemolymph where it combines with sodium and potassium cuticle. In these conditions the urine is isotonic to the body
hydrogencarbonates and water to form sodium and
potassium urates (NaHU and KHU), carbon dioxide and water fluids.
(only potassium is shown in the diagram). (2) These salts are
actively secreted into the lumen of the tubule and water
follows by osmosis. (3) As these soluble substances pass 19.4.6 Effects of changes in the
down the tubule hydrogencarbonates form and these are environment shown by invertebrates
actively reabsorbed into the haemolymph. Water follows the
hydrogencarbonates by osmosis and as the pH in the lower Invertebrates which live in sea water, such as
segment fails, due to the reabsorption of hydrogencarbonates, sea anemones, are isotonic with their environment. If
the uric acid precipitates out as crystals placed in diluted sea water they immediately take up water
704
glucose
amino acids |
water ; USO 3
StnRGnts ammonia amino acids water
salts
ammonia
urea urea
salts amino acids
hypotonic
end sac labyrinth urine
white tube bladder
200
SS
dm~3
concentration/uM
Chloride
0
Fig 19.11 Diagram summarising the structure and function of the antennal (green) gland of Astacus. The graph indicates
changes in the osmotic potential of the filtrate. (After Peters from Barrington.)
and lose ions. This is also seen by exposing Arenicola to uptake of ions by drinking water and actively secreting ions
decreasing salinities. After an initial increase in mass the back into the sea through their gills. These mechanisms of
mass returns to normal. This is explained by the fact that control are so efficient that shrimps are able to maintain
water enters by osmosis faster than ions are lost. After a their body fluid level at a steady state in external
short while conditions become stable due to loss of ions, the concentrations ranging from 2 to 110% sea water. It is
body fluids become isotonic with the environment and the thought that this ability reflects the freshwater ancestry of
mass returns to normal provided the salinity does not fall the species which have secondarily invaded sea water.
too low. Nereis is normally found in marine conditions in The brine shrimp (Artemia) is hypotonic to the extreme-
sand and mud but it can osmoregulate down to salinities of ly high salinities of its environment and maintains its body
10% sea water and inhabit estuaries. It is an osmoregulator fluids in a hypotonic state by continually removing sodium
and is able to remain hypertonic to the environment and chloride ions from the concentrated medium which it
because of the reduced permeability of its body surface and drinks. It does this by eliminating them from the
its ability to take up chloride ions from its environment haemolymph through the epithelium of the gills. It is also
(fig 19°12): able to maintain its body fluids at a steady state despite
There are some marine species that are hypotonic to sea fluctuations of 10-1 000% sea water in the external
water, such as the shrimps Palaemonetes and Leander (fig environment.
19.12). They have overcome problems of water loss and Some brackish water species, such as the mussel
(Mytilus), are able to maintain the intracellular compo-
sition of their cells at a fairly steady state despite changes in
OoaS aSo 5 f
z¢-absolute osmotici conformation
Z the composition of the external environment and the ex-
sea water ZZ
7.35 ,#-— — absolute osmotic regulation tracellular fluid. This is thought to be due to regulation of
sé in ea zat
S30)
eee ee
een ee
oboe
2 -eoeae the fluid composition by individual cells.
E
Ss
Bete
Pa
4 Many freshwater species, such as Astacus, are thought to
6 20
te =
Bia
a
api have developed from marine species via brackish water
= oe ee species by progressive improvements in the efficiency of
=o 10+ ieme sea water Pal
alaemonetes
gs oo
water and ion regulation.
E a ——— Nereis Small freshwater invertebrate animals have arelatively
&S ee L | | J pat? s :
10 20 30 35 40 Leande1 large surface area to volume ratio. Since they are
external environment /%) NaCl ----- Carcinus hypertonic to their environment they tend to take in water
and lose salts more rapidly than comparable larger
Fig 19.12 Graph showing the relationship between internal invertebrates, such as Astacus, that have a smaller surface
and external environments for osmotic conformers, osmotic area to volume ratio. They have overcome this problem, to
regulators and four marine species, the prawns, Leander and
a certain extent, by having body fluids with a lower ionic
Palaemonetes, the ragworm, Nereis and the shore-crab,
Carcinus concentration than larger forms.
705
&
____» efferent arteriole
19.4.7. | Echinodermata afferent arteriole——->
All echinoderms are marine. Nitrogenous glomerulus

waste is removed from the gills and tube-feet by diffusion as
ammonia and urea. Echinoderms have a water vascular
system containing sea water with which the cells are primitive
renal capsule
isotonic. There is no problem of osmoregulation. corpuscle of
nephron
19.4.8 Vertebrata
glomerular
All three nitrogenous waste products are filtrate
produced
excreted by vertebrates and, with few exceptions, the form here
of the waste product is related to the environmental
tubule
availability of water. Osmoregulatory mechanisms in of
vertebrates are more efficient than in invertebrates because nephron
of the reduced permeability of the body surface and the
development of the kidney. Biologists are uncertain as to
whether the earliest fish originated in the sea or in fresh
water. Many biologists favour a marine origin and see the ciliated
kidney as a later development necessary for survival in the funnel
hypotonic conditions of fresh water. In this environment it
provides a mechanism for the removal of excess water and
the retention of salts. Subsequent development of the
kidney is related to the environment of the organism, and coelom
shows a progressive increase in complexity throughout the
Fig 19.13 Structure of a primitive nephron. Filtrate produced
vertebrate classes from the fish to mammals. This increase from the glomerulus passes to the coelom via the ciliated
in complexity is associated with colonisation of the funnel or joins up with other nephrons via connecting tubules
terrestrial environment. The increased efficiency of these
mechanisms maintains the internal composition of the
body fluids within narrower limits than in the invertebrates. The vertebrate kidney relies on the principles of
The basic unit of the kidney is the nephron. Nephrons are ultrafiltration, selective reabsorption and active secretion
segmental structures formed from mesodermal nephro- for the production of urine. Urine is a liquid containing
tomes (section 21.8) which have become intimately nitrogenous waste, water and ions in excess of those
associated with blood vessels from the aorta and are linked required by the body. Ultrafiltration does not discriminate
to the coelom byaciliated funnel. The nephrons of fish between those substances which are useful to the body and
larvae show their most primitive arrangement where those which are not and energy is expended in reabsorbing
several of them open into the pericardial cavity and form a 99% of the solutes back into the blood. However, despite
collective structure known as the pronephros (fig 19.13). the apparent inefficiency in terms of energy, this mechan-
All adult fish and amphibia lose the pronephros and have a ism gives vertebrates greater flexibility to exploit new
more compact structure made up of many more nephrons habitats since it permits ‘foreign’ or ‘new’ substances to be
and found in the abdominal and tail regions of the excreted as they are encountered. It does not require the
organisms. This is called a mesonephric kidney. Its development of new secretory mechanisms to remove
nephrons have lost their connections with the coelom and these substances.
are linked together by a collecting duct which leads to the Pisces
urinogenital opening. Such a structure is ideal for the
The excretory and osmoregulatory organs of the fish are
production of dilute urine produced predominantly by
gills and kidneys. Both structures are permeable to water,
organisms living in a freshwater environment.
nitrogenous waste and ions and have a large surface area to
Reptiles, birds and mammals are adapted to life on land
facilitate exchange. The kidney, unlike the gill, is separated
where problems of water removal experienced by fish and
from the external environment by the body wall, the body
amphibians are replaced by problems of water retention.
tissues and extracellular body fluid, and can therefore
These organisms have an even more compact structure, the
exercise control over the steady-state composition of the
metanephric kidney which contains far more nephrons with
body fluids. Despite the fact that fish live in an aquatic
longer tubules for water reabsorption. These tubules
environment there are enough differences between the
eventually release concentrated urine into the central
cavities of the kidney. From here it passes to the bladder by mechanisms of excretion and osmoregulation in freshwater
a tube known as the ureter. (The detailed structure and and marine species for them to be described separately.
function of the mammalian kidney is described in section Freshwater fish. Freshwater teleosts have
oe) an internal osmotic concentration of approximately
706
300 mOsm dm™ and are hypertonic to their environment. presence but also to be able to tolerate high levels of it.
Despite having a relatively impermeable outer covering of Investigations carried out on isolated shark hearts have
scales and mucus there is a considerable osmotic influx of shown that they will only beat if perfused with a balanced
water and loss of ions through the highly permeable gills, salt solution containing urea. The body fluids contain
which also serve as the organs for the excretion of the waste between 2 and 2.5% urea, which is about 100 times the
nitrogenous substance ammonia. In order to maintain the concentration which can be tolerated by other vertebrates.
body fluids at a steady state, freshwater fish have to High concentrations of urea normally disrupt the hydrogen
continually lose a large volume of water. They do this by bonding between amino acids and denature proteins so
producing a large volume of glomerular filtrate from which inhibiting enzyme activity, but not, for some reason, in
solutes are removed by selective reabsorption into the elasmobranchs. Urea, together with inorganic ions and
capillaries surrounding the kidney tubule. The kidneys another nitrogenous waste substance trimethylamine oxide
produce a large volume of very dilute urine (hypotonic to ((CH;);N=O) which is less toxic than ammonia, increases
blood) which contains some ammonia and a number of osmotic pressure of the body fluids above that of sea water
solutes. Up to one-third of the body mass can be lost per (A sea water —1.7°C, A elasmobranchs —1.8 °C) (fig
day as urine. Ions which are lost from the body fluids are 19.14). Elasmobranchs, being therefore slightly hypertonic
replaced from food and by active uptake from the external to the environment, take in water by osmosis through their
environment by special cells in the gills. gills. This is eliminated along with excess urea and
trimethylamine oxide by the kidneys in a urine which is
Marine fish. Fish are thought to have origin- slightly hypotonic to the body fluids. The kidney has long
ated in the sea and, having successfully invaded the tubules which are used for the selective reabsorption of
freshwater environment, secondarily re-invaded the sea urea but not for the elimination of salts which enter with
and given rise to the elasmobranchs and marine teleosts. the diet. Excess sodium and chloride ions are removed
Whilst in the freshwater environment many of the basic from the body fluids by active secretion into the rectum by
metabolic activities and organ functions evolved based on the cells of the rectal gland, which is a small gland attached
an osmotic pressure of the body fluids at a level between to the rectum by a duct. The gills are relatively imper-
one-third and one-half that of sea water. On their return to meable to nitrogenous waste and any which is lost from the
the sea the body fluids retained their ancestral osmotic body is controlled by the kidney. In this way the osmotic
pressure and this produced problems of maintaining the pressure of the body fluids is maintained at a high level.
body fluids at a steady level in a hypertonic environment
(fig 19.14). Marine teleosts. Marine teleosts maintain
their body fluids at an osmotic pressure which is hypotonic
to sea water (fig 19.14). The body surfaces are relatively
2 = seawater
impermeable to water and ions due to scales and mucus but
the body fluids lose water and gain ions freely through the
rc.
gills. In order to regulate the composition of the body fluids
teleosts drink sea water, and secretory cells in the gut
remove Salts by active transport into the blood. Chloride
secretory cells in the gills actively remove chloride ions
Ao
NB
BO
OW
concentration/a°C
Blood
eooo
eee
ee from the blood into the sea and sodium ions follow to
Elasmobranchs Teleosts Reptiles Birds Mammals maintain electrochemical neutrality. The other major ions
in sea water, magnesium and sulphate, are removed in the
Fig 19.14 Approximate body fluid concentrations of marine
small volume of isotonic urine which is produced by the
vertebrates. Elasmobranchs are the only vertebrates with a
body fluid hypertonic to the environment but as the graph kidneys. The kidneys lack glomeruli and are therefore
shows the basic osmotic concentration is a little higher than unable to carry out ultrafiltration. All the contents of the
that of teleosts. The retention of urea increases the osmotic urine — such as the nitrogenous substance trimethylamine
pressure to that of sea water, as shown by freezing point oxide which gives fish their characteristic smell — and salts,
depressions (A°C = change in °C) are secreted into the kidney tubules and water follows by
osmosis.
Elasmobranchs. The basic osmotic pressure
of body fluids in elasmobranchs is approximately the same Euryhaline fish. There are several euryha-
as that of marine teleosts, that is, equivalent to a 1% salt line fish which not only tolerate slight changes in salinity
solution. In order to prevent excessive water loss to the but are able to adapt totally to freshwater and marine
hypertonic sea water elasmobranchs have increased the conditions for major periods of their lives. Depending
osmotic pressure of their body fluids by synthesising and upon the direction in which they move to spawn these fish
are described as anadromic or catadromic. Anadromic
retaining urea within their tissues and body fluids. The
(ana, up; dramein, to run) species such as the salmon
majority of the cells in the body, with the exception of brain
(Salmo salar) hatch in fresh water and migrate to the sea
cells, appear capable of synthesising urea and their
where they mature before returning to fresh water to
metabolic activities appear not only to depend on its
707
(a) Amphibia
Amphibians are believed to have developed from an
ancestral stock of freshwater fish, and have inherited
similar problems of osmoregulation in that their blood
hypertonic
fluids are hypertonic to their freshwater environment. The

water,
skin of a frog is permeable to water, and most of the water
large volume from the environment is absorbed across the skin. The
See
it very dilute
urine excess water which enters the body fluids is removed by
containing ultrafiltration through the many large glomeruli of the
ammonia
kidney.
The amphibian kidney has been extensively used in
(0)
investigating the physiology of the kidney because its large
glomeruli are situated near the surface of the kidney. These
iso-/hypertonic are accessible to micropipettes which can be inserted into
(Li the glomerulus and tubules and the filtrate removed for
analysis. In this way the effectiveness of ultrafiltration and
trimethylamine oxide selective reabsorption can be demonstrated. Amphibia
excess urea }kidney
some water water produce a large volume of very dilute urine (hypotonic to
ay
ao }rectal gland body fluids). This contains urea which is eliminated by
secretion into the tubule as well as by ultrafiltration. The
advantage of this mechanism is that it allows amphibians, in
()
dry conditions, to reduce the amount of ultrafiltration by
sea water
the glomerulus and thereby reduce water loss in the urine;
at the same time the tubules continue to receive blood from
) hypotonic the renal portal system from which they still actively secrete
urea into the tubule. In this respect the mechanism is
water
ae opposite to that seen in elasmobranchs where urea is
actively reabsorbed by the tubule.
salts small volume isotonic urine
containing trimethylamine Some salts are inevitably lost in the urine and by
< oxide diffusion through the skin, but these are replaced from the
active transport of C17,
Nat follows passively diet and by their active uptake from the environment by the
Fig 19.15 Excretion and osmoregulation in (a) freshwater
skin which is the main organ of osmoregulation. The larval
teleosts, (b) elasmobranchs and (c) marine teleosts stage of the amphibia, the tadpole, is completely aquatic
and excretes ammonia through its gills, but at metamor-
phosis both the nitrogenous product and the mechanism of
spawn. Catadromic (cata, down) species such as the eel excretion changes, as described earlier in this section.
(Anguilla vulgaris) move in the opposite direction. They Frogs are able to store water in their bladders and the
hatch in sea water and migrate to fresh water where they many subcutaneous lymph spaces. This replaces water lost
mature before returning to the sea to breed. When the eel by evaporation during periods when they are on land.
moves from fresh water to sea water it loses about 40% of Toads are able to tolerate dry conditions for longer periods
its body mass in ten hours. To compensate for this and of time because their skin is less permeable to water and
remain hypotonic it drinks sea water and eliminates ions by because their kidneys are able to reabsorb water from the
active secretion from the gills. When the eel moves from glomerular filtrate and produce a more concentrated urine.
sea water to fresh water the mass initially increases as water The permeability of amphibian skin is known to be
enters by osmosis but it is able to achieve an osmotic steady controlled by an antidiuretic hormone produced by the
state after only two days. Whilst in fresh water, salts are posterior pituitary gland. The mechanism of control of
taken up by active transport through the gills.
permeability is thought to be similar to that of the
These two species show that the active transport mammalian kidney tubule.
mechanisms in the gills are able to work in two directions.
Whether this involves a change in the direction of pumping Water balance in terrestrial organisms
of the same cells or the operation of different sets of cells is The efficient functioning of animal cells relies on the
not known. It is thought that hormones from the pituitary maintenance of the steady state of the intracellular fluid of
gland and the adrenal cortex may influence these mechan- cells. Homeostatic exchanges in water content between
isms. In both types of fish a period of ‘lying up’ is necessary cells, tissue fluid, lymph, plasma and the environment
when they enter freshwater environments to allow their present problems to both aquatic and terrestrial forms of
osmoregulatory mechanisms to adapt to the new surround- life. Aquatic organisms gain or lose water by osmosis
ings. through all permeable parts of the body surface depending
708
on whether the environment is hypotonic or hypertonic. Terrestrial reptiles reduce their water loss by having a
Terrestrial organisms have the problem of losing water and relatively impermeable skin covered by keratinised scales.
many mechanisms are employed to maintain a steady-state The gaseous exchange organs are lungs situated inside the
water balance as summarised in table 19.5. This steady body to reduce water loss. The tissues produce insoluble
state is achieved by balancing water loss and water gain, as uric acid which can be excreted without the loss of much
shown in table 19.6. water. Excess sodium and potassium ions require water for
Table 19.5. Summary of water conservation mechanisms their removal but, since conservation of water is critical,
these combine with uric acid to form insoluble sodium and
shown by terrestrial organisms
potassium urates which are excreted with uric acid. The
glomeruli are small and produce only enough filtrate to
Organism Water conservation mechanism
wash out the uric acid from the kidney tubules into the
insect impermeable cuticle cloaca where more water is reabsorbed. Many terrestrial
trachea and spiracles reptiles have no glomeruli at all.
Malpighian tubules There is no special mechanism for the removal of salts in
uric acid as nitrogenous waste terrestrial reptiles and the tissues are able to tolerate
cleidoic egg
increases of up to 50% the normal level of salts following
reptile scales and keratinised skin the ingestion of food or when water loss is excessive.
lungs
metanephric kidney Marine reptiles, such as the Galapagos iguana and the
uric acid as nitrogenous waste edible turtle (Chelone mydas) obtain a great deal of salt
cloacal reabsorption’ from their food. Their kidneys are unable to remove the
cleidoic egg rapid influx of salt from the body fluids and rely on
behavioural responses to heat specialised salt glands in the head. These glands are able to
physiological tolerance to dehydration
secrete a concentrated solution of sodium chloride which is
bird feathers
several times stronger than sea water. In the turtle the salt
lungs
metanephric kidney glands are found in the orbit of the eye and ducts lead from
uric acid as nitrogenous waste them to the eyes. They give the impression of ‘crying’ and
cleidoic egg produce tears with a very high salt concentration.
cloacal reabsorption
The cleidoic egg
mammal keratinised skin and hair
lungs An important characteristic of both reptiles and birds
metanephric kidney which has enabled them to become totally independent of
hypertonic urine containing urea water in their life cycle is the cleidoic egg (fig 20.52). This is
viviparity
behavioural response to heat an egg enclosed in a tough shell which protects the embryo
restricted ecological range from dehydration. During embryonic development an
physiological tolerance to dehydration outgrowth of the hindgut produces a sac-like structure
called the allantois which stores uric acid produced by the
embryo. Since uric acid is insoluble and non-toxic it forms
Table 19.6. Summary of water balance mechanisms in an ideal storage excretory material for the embryo. In the
terrestrial organisms later stages of development the allantois becomes vascula-
rised and pressed up against the shell where it functions as a
water loss from body = water gained by body gaseous exchange organ.
evaporation from body surface drinking Birds

evaporation from gaseous ex- water in food
uptake through body surface
Birds are believed to have developed from a stock of
change surface
water in faeces respiration (metabolic water) terrestrial reptiles, such as snakes and lizards, and
water in urine inherited the same problems. The skin of birds is relatively
water in secretions such as tears impermeable to water and the evaporation rate is kept low
ee eee ao by the presence of feathers and the absence of sweat
glands. However water loss from gaseous exchange surfaces
Reptilia is considerable due to the high ventilation rate and the
Reptiles are primarily adapted to life in the terrestrial maintenance of a relatively high body temperature. As a
environment and show many morphological, biochemical consequence of the increased metabolic rate some small
and physiological adaptations to life on land. However, all birds may lose up to 35% of their body mass per day.
three orders, the turtles, lizards and snakes, and crocodiles Nitrogenous waste is eliminated as uric acid in urine
and alligators, have species which are secondarily adapted which is hypertonic to the body fluids. The urine passes into
to life in the freshwater and marine environments. In all the cloaca where more water is removed from the urine and
cases the excretory and osmoregulatory mechanisms of faeces before the almost solid residue is released from the
these organisms show adaptations to these environments. body.
709
The kidney of birds has small glomeruli and all the blood Fig 19.16 7S through mammalian kidney showing the
supplying the tubule, where reabsorption of water and position of a cortical nephron and a juxtamedullary nephron
secretion of salts occurs, comes from the glomerulus, which fibrous capsule
relies on a relatively high blood pressure for efficient
functioning. Consequently the production of a large
volume of glomerular filtrate and the subsequent reabsorp- juxtamedullary
nephron
tion of most of its water and salts have become linked. The
surface area of the tubule is increased, to facilitate this
reabsorption, by the development of a loop of Henle. The \ cortical
i nephron
physiology of this structure enables the concentration of
L> \ collecting duct
uric acid in the urine to reach 21% which is approximately
three thousand times its concentration in the body fluids. cortex ;
papilla
Some marine birds, such as the penguin, gannet,
cormorant and albatross, which feed on fish and sea water,
medulla pk ee of
absorb large quantities of salts. These are removed from col lecting duct
the body fluids by specialised salt-secreting cells in the salt, FY |
aN
/ pelvis
or nasal, gland. The nasal glands are similar to those of
marine reptiles and are situated in the orbit. They secrete a calyx
sodium chloride solution that is four times stronger than pyramid

that of the body fluids. The nasal glands are composed of ureter
many lobules made up of a large number of secretory
tubules that open into a central cavity leading to the nasal
cavity where the salt is released as large droplets or blown
out in a fine spray.
A transverse section of the kidney shows two distinct

19.5 The mammalian kidney regions, an outer cortex and an inner medulla (fig 19.16).
The cortex is covered by a fibrous capsule, and contains
The kidney is the major excretory and glomeruli which are just visible to the naked eye. The
osmoregulatory homeostatic organ of the mammalian body medulla is composed of tubules, collecting ducts and blood
and has functions which include removal of metabolic vessels grouped together to form renal pyramids. The
waste products and ‘foreign’ molecules; regulation of the apices of the pyramids are called papillae, and they project
chemical composition of body fluids by removal of into the pelvis which is the expanded origin of the ureter
substances in excess of immediate requirements; regula- (fig 19.16). A large number of blood vessels run through
tion of the water content of body fluids, and thus the the kidney and supply a vast network of blood capillaries.
volume of the body fluids, and regulation of the pH of body
fluids. 19.5.2 Nephron — gross structure and blood
The kidney has a rich blood supply and regulates the supply
blood composition at a steady state. This ensures that the
The basic structural and functional unit of the
composition of the tissue fluid, and consequently the cells
kidney is the nephron and its associated blood supply. Each
bathed by it, is maintained at a level which coincides with
kidney, in Man, contains an estimated one million
the optimal conditions of the cells and enables them to
nephrons each having an approximate length of 3 cm. This
function effectively and efficiently at all times.
offers an enormous surface area for the exchange of
materials.
19.5.1 Position and structure of kidneys
Each nephron is composed of six regions having
There are a pair of kidneys in Man situated at strikingly different anatomical features and physiological
the back of the abdominal cavity, on either side of the functions:
vertebral column, in the region of the thoracic and lumbar (1) renal corpuscle (Malpighian body), composed of
vertebrae. Each kidney weighs about 0.5% of the total Bowman’s capsule and glomerulus,
body mass, and the left kidney lies slightly anterior to the (2) proximal convoluted tubule,
right. (3) descending limb of the loop of Henle,
The kidneys receive blood from the aorta via the renal (4) ascending limb of the loop of Henle,
arteries, and the renal veins return blood to the inferior (S) distal convoluted tubule, and
vena cava. Urine formed in the kidneys passes bya pair of (6) collecting duct.
ureters to the urinary bladder where it is stored until it can The structural relationship between these regions is shown
be released conveniently via the urethra. in fig 19.17.
710
-—— efferent arteriole Z
afferent arteriole_»
inher <,
wits sen
renal ; | (J () :
distal OeC
corpuscle ) Bowman’s convoluted R
capsule tubule att
le
proximal x
convoluted
tubule
thick A ;
ascending collecting
limb duct
M
’ Es
loop of Henle D
U
Ie
| | L
(ao A
1 |
ft thin duct of
thin
descending ascending renal papilla Bellini
limb limb
Fig 19.17 Diagram showing the structure of a nephron. (Not to scale.)
There are two types of nephrons, cortical nephrons and medulla. They have long descending and ascending tubules
juxtamedullary nephrons, which differ in their positions in of the loop of Henle which extend deep into the medulla
the kidney. Cortical nephrons are found in the cortex and (fig 19.18). The-significance of the two types of nephrons
have relatively short loops of Henle which just extend into relates to differences in their use. Under normal conditions
the medulla. Juxtamedullary nephrons have their renal of water availability the cortical nephrons deal with the
corpuscle close to (= juxta) the junction of the cortex and control of plasma volume, whereas when water is in short
(6) efferent arteriole

(a) afferent arteriole
glomerulus
Cc => pertabelan interlobular artery
O capillaries
R
a
E
x
arcuate artery —-»
arcuate vein——» |)
wart as tateSpee
vein
-
E
D
U loop of vasa recta
LD collecting
L duct Henle
A
——=
nephron
Fig 19.18 Nephrons and their associated blood supply; (a) cortical nephron, (b) juxtamedullary
Al
supply increased water retention occurs through the Processes
juxtamedullary nephrons. (1) Ultrafiltration. All small molecules, such as water,
Blood enters the kidney by the renal artery which glucose and urea, are filtered out of the blood plasma in
progressively branches into interlobar arteries, arcuate the glomerulus and produce a filtrate in Bowman’s
arteries and interlobular arteries before passing to the capsule which passes into the tubule of the nephron.
glomerulus of the renal corpuscle as an afferent arteriole. (2) Selective reabsorption. All substances useful to the
A reduced volume of blood leaves the glomerulus by an body and required to maintain the water and salt
efferent arteriole and flows through a network of peritubu- composition of the body fluid at a steady state are
lar capillaries in the cortex which surround the proximal removed from the filtrate and reabsorbed into the
and distal convoluted tubules of all nephrons and the loop blood capillaries, for example glucose in the proximal
of Henle of cortical nephrons. Arising from these capillar- convoluted tubule.
ies are the capillaries of the vasa recta which run parallel to (3) Secretion. Further substances not required by the body
the loops of Henle and the collecting ducts in the medulla. may be secreted into the filtrate, by cells of the
The function of both of these vascular networks is to return nephron, before it leaves the kidney as urine, for
blood, containing substances which are useful to the body, example K*, H* and NH," in the distal convoluted
to the general circulation. Blood flow through the vasa tubule.
recta is much less than through the peritubular capillaries
and this enables a high osmotic pressure to be main- Mechanisms
tained in the interstitial tissue of the medulla. This high (1) Active transport. Molecules and ions are secreted into
osmotic pressure is vital in producing a concentrated urine. and out of the filtrate as described in (2) and (3) above,
for instance the uptake of glucose into the peritubular
19.5.3 The role of the kidney capillaries surrounding the proximal tubule and the
Apart from their function in the removal of removal of NaCl from the thick ascending limb.
metabolic waste, the kidneys are homeostatic organs and (2) Differential permeability. Various regions of the
maintain the body fluid composition at a steady state nephron are selectively permeable to ions, water and
despite wide fluctuations in water and salt uptake. A urea, for example the proximal tubules are relatively
shortage of water, excess water uptake, excessive sweating, impermeable compared with the convoluted tubule;
a shortage of salts or excess of salts would all have serious the permeability of the collecting duct can be modified
consequences for the body if the kidney was unable to by hormones.
adapt its function to these changes. Yet it is only the last (3) Concentration gradients. A concentration gradient,
two regions of the nephron, the distal convoluted tubule varying from 300 mOsm kg" of water in the cortex to
and the collecting ducts, where changes in function occur to 1 200 mOsm kg™' of water at the papillae, is main-
regulate the body fluid composition. The rest of the tained within the interstitial region of the medulla of
nephron up to the distal convoluted tubule is stereotyped Man asa result of (1) and (2) above.
and functions in the same way in all physiological states. (4) Passive diffusion and osmosis. Sodium and chloride ions
The waste product of kidney function is urine whose and urea molecules will diffuse either into or out of the
volume and composition varies, reflecting the physiological filtrate, according to the concentration gradient, wher-
state of the body. Normally there is a copious supply of ever the nephron is permeable to them. Water
dilute urine, but at times when the body is short of water a molecules will pass out of the filtrate into a hypertonic
concentrated urine is produced. interstitial region of the kidney wherever the nephron is
permeable to water.
19.5.4 Basic principles of kidney function (S) Hormonal control. The regulation of water balance in
the body and salt excretion is achieved by the effects of
A logical explanation of kidney function based hormones acting on the distal convoluted tubule and
on following the course of filtrate through the nephron is
collecting ducts, such as anti-diuretic hormone, aldo-
made difficult because events in one region of the nephron
sterone and other hormones.
have important consequences for events in its other
regions.
Urine formation and the regulation of the steady state of
19.5.5 Methods of study of kidney function
the body is a dynamic process involving the removal of Investigations into the functioning of the
substances from one part of the nephron to another, such as kidney may be carried out directly on the exposed kidney,
the collecting duct to ascending limb, and from nephron to as in the case of studying the formation of urine, or
capillaries surrounding the nephron. indirectly, by analysing the relative composition of plasma
The purpose of this section is to outline the general and urine.
principles which underlie nephron function in terms of The American physiologist A. N. Richards showed that
processes and mechanisms. This provides a background for the composition of plasma and glomerular filtrate was
the detailed study of later sections. similar, by inserting micropipettes into the renal corpuscles
TA
of amphibian kidneys and removing samples of filtrate for
analysis. Amphibian kidneys were used for these studies
because their renal corpuscles are relatively large and near
to the surface of the kidney. This technique of renal
micropuncture and withdrawal of fluid has been extended
to other parts of the nephron and to mammals where it has
yielded much information concerning the functions of
these various regions.
Despite these advances there are still many kidney
functions which are far from clear, for example the function
of the loop of Henle. In these cases current knowledge is
only hypothetical and based upon models of how these
functions may be carried out.
Various techniques, based on the measurement of
plasma clearance, have been used to investigate the rates of
filtration and reabsorption by the kidney and the rate of
blood flow to the kidney. Plasma clearance is a measure of
the rate at which substances are removed from the plasma
by the kidney.
The rate at which plasma is filtered by the glomeruli, the
glomerular filtration rate (GFR) can be calculated directly
from the plasma clearance value for a substance, such as
the polysaccharide inulin (C,H,,O;),,, which is filtered from
the blood and not reabsorbed or secreted. For Man the
glomerular filtration rate for both kidneys is about
125 cm’ per minute.
19.5.6 Nephron — structure and function

The renal corpuscle consists of a blind-ending ®
tube which invaginates to form a double-walled epithelial Fig 19.19 Glomeruli and arterioles of dog kidney injected
capsule called Bowman’s capsule. This encloses the with silicone rubber. The tissues have been dissolved away
glomerulus, consisting of a knot of about 50 parallel
capillaries originating from a single afferent arteriole and
rejoining to form an efferent arteriole (fig 19.19). The
entire structure of the renal corpuscle is related to its
function of filtering blood. The capillary walls are com- afferent efferent
arteriole ~~, arteriole
posed of a single layer of endothelial cells with openings
between them of diameter 50-100 nm. These cells are capillary capillaries
showing of the
pressed up against a basement membrane (basal lamina) fenestrations ; glomerulus
which completely envelops each capillary and forms the basement LY,
only continuous structure separating the blood in the membrane e iY D redcsie
capillary from the lumen of Bowman’s capsule (fig 19.20). podocyte a showing
showing AY primary
The inner layer of Bowman’s capsule is composed of cells secondary processes
called podocytes (podos, foot; cytos, cell) and these processes
resemble starfish in having arms (primary processes) which
give off structures resembling tube-feet called secondary or slit-pore squamous
epithelial
foot processes. The foot processes support the basement cells of
membrane and capillary beneath it and gaps of 25 nm Bowman’s
between the foot processes, called slit-pores, facilitate the capsule
process of filtration. The outer cells of Bowman’s capsule

are unspecialised squamous epithelial cells (figs 19.21 and cells of
proximal
19222); convoluted
Ultrafiltration is the removal from the blood of all tubule
substances with a molecular mass (RFM) less than 68 000

and the formation of a fluid called glomerular filtrate. Fig 19.20 Diagram of a renal corpuscle showing typical cells
Both kidneys receive a total of about 1 200 cm? of of the glomerulus and Bowman's capsule
v3
i Rtg
rte 2:
Pee a
e aluriey
foot processes
P a
Fig 19.21 (a) Electronmicrograph showing the structure of the glomerular capillary and wall of Bowman’s capsule. Lumen of
capillary (C) contains plasma which passes through fenestrations in the capillary wall as indicated by the arrows. (B) is the
basement membrane which acts as the filter between the capillary (C) and the cavity of Bowman's capsule (U). Foot processes
(FP) of a podocyte (P) rest on the basement membrane and slit-pores (SP) can be seen between the foot processes.
(b) Scanning electron micrograph of glomerular capillary
Fig 19.22 Diagram showing the filtration path between the blood per minute which means that all the blood in the
plasma in a glomerular capillary and the filtrate within the circulatory system passes through the kidney every 4-5
lumen of Bowman's capsule minutes. This volume of blood contains 700 cm? of plasma
capsule basement membrane capillary of which 125 cm* per minute is filtered out in the renal
corpuscle. The substances forced out of the glomerular
;
capillaries pass through the capillary fenestrations, base-
ment membrane and slit-pores by the pressure developed
secondary endothelial
within the capillaries. Most of the pressure is derived from
process of cell of the hydrostatic pressure of the circulating blood, but the
podocyte capillary
effective pressure forcing fluid out of the glomerulus, the
filtration pressure, is the result of a balance of other
pressures shown in fig 19.23 and summarised by the
expression:
slit-pore
filtration__hydrostatic glomerular colloid osmotic , hydrostatic glomerular
pressure blood pressure ~ \ pressure filtrate pressure
filtration
pathways
The blood pressure in the glomerular capillary can be
FILTRATE PLASMA varied by changes in the diameter of the afferent and
efferent arterioles which are under nervous and hormonal
control. Constriction of the efferent arteriole decreases the
blood flow out of the glomerulus, raises the hydrostatic
pressure within the glomerulus and, if the condition
persists, it leads to substances with molecular mas-
ses (RFM) greater than about 68000 entering the glom-
erular filtrate.
Glomerular filtrate has a chemical composition similar to
714
that of blood plasma. It contains glucose, amino acids,
The kidney tubules and selective
vitamins, some hormones, urea, uric acid, creatinine, ions
reabsorption and secretion
and water. White and red blood corpuscles, platelets and
Ultrafiltration produces a steady supply of glomerular
plasma protein molecules, such as albumins and globulins,
filtrate, about 125 cm° per minute in Man. If this was
are prevented from passing out of the glomerulus by the
removed as urine it would produce 180 dm’ per day. Since
basement membrane which acts as afilter. Blood passing
only 1 500 cm? of urine is produced each day, a great deal
from the glomerulus has an increased colloid osmotic
pressure due to the increased concentration of plasma
of reabsorption must occur. In fact, of the 125 cm’ of
filtrate produced per minute 124 cm’ is reabsorbed. The
proteins and a reduced hydrostatic pressure.
main regions of reabsorption are shown in fig 19.24.
Ultrafiltration is an entirely passive and indiscriminate
(in terms of substances which are valuable) process.
—- efferent Substances which are useful and vital are removed from the
arteriole
afferent —-> plasma along with excretory and other ‘waste’ substances.
arteriole The function of the kidney tubules is to selectively reabsorb
substances of further use to the body and those required to
maintain the composition of the body fluids at a steady rate.
Further waste substances may be added to the tubules by
active secretion from the blood capillaries surrounding the
colloidal osmotic tubules.
pressure 30 mm Hg
lomerular ; Proximal convoluted tubule. The proximal

Sa hydrostatic blood
network PRES SUE
convoluted tubule is the longest (14mm) and widest
70 mm Hg (60 um) part of the nephron and conveys filtrate from
Bowman’s glomerular Bowman’s capsule to the loop of Henle. It is composed of a
capsule filtrate
pressure single layer of epithelial cells with extensive microvilli
20 mm Hg about 1 um in length forming a brush border on the surface
lining the tubule. The outer membrane of the epithelial cell
rests on a basement membrane and is invaginated to form a
filtration pressure=70-(30+20)
=20 mm Hg labyrinth of basal channels. Adjacent membranes of tubule
cells are separated by intercellular spaces and fluid
circulates through basal channels and intercellular spaces
Fig 19.23 The direction and magnitude of pressures as shown in fig 19.25. This fluid bathes and links the cells of
influencing the filtration pressure in the human glomerulus the proximal convoluted tubule and the surrounding
network of peritubular capillaries. The proximal convo-
luted tubule cells have numerous mitochondria concen-
trated near the basement membrane where they provide
ATP for membrane-bound carrier molecules involved in
active transport.
Selective reabsorption. The large surface

area, the numerous mitochondria of the proximal convo-
, luted tubule cells and the proximity of the peritubular
100 on
capillaries are adaptations for the reabsorption of sub-
stances from the glomerular filtrate. Over 80% of the
glomerular filtrate is reabsorbed here, including all the
Eu cm?
glucose, amino acids, vitamins, hormones and 85% of the
sodium chloride and water. The mechanism of reabsorp-
tion occurs as follows. (1) Glucose, amino acids and ions
that have diffused into the cells of the proximal convoluted
tubule from the filtrate are actively transported out of the
cells and into the intercellular spaces and basal channels by
carrier mechanisms in the cell membranes. (2) Once in
these spaces and channels they enter the extremely
permeable peritubular capillaries by diffusion and are
urine 1 cm? carried away from the nephron. (3) The constant removal
of these substances from the proximal convoluted tubule
Fig 19.24 Diagram showing the principal regions of
reabsorption of renal filtrate cells creates a diffusion gradient between the filtrate in the
WAS
filtrate solutes, such as glucose, from the
Fig 19.25 Diagram showing the suggested mechanism of reabso. rption of glomerular
The carrier molecules of active transport
lumen of the proximal convoluted tubule into the surrounding peritubular capillaries.
apply to the description of glucose uptake
mechanisms are thought to be situated in the basement membrane. The numbers
given in the text
lumen
epithelial cell of proximal
convoluted tubule intercellular space
peritubular capillary
: ili
Gy B microvilli
glucose
basal channel z5
filtrate
Go)
eke - ena i ©) g lucose > @)
glucose & Uw)
glucose
_____ tight junction

basement membrane
with basal infoldings UU) (Cuan)
diffusion ———»> glucose

pathway
active hema
transport
lumen and the cells, down which further substances pass. Table 19.7. The composition of plasma and urine and changes
Once inside the cells they are actively transported into the in concentration occurring during urine formation in Man
spaces and channels and the cycle continues. ee eee
The active uptake of sodium and accompanying anions Plasma % Urine % Increase
POL Pan ARIE, Se Rr
reduces the osmotic pressure in the tubular filtrate and an —
water 90 95
equivalent amount of water passes into the peritubular 8 0 _—
protein
capillaries by osmosis. Most of the filtered solutes and glucose 0.1 0 —
water are removed from the filtrate at a fairly constant rate. urea 0.03 Z 67x
This produces a filtrate in the tubule which is isotonic with uric acid 0.004 0.05 12x
creatinine 0.001 0.075 75X
blood plasma in the peritubular capillaries.
Na® O32 0.35 1x
About 50% of the urea from the filtrate is reabsorbed, by 0.0001 0.04 400 x
NH,"
diffusion, into the peritubular capillaries and passes back Ka 0.02 0.15 7x
into the general circulation whilst the remainder is excreted Mg” 0.0025 0.01 4x
in the urine. Gils 0.37 0.60 2x
Small molecular mass (RFM) proteins, that is those less PO,” 0.009 0.27 30x
than 68 000, passing into the tubule during ultrafiltration
SO} 0.002 0.18 90x
are removed by pinocytosis at the base of the microvilli.
They are enclosed in pinocytotic vacuoles to which primary
lysosomes (section 7.2.8) are attached. Hydrolytic en- The ability to produce a hypertonic urine is found only in
zymes in the lysosomes digest the proteins to amino acids the two groups of vertebrates, the birds and mammals,
which are either utilised by the tubule cells or passed on, by which possess a loop of Henle. The concentration of urine
diffusion, to the peritubular capillaries. produced is directly related to the length of the loop of
Finally, active secretion of creatinine and ‘foreign’ Henle and the thickness of the medulla relative to the
substances occurs in this region. These substances are cortex, which both increase progressively in animals living
transported from the interstitial fluid bathing the tubules in drier habitats. For example, the beaver (Castor), a
into the tubular filtrate and eventually removed in the semi-aquatic mammal, has a thin medulla, a short loop of
urine. Henle and produces a large volume of dilute urine
(600 mOsm kg™' of water), whereas the desert-dwelling
19.5.7 Urine formation
kangaroo rat (Dipodomys) and the jerboa (hopping
Urine is formed by the exchange of solutes and mouse — Dipus) have thick medullas, long loops of Henle
water between the filtrate leaving the proximal convoluted and produce small volumes of highly concentrated urine
tubule and all the structures distal to it (table 19.7). (6 000 and 9 000 mOsm kg" of water respectively).
716
Descending and ascending limbs of the convoluted tubule to the longer, thin (12 um) descending
loop of Henle limb which is freely permeable to water. (2) Since the
Before describing the structure and detailed function of the interstitial region of the medulla is hypertonic due to high
various regions of the loop of Henle it is necessary to concentrations of sodium chloride and urea, water is drawn
describe its overall function. The loop of Henle, in out of the filtrate by osmosis and carried away by the vasa
conjunction with the capillaries of the vasa recta and recta. (3) This reduces the volume of the filtrate by 5% and
collecting duct, creates and maintains an increasing makes it hypertonic. At the apex of the medulla (the
Osmotic gradient in the medulla from the cortex to the papilla) the descending limb of the loop of Henle makes a
papilla due to the build-up of increasing concentrations of hairpin turn and becomes the ascending limb which is
sodium chloride and urea. This, in turn, provides the impermeable to water along its entire length. (4) The first
conditions for the progressive removal of water, by part of the ascending limb, the thin ascending limb, is
osmosis, from the tubule into the interstitial region of the permeable to sodium chloride and urea. Sodium chloride
medulla. The reabsorbed water is then removed from the diffuses out of the limb as it passes away from the
medulla by the blood vessels of the vasa recta. Consequent- hypertonic region of the medulla towards the cortex
ly a hypertonic urine is produced in the collecting duct. whereas urea diffuses into the limb. (5) The epithelium of
At present the precise mechanism by which the loop of the thick ascending limb is composed of flattened cuboidal
Henle concentrates sodium chloride towards the papilla cells with a rudimentary brush border and many mitochon-
of the medulla is not completely understood. The extent of dria. These cells are the site of the active transport of
the permeability of the descending tubule to sodium sodium and chloride ions out of the filtrate. (6) The loss of
chloride is not yet understood, and whilst this affects the sodium and chloride ions from the filtrate increases the
precise details of the mechanism it does not alter what has hypertonicity of the medulla and results in a hypotonic
been described above, or what follows. The movement of filtrate passing to the distal convoluted tubule.
ions, urea and water between the loop of Henle, vasa recta The loop of Henle as a countercurrent
and collecting duct is described by the following points multiplier
which refer to fig 19.26. (1) The first part of the descending
The loop of Henle functions as a countercurrent multiplier
limb is short, relatively wide (30 um) and impermeable to
for the following reasons:
ions, urea and water. It conveys filtrate from the proximal
(1) the close proximity of the descending and ascending
limbs,
Fig 19.26 Summary diagram showing the movement of ions, (2) the permeability of the descending limb to water,
urea and water in the medulla that produce an increasing (3) the impermeability of the descending limb to solutes,
osmotic gradient within the medulla. The numbers in boxes (4) the permeability of the thin ascending limb to solutes,
are the osmotic concentrations in mOsm kg water. The
numbers in circles refer to details in the text. The tonicity and
shown in various regions of the nephron is relative to blood (5S) the active transport mechanisms of the thick ascending
plasma outside the kidney limb.
descending ascending collecting 300
limb limb 4 duct
isotonic (6) hypotonic

300 100
vasa © o>
k+x.0
NaCl => urea
recta (> =p urea x
eS increasing
osmotic
H,O pressure in
interstitial
H,O fluid
H,O
H,O
Ue TS ee ee ee een ae urea
Ss ; a
> 1200
aia seta ie
<+— movement of water ! G) hypertonic
<== active transport of NaCl a it me
On passive movement of Na™ and Cl Ne Nae = a
Gace
geass movement of urea a 1200
a a
717
The values for osmotic concentrations of the filtrate in both The vasa recta as a countercurrent
exchanger
limbs also suggest that the loop of Henle functions as a
countercurrent multiplier. In such a system the concentra- The narrow descending capillary and wider ascending
tion difference at any given level between the fluids in the capillary of the vasa recta run parallel to each other and
ascending and descending limbs may be slight, but as this is give off branched loops at different levels along their
maintained over a long distance, as in the loop of Henle, length. They are situated close to the limbs of the loop of
the difference becomes additive so that the final concentra- Henle but substances are not transferred directly from
tion of filtrate at the base of the loop is much greater than at limbs to blood vessels. Instead they pass through the
either end. The longer the loop the greater the concentra- interstitial region of the medulla where, due to the slow
tion difference. rate of blood flow through the vasa recta, sodium chloride
The removal of sodium chloride from the ascending limb and urea are trapped and an osmotic gradient is main-
increases the concentration of the interstitial region of the tained. The cells of the vasa recta are freely permeable to
medulla and water is drawn out of the descending limb. ions, urea and water, and as the vessels run side by side
This water immediately passes into the vasa recta, thereby they function as a countercurrent exchange system. As the
maintaining high concentrations of solutes in the interstitial descending capillary enters the medulla the progressively
region and in the filtrate, as shown in fig 19.27. Filtrate increasing hypertonic condition of the interstitial region
flows around the loop and countercurrent exchange draws water out of the plasma in the capillary by osmosis,
continues. The constant cycling of solutes, both passively and sodium chloride and urea enter by diffusion (fig 19.28).
and actively, ensures a permanently high concentration of As blood flows from the medulla in the ascending capillary
sodium chloride in the medulla. At the same time another the reverse happens and the decreasing osmotic pressure of
system, a countercurrent exchange system (see below), the interstitial region enables water to re-enter the plasma
operates in the vasa recta, and these two countercurrent whilst sodium chloride and urea leave. The adaptive
mechanisms operate together to ensure the gradient of significance of this mechanism is that it allows the osmotic
hypertonicity in the medulla. concentration of plasma Jeaving the kidney to remain at a
steady state irrespective of the osmotic concentration of
Fig 19.27 Model illustrating the countercurrent multiplier plasma entering the kidney. Finally, and most importantly,
formed by the tubules of the loop of Henle. Na* and Cl” ions since all the movements of solutes and water are passive,
are actively transported out of the thick ascending limb into
the vasa recta countercurrent exchanger makes no metabo-
the interstitial region. The solute concentration of the
interstitial region rises. Water leaves the descending limb. The lic demand on the kidney.
numbers indicate the osmotic concentration in mOsm kg~'
water. An arbitrary concentration gradient of 200 mOsm ko™*
water exists between descending and ascending limbs in the
model
descending limb ascending limb EG
O
300 R
o
E
xX
H,O
H,O
increasing
osmotic , M
concentration E
H,O
D
U
L
L
A
H,O
interstitial region interstitial region
active transport
of NaCl H,0
<¢-—-—— passive movement
of NaCl
<—— movement of v
water 1 200
718
Fig 19.28 ts The passive movement of water, ; ions and urea between adjyacent vessels of the vasa recta
and between these and
the interstitial fluid of the medulla. The vessels of the vasa recta form a countercurrent exchange system.
All values are given
in mOsm kg~' water
c
O
R 300 |
I
ail
\
|
x 350
300 300
350 water 400
sodium chloride
400 -—----pPeP #—-—————_———___ —- —-—+——-—-—»p 400
ee pe urea
=== === = =
500
M
E 600 600
D
U
Ie
13 700
A
300 > 800

900
1000 1000
—» movement ofwater = ——————— 1 100-q- — — —— — — — — — >

—— —p movement of sodium chloride =======>==
=—=—-=> movement of urea
1200 1 200 1200
Distal convoluted tubule

The distal convoluted tubule loops backward towards the 19.6 = Anti-diuretic hormone (ADH) and
renal corpuscle and passes through the cortex. The cells of the formation of a hypertonic or
the distal tubule have brush borders and abundant hypotonic urine
mitochondria and this region is the site of the mechanisms
The body maintains the osmotic pressure of
for the fine control of salt, water and pH balance of the
the blood at an approximately steady state by balancing
blood. The hormonal control of the permeability of the
water uptake from the diet with water lost in evaporation,
distal convoluted tubule to water is linked to that of the
sweating, egestion and urine. The precise control of
collecting duct; the two are described later. The control of
osmotic pressure, however, is achieved primarily by the
salt and pH balance are described in section 19.7.
effect of ADH on the permeability of the distal convoluted
Collecting duct tubule and collecting duct.
The collecting tubule passes from the distal convoluted When small amounts of water are ingested, or excessive
tubule in the cortex down through the medulla where it sweating occurs, or large amounts of salt are ingested,
joins up with several other ducts to form larger ducts, the osmoreceptors in the hypothalamus detect a rise in blood
ducts of Bellini. The permeability of the walls of the duct to osmotic pressure. Nerve impulses are set up and pass to the
water and urea is controlled by anti-diuretic hormone posterior pituitary gland where ADH (vasopressin) is
(ADH), and together with the distal convoluted tubule released. ADH increases the permeabilities of the distal
convoluted tubule and collecting duct, water is withdrawn
these structures can produce a hypertonic or hypotonic
from the filtrate into the cortex and medulla and a reduced
urine according to the body’s demand for water.
wA9
Lp Nat and CI ions ADH absent
ADH present
EG ——» water movement
O
R ae = | ====> urea movement
i 300
E ---— ~~. |
|
* Se 3008
| [
| |
| |
ont 400 eae
| ae
ea GOO |
<< aa |
|
M |
E 22ee= 500mm |
U | |
Ik | |
1 } 1000 |!
qtr om |
| |
| |
| |
; 1200 |
(a) |
volume of hypertonic urine (1 000 mOsm kg’ of water) is Fig 19.29 Diagram illustrating the effect of ADH on the
released from the kidney. permeabilities of the distal convoluted tubule and collecting
duct to water and urea
ADH also increases the permeability of the collecting
duct to urea, which diffuses out of the urine into the
interstitial region of the medulla. Here it increases the
osmotic concentration resulting in the removal of an
increased volume of water from the thin descending limb
(fig 19.29). 19.7 Control of blood sodium level
When there is a high intake of water and the osmotic
pressure of the blood begins to fall ADH release is The maintenance of the plasma sodium level
inhibited, the walls of the distal convoluted tubule and at a steady state is controlled by the hormone aldosterone
collecting duct become impermeable to water, less water is which secondarily influences water reabsorption. A de-
reabsorbed as the filtrate passes through the medulla and a crease in blood volume stemming from a loss of sodium
large volume of hypotonic urine is excreted (fig 19.29). stimulates a group of secretory cells, the juxtaglomerular
Table 19.8 shows a summary of the events involved in complex, situated between the distal convoluted tubule and
regulating water balance, and the control mechanisms the afferent arteriole to release an enzyme, renin. Renin
involved in regulating water and salt balance are shown in activates a plasma globulin, produced in the liver, to form
fig 19.30. the active hormone angiotensin and this releases aldoster-
Failure to release sufficient ADH leads to a condition one from the adrenal cortex. This stimulates the active
known as diabetes insipidus in which large quantities of uptake of sodium from the filtrate into the plasma in the
hypotonic urine are produced (diuresis). The fluid lost in peritubular capillaries. This uptake for sodium ions causes
the urine has to be replaced by excessive drinking. the reabsorption of an osmotically equivalent amount of
water.
Table 19.8. Summary of the changes produced in the epi-
thelium of the distal convoluted tubule and collecting duct 19.7.1 Homeostatic control of plasma,
in response to ADH water and sodium levels
When the body has free access to water and a
Blood osmotic normal sodium diet, no ADH or aldosterone is released
pressure ADH Epithelium Urine and the epithelium of the distal convoluted tubule and
rises released permeable concentrated collecting duct remains impermeable to ions, urea and
water and the urine is copious and dilute, a condition
falls not released impermeable dilute known as diuresis. The homeostatic control mechanisms
are summarised in fig 19.30.
720
KIDNEY
permeability of distal convoluted

decrease in ADH output tubule and collecting duct
(effector) decreases (effector)
osmoreceptors in hypothalamus
(detector) ~
oe copious dilute urine produced

— output
fall in plasma Meg oe:
osmotic pressure Cas =
increased water Gack ~ =
uptake (input) a ~,
= ~ .
normal plasma osmotic level ~w~_] normal plasma osmotic level
(set-point) ioe _ (set-point)
- 1
decreased water
uptake (input) e Sa
rise in plasma - pack
osmotic pressure ie fee
_— neg" reduced volume of
=e concentrated urine produced
ed (output)
osmoreceptors in hypothalamus 4. —~
(detector)
KIDNEY
permeability of distal convoluted
increase in ADH output tubule and collecting duct
(effector)
increases (effector)
Fig 19.30 Summary diagram of the control of plasma osmotic pressure
19.8 Control of blood pH the plasma pH from its normal level of 7.4, however, are
counteracted by the distal convoluted tubule. This excretes
Hydrogencarbonate and phosphate buffers in hydrogen ions and retains hydrogencarbonate ions if the
the blood prevent excess hydrogen ions (H"), produced by pH falls, and excretes bicarbonate ions and retains hydro-
metabolic activities, from decreasing the pH of the blood. gen ions if the pH rises. This may produce changes in the
Carbon dioxide released into the blood during respiration pH of the urine from 4.5 to 8.5. A fall in pH also stimulates
is regulated by this system and prevented from causing the kidney cells to produce the base ion ammonia (NH,")
changes in plasma pH prior to its excretion from the lungs. which combines with acids brought to the kidney and is
Excessive changes in blood chemistry which would change then excreted as ammonium salts.
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Chapter Twenty
a
Reproduction
The ability to reproduce, that is to produce a new

20.1.1 Asexual reproduction
generation of individuals of the same species, is one of the
fundamental characteristics of living organisms. It involves Asexual reproduction is the production of
the transmission of genetic material from the parental offspring from a single organism without the fusion of
generation to the next generation, thereby ensuring that gametes. Except in plants with alternation of generations
the characteristics, not only of the species but also of the (section 3.3.1), meiosis is not involved and the offspring are
parental organisms, are perpetuated. The significance of identical to the parent. Identical offspring from a single
reproduction in a given species is to replace those members parent are referred to as a clone. Members of a clone only
of the species that die, thus ensuring continuity of the differ genetically as a result of random mutation. Higher
species, and also to allow increase in total numbers of the animals do not naturally reproduce asexually, though
species where conditions are suitable. recent successful attempts, discussed later, have been
A new individual normally has to go through a period of made to clone certain species artificially.
growth and development before it reaches the stage at There are several types of asexual reproduction. Further
which it can reproduce itself (chapter 21). Some members details of the process in some of the individuals referred to
of a species will die before they reach reproductive age, due below are given in the relevant sections of chapters 2, 3
to predation, disease and accidental death, so that a species and 4.
will only survive if each generation produces more Fission
offspring than the parental generation. Population sizes Fission occurs in unicellular organisms and is the division of
will fluctuate according to the balance between rate of the cell into two or more daughter cells identical to the
reproduction and rate of death of individuals (section parent cell. Replication of DNA and, in the case of
12.6). There are a number of different reproductive eukaryotes, nuclear division precedes cell division. Norm-
strategies, all with certain advantages and disadvantages ally two identical daughter cells are produced, a process
which are described in this chapter. called binary fission. This occurs in bacteria, many
protozoans, such as Amoeba and Paramecium, and in some
unicellular algae, for example Euglena. Under suitable
conditions it results in rapid population growth, as
described in section 2.2.4 for bacteria.
20.1 Asexual and sexual reproduction Multiple fission, in which repeated divisions of the parent
nucleus are followed by division into many daughter cells,
There are two basic types of reproduction, occurs in the Sporozoa, a group of protozoans including the
asexual and sexual. Asexual reproduction is reproduction malaria parasite Plasmodium. The stage undergoing
by a single organism without production of gametes. It multiple fission is called the schizont and the splitting
usually results in the production of identical offspring, the process schizogony. In Plasmodium schizogony occurs
only genetic variation arising as a result of random immediately after infection when the parasite enters the
mutations among the individuals. liver. About 1000 daughter cells are released, each
Genetic variation is advantageous to a species because it capable of invading a red blood cell and producing up to a
provides the ‘raw material’ for natural selection, and hence further 24 daughter cells by schizogony. Such prolific
evolution. Offspring showing most adaptations to the powers of reproduction compensate for the large losses
environment will have a competitive advantage over the associated with the difficulties of successful transfer
other members of the species and be more likely to survive between hosts, namely Man and the vector organism, the
and pass on their genes to the next generation. The species mosquito.
therefore has the capacity to change, that is to undergo
Spore formation (sporulation)
speciation (section 25.7). Increased variation can be
achieved by the mixing of genes from two different A spore is a small reproductive unit which is usually
individuals, a process known as genetic recombination. microscopic and unicellular, containing a small amount of
This is the essential feature of sexual reproduction and cytoplasm and a nucleus. Spores are produced by bacteria,
occurs in a primitive form in some bacteria (section 2.2.4). protozoans, all groups of green plants and all groups of
R23
parent
fungi. They vary in type and function and are often Fig 20.1 (above) Budding of Hydra, a coelenterate
produced in special structures. For example, spores of
Rhizopus and Dryopteris are produced in sporangia; the
microspores (pollen) and megaspores (embryo sacs) of Fig 20.2 (below) Asexual reproduction in Bryophyllum.
seed-bearing plants are produced in sporangia called pollen Young plantlets are seen along the leaf margins
sacs and ovules respectively.
Often spores are produced in large numbers and are very
light, being dispersed easily by air currents as well as by
animals, particularly insects. Being small they usually have
minimal food stores and there is great wastage as many fail
to find a suitable place for germination. In these cases they
are primarily a means of rapid multiplication and spread of
species, particularly of fungi.
Strictly speaking, the spores of bacteria are not repro-
ductive structures but means of surviving adverse condi-
tions, since each bacterium produces only one spore.
Bacterial spores are among the most resistant known. For
instance, they will often survive strong disinfectants and
boiling in water.
Although this section is concerned with asexual repro-
duction, it should be pointed out that some spores are
sexual spores, that is they take part in sexual reproduction.
Examples are the zoospores of Chlamydomonas which
sometimes function as gametes, and the zygospores of
Rhizopus and Spirogyra. Zygospores are products of
sexual reproduction and are relatively large spores with
food stores and a protective outer coat. They can remain
dormant during periods of adverse conditions.
724
Note that one organism may produce more than one type Perennating organs may be stems, roots, or whole shoots
of spore, for example Rhizopus produces sexual and (buds), but in all cases the food stored is derived mainly
asexual spores, and higher plants produce micro- and from photosynthesis of the current year’s foliage leaves.
megaspores in an asexual process. The food is translocated to the storage organ and then
usually converted to an insoluble storage product such as
Budding
starch. During the period of adverse conditions the aerial
Budding is a form of asexual reproduction in which a new parts of the plant die and the perennating organ undergoes
individual is produced as an outgrowth (bud) of the parent, a period of dormancy underground. At the beginning of the
and is later released asa self-supporting, identical copy of next growing season the food reserves are mobilised by
the parent. It takes place in a number of groups of enzymes and buds become active, growing at the expense
organisms, notably the coelenterates, for example Hydra of the stored food. If more than one bud grows reproduc-
(fig 20.1), and unicellular fungi, such as yeasts. In the latter tion has taken place. This sequence of events is synchro-
case budding differs from fission, which also occurs, in that nised closely with the seasons, being controlled by such
the two parts produced are not of equal size. factors as day length (photoperiod) and temperature,
An unusual form of budding is shown by the succulent environmental variables whose profound effects on growth
plant Bryophyllum, a xerophyte commonly used as a and development are described in chapter 15.
decorative houseplant. It forms miniature plants, complete Some of the organs of vegetative propagation and
with small roots, along the margins of its leaves (fig 20.2). perennation are described below.
These ‘buds’ eventually fall off and establish themselves as
independent plants. Bulb. A modified shoot, for example onion
(Allium), daffodil (Narcissus) and tulip (Tulipa). An organ
Fragmentation of perennation as well as vegetative propagation.
Fragmentation is the breaking of an organism into two or A bulb has a very short stem and fleshy storage leaves. It
more parts, each of which grows to form a new individual. is surrounded by brown scaly leaves, the remains of
Fragmentation occurs in filamentous algae, such as previous year’s leaves after their food stores have been
Spirogyra. Here breakage into two may occur anywhere used. The bulb contains one or more buds. If more than
one grows, each forms a shoot which produces a new bulb
along the length of the filament.
Fragmentation also occurs among certain lower animals at the end of the growing season, this being vegetative
which, unlike more advanced animals, retain strong propagation. Roots are adventitious, that is they grow from
the stem with no tap root.
powers of regeneration from relatively undifferentiated
A typical bulb is illustrated in fig 20.3.
cells. For example, the bodies of ribbon worms, a group of
primitive, mainly marine worms, break up particularly
easily into many pieces, each of which can regenerate anew brown scale leaf, protective, foliage leaf of
individual. While in this case the process is a natural and season before last
controlled one, there are examples of animals that fleshy leaves storing food, swollen bases of
last season’s foliage leaves
normally only regenerate as a result of accidental frag- apical bud next season’s plant, contains
mentation, such as starfish. These animals have been the embryo stem, foliage leaves and a flower
subjects of experiments on regeneration, a commonly used apical bud of next season’s bulb
example being the free-living flatworm Planaria. These axillary bud, may also form a new plant
experiments have contributed to our understanding of the (compare apical bud)
short, flattened stem
process of differentiation (section 22.8).
remains oflast season’s adventitious roots
Vegetative propagation (vegetative
reproduction) Fig 20.3 Diagrammatic section through a dormant bulb
Vegetative propagation or vegetative reproduction is a
form of asexual reproduction in which arelatively large,
usually differentiated part of a plant body becomes Corm. A short, swollen, vertical under-
detached and develops into an independent plant. In ground stem, as in Crocus and Gladiolus. An organ of
essence it is similar to budding. Specialised structures often perennation as well as vegetative propagation.
develop for the purpose, including bulbs, corms, rhizomes, A corm consists of the swollen base of a stem surrounded
stolons and tubers. Some of these also store food and are by protective scale leaves; there are no fleshy leaves, unlike
means of surviving adverse conditions, such as cold periods bulbs. Scale leaves are the remains of the previous season’s
or drought. Plants possessing them can therefore survive foliage leaves. Roots are adventitious. At the end of the
from one year to the next and are either biennial, flowering growing season contractile roots pull the new corm down
and dying in their second year, or perennial, surviving from into the soil. The corm contains one or more buds which
year to year. The structures are called perennating organs, may result in vegetative propagation (compare bulb).
and include bulbs, corms, rhizomes and tubers. A typical corm is illustrated in fig 20.4.
Ted
apical bud, next season’s plant, contains parent plant stolon, shoot arches over
embryo stem, foliage leaves and a flower and roots from a node,
the bud at this node
next season’s stem and corm grows into a new shoot
S@—oniillary bud, may also form a new plant leaf
(compare apical bud) apical bud
brown scale leaves, protective, shrivelled axillary new shoot
remains of last season’s foliage leaves bud
corm, formed by swelling of stem base with
food at end of growing season adventitious roots
old corm of last year
adventitious roots, some are contractile and
\ pull new corm down into soil
Fig 20.6 (above) Generalised plan of a stolon

Fig 20.4 Diagrammatic section through a dormant corm
Fig 20.7 (below) Plan of a strawberry runner
Rhizome. A horizontally growing under- parent plant runner growing
ground stem, such as in Iris, Solomon’s seal (Poly- new plants growing
from axillary buds
from axillary bud
in axis of scale leaf
gonatum), where it is short and swollen with stored food,
and couch grass (Agropyron repens), mint (Mentha) and apical bud
Michaelmas daisy (Aster spp.) where it is long and thin. It is

usually an organ of perennation as well as vegetative
propagation.
A rhizome bears leaves, buds, and adventitious roots.
The leaves may be scale-like (small and thin, whitish or lished. The runner may represent the main stem or grow
brownish in colour) as in couch grass, green foliage leaves from one of the lower axillary buds on the main stem, as
only as in Jris, or a mixture of green and scale leaves as in illustrated in fig 20.7. In strawberry, scale leaves and
Solomon’s seal, where both types are borne on aerial axillary buds occur at every node, but roots and foliage
shoots. An Iris rhizome is illustrated in fig 20.5. leaves arise only at every other node. All axillary buds may
Stolon. A creeping, horizontally growing give rise to new runners.
stem that grows along the surface of the ground, for Tuber. A tuber is an underground storage
example blackberry (Rubus), gooseberry, blackcurrant organ, swollen with food and capable of perennation.
and redcurrant (all Ribes spp.). It is not an organ of Tubers survive only one year and shrivel as their contents
perennation. (See also runner below.) Roots are adven- are used during the growing season. New tubers are made
titious, growing from nodes. at the end of the growing season, but do not arise from old
A plan ofa typical stolon is illustrated
in fig 20.6. tubers (in contrast to corms, which arise from old corms).
Runner. A type of stolon that elongates Stem tubers are stem structures produced at the tips of
rapidly, as in strawberry (Fragaria) and creeping buttercup thin rhizomes, as in potato (Solanum tuberosum) and
(Ranunculus repens). artichoke (Helianthus tuberosum). Their stem structure is
A runner bears scale leaves with axillary buds and the revealed by the presence of axillary buds in the axils of
buds give rise to adventitious roots and new plants. The scale leaves. Each bud may grow into a new plant during
runners eventually decay once the new plants are estab- the next growing season.
remains of vascular bundles flower stalk

from apical bud which turns up
circular leaf scars at nodes remains of leaves foliage leaves
after flowering, food from the leaves passes back to the
rhizome and enters an axillary bud which continues
growth ofthe rhizome
old part of stem does not
die away as quickly as in ast year’s leaves withering, brown and scaly
bulbs and corms, and lasts
for several years
H eee of concealed axillary bud; it becomes swollen
adventitious roots grow
mainly from nodes
new rhizome branch
A with food from the leaves and continues growth of the
rhizome; at the end of the new growth an apical bud
will develop; at the beginning of the next season this
grows and produces leaves and flowers at the expense
growing from an axillary ofthe food store
bud, therefore vegetative
propagation
Fig 20.5 Diagrammatic structure of an \ris rhizome
726
scale leaf scar : aS aT ery
‘eye’, spiral arrangement of
axillary bud eyes around tuber;
each bud can give rise NGGeti: tuber swollen adventitious
to anew shoot root; stores inulin, a polymer of
in the next growing fructose
season
: me lenticels in outer corky layer
pes
= apical bud
rhizome of this year’s plant, bears Fig 20.9 (above) Root tubers of Dahlia
stem tuber, stores starch
small scale leaves with axillary
buds; it leaves a scar on the tuber
when the latter becomes detached
at the end of the growing season
Fig 20.8 (above) Stem tuber of potato Fig 20.10 (below) Tap roots of carrot and turnip
buds (axillary and apical)
oS
cortex stele parenchymatous leaf bases of current
cells swollen with food season’s shoot; aerial
parts die at end of
lateral root growing season
swollen tap root swollen
with food produced by
vascular tissue of foliage leaves of current
lateral root : season
vascular tissue of tap
root
small lateral roots
normal part of roots
Carrot LS Carrot entire Turnip entire
Root tubers are swollen adventitious roots, for example Cuttings are parts of the plant which, when removed and
Dahlia and lesser celandine (Ranunculus ficaria). New given suitable conditions, develop roots and leaves and
plants develop from axillary buds at the base of the old become independent plants. In this way favoured varieties
stem. can be artificially propagated without change. A rooting
A stem tuber of potato is illustrated in fig 20.8 and root hormone is often added to stimulate rooting. The shoots of
tubers of Dahlia in fig 20.9. Pelargonium and Coleus, twigs of willow (Salix spp.) and
Forsythia and leaves of Begonia and African violet
Swollen tap roots. A tap root is a main root
(Saintpaulia ionantha) can all be used for successful
that has developed from the radicle, the first root of the
cuttings.
seedling. Tap root systems are characteristic of dicotyle-
Another important, commonly used means of artificial
donous plants. Tap roots may become swollen with
propagation is grafting. Grafting in plants means the
parenchymatous food-storing tissue, as in carrot (Daucus),
transplantation of part of one plant (a shoot or bud) on to
parsnip (Pastinaca), swede (Brassica napus), turnip
the lower part of the shoot of another plant. The donor
(Brassica rapa) and radish (Raphanus sativus). Together
plant is called the scion and the recipient the stock. The
with buds at the base of the old stem, just above the tap
stock is cut off above the graft. The resulting plant usually
root, they form organs of perennation and vegetative
has the root system of the stock and the shoot system,
propagation. Two types of swollen tap root are shown in
including the flowers and fruit, of the scion. The method is
fig 20.10. commercially important for the propagation of rose bushes
Swollen tap roots are characteristic of biennial plants,
and fruit trees, particularly apple trees. The technique has
plants that grow vegetatively during the first year of growth
the twin advantage of combining advantageous features
and survive winter by means of an underground storage
from two different varieties or species, and allowing rapid
organ. They produce flowers and seeds during the second production of large numbers of the new scion/stock
year of growth, at the end of which they die. combinations for marketing. Occasionally successful grafts
Apart from the specialised organs of vegetative propaga- of more than one scion on to a stock have been marketed,
tion described above, some unmodified plant parts have an example being apple trees that bear both eating and
the ability to regenerate new plants if detached from the cooking apples.
parent plant, such as the fallen leaves of the succulent plant
Sedum. The taking of cuttings by gardeners and horticul-
turalists can be regarded as a form of vegetative propaga-
tion.
tel
Cloning of higher plants and animals Meiosis is an essential feature of life cycles in which
sexual reproduction occurs because it provides a mechan-
As already mentioned, the production of identical ism for reducing the amount of genetic material by half.
offspring by asexual reproduction is called cloning. Since This ensures constancy in the amount of genetic material in
the early 1960s techniques have been developed which
each sexually reproducing generation, since fertilisation
have allowed successful cloning of certain higher plants and
doubles the amount of genetic information. During meiosis
animals. These arose from attempts to demonstrate that
random segregation of chromosomes (independent assort-
the nuclei of mature, non-developing cells still contain all
ment) and exchange of genetic material between homo-
the information required to code for an entire organism
logous chromosomes (crossing-over) results in new com-
and that cells become specialised as a result of the switching
binations of genes being brought together in the gamete
on and off of genes rather than by the loss of certain genes
and this reshuffling increases genetic diversity (section
(section 22.7). The first success was achieved by Professor
22.3). The fusion of haploid gametic nuclei is called
Steward of Cornell University, who showed that individual
fertilisation or syngamy and it results in the production of a
cells from a carrot root (the part we eat), when grown in a
diploid zygote, that is a cell containing a set of chromo-
medium containing suitable nutrients and hormones, could
somes from each parent. This combination of two unique
be induced to start dividing again and to produce new
sets of chromosomes (genetic recombination) in the zygote
carrot plants.
forms the genetic basis of variation within species. The
Success with higher animals followed shortly when
zygote grows and develops into the mature organism of the
Gurdon, working at Oxford University, achieved the first
next generation. Sexual reproduction, therefore, involves
successful cloning of a vertebrate. The process does not
the alternation, within the life cycle, of diploid and haploid
occur naturally among vertebrates but, by taking a cell
phases, and the form adopted by the organism in each of
from the intestine of a frog and introducing its nucleus into
these phases varies according to species, as shown in fig
an egg cell whose own nucleus had been destroyed by
2013200332
ultra-violet radiation, he was able to grow a tadpole, and
Gametes are usually of two types, male and female, but
thence a frog, identical to the parent from which the
in some primitive organisms they are of one type only. (See
nucleus was transplanted (fig 20.11).
isogamy, anisogamy and oogamy in section 20.2). If the
Not only did experiments like these establish that
gametes are of two types, they may be produced by
differentiated (specialised) cells still contain all the in-
separate male and female parents or by a single parent
formation needed to make the whole organism, but they
bearing both male and female reproductive organs. Species
suggested that similar techniques might successfully be
that have separate male and female individuals are
used in cloning more advanced vertebrates, including Man.
The cloning of desirable animals such as prize bulls,
described as unisexual, such as Homo (Man) and most
other animals. Some flowering plants are unisexual and in
racehorses and so on, might be as advantageous as the
cloning of plants which it has been noted already takes the case of monoecious plants there are separate male and
place. However the application of the technique to Man female flowers on the same plant, for example Corylus
would be open to serious moral questions. Theoretically (hazel) and Cucurbita (cucumber), whilst in dioecious
any number of genetically identical copies of the same man plants one plant bears male flowers only and the other
or woman might be made. Although superficially it might female flowers only, as in Ilex (holly) and Taxus (yew).
seem that, for example, brilliant scientists or artists might Hermaphroditism
be perpetuated in this way, it has to be remembered that
the degree to which environment influences development is Species capable of producing both male and female
not fully known, and any cloned cell would have to go gametes within the same organism are described as
through all the phases of development once again includ- hermaphrodite, or bisexual. Many protozoans, for example
ing, in the case of Man, embryo, foetus, baby and Paramecium, coelenterates such as Obelia, platyhelminths
childhood. such as Taenia (tapeworm), oligochaetes such as Lumbri-
cus (earthworm), crustacea such as Balanus (barnacle),
molluscs such as Helix (garden snail), some fish, lizards and
20.1.2 Sexual reproduction birds, and most flowering plants are hermaphrodite. This is
Sexual reproduction is the production of thought to be the most primitive form of sexual reproduc-
offspring by the fusion of the genetic material of haploid tion and is seen in many primitive organisms. It represents
nuclei. Usually these nuclei are contained in specialised sex an adaptation to sessile, slow-moving and parasitic modes
cells, or gametes, and at fertilisation they unite to form a of life. One of the advantages of hermaphroditism is the
diploid zygote which develops into the mature organism. ability to carry out self-fertilisation, an essential for certain
Gametes are haploid, carrying a set of chromosomes endoparasites, such as Taenia, which live a solitary
derived by meiosis, and they provide the link between one existence. However, in the majority of species fertilisation
generation and the next. (Sexual reproduction in flowering of gametes from different organisms occurs, and many
plants involves fusion of nuclei rather than cells but genetic, anatomical and physiological adaptations exist
generally these nuclei are referred to as gametes.) which prevent self-fertilisation and favour cross-
728
wild-type
female
(recipient)
Fig 20.11 A clone of toads (Xenopus laevis) produced by nuclear transplantation. (above) A single tail-bud embryo was
obtained from a cross between two albino mutants (donor parents). Its cell were dissociated and their nuclei transplanted
into u.v.-treated (nucleus destroyed) unfertilised eggs of the wild-type female (recipient) shown.
(below) The group of 30 toads (nuclear-transplant clone) are all female and albino; they were obtained froma total of 54
nuclear transfers
129
fertilisation. For example, self-fertilisation is prevented in into the population. The main advantage of parthenogene-
many protozoans by ‘genetic incompatability’, in many sis in aphids is the speed with which the population can
increase its numbers, since all mature members of the
flowering plants by the structure of the androecium and
gynaecium and in many animals by the production of eggs population are egg-layers. This is particularly important at
and sperm at different times by the same organism. times when the environment is favourable and can support
a large population, as during the summer months.
Parthenogenesis Among plants parthenogenesis occurs widely and takes
Parthenogenesis is a modified form of sexual reproduction various forms. One form is called apomixis, but it is not
in which a female gamete develops into a new individual strictly a form of sexual reproduction, it only mimics its
without being fertilised by a male gamete. The process action. It occurs in some flowering plants in which a diploid
occurs naturally in both animal and plant kingdoms and has cell of the ovule, either from the nucellus or megaspore,
the advantage, in some cases, of accelerating the normal develops into a functional embryo in the absence of a male
reproductive rate. gamete. The rest of the ovule develops into the seed and
There are two forms of parthenogenesis, haploid and the ovary into the fruit. In other forms a pollen grain is
diploid, depending upon the chromosome number of the necessary to trigger parthenogenesis, even though the
female gamete. Many insect species, including ants, bees, pollen grain does not grow. In such cases the pollen grain
and wasps, utilise haploid parthenogenesis for the produc- induces the hormonal changes necessary for embryo
tion of specific types of organism within the social group. In development, and such cases are, in practice, difficult to
these species meiosis occurs and haploid gametes are distinguish from true sexual reproduction.
formed. Some eggs undergo fertilisation and develop
normally into diploid females, whereas unfertilised (hap-
20.1.3 The origins of sexual reproduction
loid) eggs develop into fertile haploid males. For example,
the queen honeybee (Apis mellifera) lays fertilised eggs Any discussion based upon events occurring at
(2n = 32) which develop into females (queens or workers) an unknown stage in the history of life is purely speculative
and unfertilised eggs (n =16) which develop into males and any conclusions which emerge from the discussion
(drones) which produce sperm by mitosis and not meiosis. must be tentative. The origins of sexual reproduction
The details of the development of the three types of represent such a situation. It is thought that asexual
honeybee are summarised in fig 20.12. In social insects this methods of reproduction represent the primitive form of
mechanism has the adaptive significance of controlling the reproduction which existed as a means of ‘copying’
numbers of each type of offspring. generations which were genetically identical. At a later
Aphids show diploid parthenogenesis in which the stage, mechanisms are thought to have arisen enabling
egg-producing cells of the female undergo a modified form nucleic acid to be exchanged between organisms. This may
of meiosis involving total non-disjunction (section 22.3). have involved the fusion of whole organisms followed by
All the chromosomes enter the egg cell and not the polar meiosis and, indeed, this possibility would appear to
bodies. The egg cells develop within the mother and young precede the production and fusion of gametes. It is thought
females are released viviparously, that is the parents do not that the mechanisms involved in the exchange of genetic
lay eggs but bear live young. This can occur for several material as shown by bacteria and described in section 2.2.4
generations, particularly during the summer, until a cell represent a primitive stage in the development of sex. The
undergoes almost complete non-disjunction and produces advantage of phenotypic variation resulting from meiosis
a cell containing all the autosomes plus a single X and genetic recombination doubtless played a major role in
chromosome. This cell develops parthenogenically into a the development of more complex forms of life and types of
male aphid. These autumn stage males and parthenogeni- gametes, ranging from identical gametes (isogametes) to
cally produced females, having undergone meiosis, pro- the heterogametic stage of motile male gametes, re-
duce haploid gametes for sexual reproduction. Fertilisation presented by sperms and antherozoids, and non-motile
occurs and the diploid eggs laid by the females overwinter female gametes, represented by eggs (ova). The fact that
and hatch to produce parthenogenic females in the spring. every major group of organisms carries out sexual
These offspring are viviparous. The parthenogenic genera- reproduction suggests that it has advantages over asexual
tions alternate with a normal sexually reproduced genera- reproduction, the latter though rare in animals above the
tion which introduces genetic recombination and variation level of platyhelminths, is retained in plants. Indeed the
queen
mitosis NY
(fertile female)
(fed on royal jelly)
fertilised egg (fed on honey and pollen)

unfertilised egg drone ————
(2n = 32) (n= 16) parthenogenesis
(fed on honey and pollen)
worker Fig 20.12 Summary diagram showing the role of
(sterile female) parthenogenesis in the life cycle of the honey bee colony
730
Table 20.1. Comparison: of asexual reproduction with majority of plant species show alternation of asexual and
sexual reproduction sexual modes of reproduction in their life cycles, as
—————————
oe described in section 20.2. One reason for this is that sessile
Asexual reproduction Sexual reproduction organisms have problems of exchanging gametes and in
(omitting bacteria) many cases retain the option of reproducing asexually.
a a i
One parent only Likewise the ecological problem of finding favourable
Usually two parents
No gametes are produced
habitats for offspring is aided by parents established in a
Gametes are produced. These
are haploid and nuclei of two
habitat reproducing asexually. Problems of overcrowding,
gametes fuse (fertilisation) to however, work counter to this and help limit the size of
form a diploid zygote plant populations.
Meiosis absent Meiosis present at some stage in A summary of some of the essential and typical features
life cycle to prevent of asexual and sexual reproduction is given in table 20.1.
chromosome doubling in every
generation*
20.1.4 Variety of life cycles
Offspring identical to parent Offspring are not identical to
parents. The various stages of development through
They show genetic variation as a which the members of a species pass from the zygote of one
result of genetic recombination
generation to the zygote of the next are called the life cycle.
Commonly occurs in plants, Occurs in the majority of plant Life cycles vary in complexity and in some cases involve
lower animals and micro- and animal species
organisms two or more generations that differ in their morphology
Absent in higher animals (appearance) and reproduction, a phenomenon sometimes
Often results in rapid Less rapid increase in numbers known as alternation of generations. This term is better
production of large numbers confined to land plants and some advanced algae which
of offspring show alternation between a diploid, spore-producing
generation called the sporophyte and a haploid, gamete-
*A common source of confusion is the role played by mitosis and meiosis producing generation called the gametophyte (section
in reproduction. A proper discussion of this depends on a study of 3.3.1). An alternation of asexual and sexual generations
different life cycles and the points at which the two types of nuclear
division occur in the life cycles. These are shown in fig 20.13 and described also occurs in some coelenterates, but here both genera-
in section 20.1.4. tions are diploid and the haploid stage is represented only
A e.g. Chlamydomonas, C e.g. Obelia

Spirogyra, Rhizopus Three different
The adult (sexual) organism sexual organism morphological forms occur sexual generation
is haploid. The only diploid (1) (polymorphism). All are
stage is the zygote. The first diploid. The diploid sexual medusa “meiosis
nuclear division of the generation alternates witha
germinating zygote is by diploid asexual generation.
meiosis, resulting in a return The only haploid cells are gametes
ametes the gametes. (n)
to the haploid condition. 8 (n)
Gametes are produced by y
NB Gametes are not meiosis. peaeal aoe
produced by meiosis.
generation (2n)
LS
BS (2n) ra
S ~ zygote
(2n) reproductive polyp <—— feeding polyp
(2n) (2n)
D e.g. Laminaria (a brown

B_ e.g. Fucus, vertebrates and
sexialOreaniem alga), all land plants i.e.
most other animals sexual generation
bryophytes, pteridophytes
an and spermatophytes. (n)
The only haploid cells are gametophyte
the gametes. meiosis Alternation of haploid and
Gametes are produced by diploid generations occurs.
meiosis. NB Gametes are not
produced by meiosis. gametes
gametes (n)
spores Sv
(n) zygote
\é “0
(Qn)
ee GK acon
zygote generation
(2n)
sporophyte
ig 20. eh life cycles. A, B, C, D represent four different,

Representative j j
commonly occurring, types of life cycle. Note that
though
a a Bonus once in each life cycle. Asexual reproduction of the sexual organism is not shown in A, B and D,
diploid. )
may be possible depending on the species. (n = haploid; 2n =
Jish
by the gametes. This form of alternation of generations is (iii)Gametes are always produced bymeiosis.
sometimes described as metagenesis (section 4.4.3). Dur- . (iv) Meiosis always produces haploid cells|a ,
ing the life cycle of these coelenterates, different forms of ‘mitosis always produces. diploid cells.
the individuals of the species alternate with each other, a (v)Mitosis occurs only in diploid cells.
phenomenon known as cyclic polymorphism. In Obelia, (b) Give examples of ge. to the other our
for example, three forms occur in the life cycle, namely two —
statements.
types of polyp and a medusa (fig 4.11). Polymorphism
occurs in some other organisms, for example, the primrose
(Primula) has two types of flower, thrum-eyed and 20.2 Sexual reproduction in plants
pin-eyed.
The life cycles of parasites are often complex and may All green plants that live on land, together
involve several generations. Each generation is adapted to with some advanced algae, show alternation of generations
a particular situation, either for survival within a host or for
in which a haploid gametophyte generation alternates with
transfer between hosts (for example, the life cycle of a diploid sporophyte generation as shown in fig 20. 13D. It
Fasciola, section 4.5.3). The additional generations pro-
is the gametophyte which undergoes sexual reproduction,
duced by asexual reproduction (polyembryony) serve to producing gametes by mitosis.
increase the numbers of the species. Some representative
The remaining green plants are all algae, and show a
life cycles are shown in fig 20.13.
range of types of sexual reproduction which are summa-
Confusion sometimes occurs over the respective roles
rised in section 3.2.3. In the simple types (generally
played by mitosis and meiosis in asexual and sexual
regarded as primitive) the gametes are all identical and the
reproduction and for this reason the significance of each
process is called isogamy; more complex (or advanced)
within the life cycle is described again here in the context of
types show anisogamy in which dissimilar gametes are
life cycles. Meiosis occurs only in life cycles in which sexual
found, the extreme form of anisogamy being oogamy,
reproduction occurs. When two haploid gametic nuclei
where a motile male gamete swims to a sedentary female
fuse, the zygote produced has the diploid number of
gamete.
chromosomes and starts the next life cycle. Unless meiosis
On leaving water, both plants and animals have evolved
occurs somewhere in this life cycle, the gametic nuclei
strategies over millions of years to cope with dry land. The
produced will be diploid and the zygote would then be
most advanced groups of plants and animals, namely
tetraploid (have four sets of chromosomes). Meiosis is
flowering plants and mammals respectively, owe much of
therefore necessary to prevent chromosome doubling in
their success to adaptations for sexual reproduction, and
every generation in those life cycles in which sexual
certain analogies can be drawn between the two groups.
reproduction occurs. It does not necessarily occur imm-
For example, the bringing together of male and female
ediately before gamete formation, as life cycles A and D
gametes does not rely on gametes being released into
in fig 20.13 show. An additional advantage of meiosis is that
water, as it does for example with bryophytes or amphi-
it contributes to variation by the processes of independent
bians, and there is protection and nourishment of the
assortment of chromosomes and crossing-over of chromo-
developing embryo by the parent. A fundamental dif-
somes (section 22.3). As a direct result, as in life cycles B
ference is the origin of gametes by mitosis from a
and C, or indirectly, as in A and D, the gametes show
gametophyte generation in flowering plants, whereas in
variation. In the special case of alternation of generations
(life cycle D) meiosis occurs in the production of spores mammals there is not alternation of generations and
and not gametes. Variation therefore occurs among the gametes are produced by meiosis from a diploid parent. As
will be shown, however, the gametophyte generation of
spores and the gametophytes that develop from the spores.
This variation is passed on to the gametic nuclei which are
flowering plants is extremely reduced.
produced in the gametophyte by mitosis. In all cases,
20.2.1 The life cycle of flowering plants
however, meiosis can be said to form the basis of sexual
(angiosperms)
reproduction. In life cycles A, B and C, that is life cycles
where alternation of generations does not occur, asexual The ways in which flowering plants have
reproduction results in the production of genetically adapted to life on land have been described in chapter 3.
identical offspring, and involves mitosis. The major adaptations made in reproduction are the
production of seeds and fruits to nourish and protect the
20.1 (a) ‘Which one of the {following . embryo plants and to aid in their dispersal, the absence of
_statements is true, according to the information giver swimming male gametes, and the extreme reduction of the
so far in this chapter? sexual gametophyte generation. Male gametes are carried
(i) Asexual reproduction—always results in ‘the inside pollen grains to the female parts of the plant, a
production of identical hate process called pollination which is followed by the
(ii) Gametes are always nape. production of a pollen tube carrying male nuclei.
An outline of the life cycle of flowering plants is given in
(oz
(a) stigma (b) flower
one ohmore anther
carpels ii t
-> GYNAECIUM j °" ee a et
(female parts)
lity
P
ovary nectary pedicel
ovules
(one or more) SSS
bract
petal____, COROLLA
\ PERIANTH solitary flower raceme main axis
(small leaf-like structure) sepal____,» CALYX (no peduncle) e.g. tulip continues to grow e.g.
Collective names are indicated by b (Tulipa) bluebell (Endymion non-
arTows, e.g. ‘petal—» COROLLA’ receptacle (origin of perianth) scripta), wallflower
means that the collection of petals (Cheiranthus cheiri)
is known as the corolla pedicel (flower stalk)
|
aa
modified racemes
platform for insects youngest flower oldest flower
youngest
peduncle flower
corymb pedicels grow spike flowers are sessile simple umbel pedicels compound umbel more cyme main axis forms a cyme where growth is
to same height e.g. (lack pedicels) e.g. arise from same point common than simple a flower and growth continued by two
candytuft (Iberis) plantain (Plantago) umbel, peduncles arise continues by lateral lateral buds e.g.
from same point e.g. buds e.g. buttercup campion (Silene)
carrot (Daucus), (Ranunculus)
hogweed (Heracleum)
Fig 20.14 (a) LS of a generalised flower. (b) Arrangement of flowers in groups (inflorescences) or in isolation. Some common
types of inflorescence are shown which can be described as racemose or cymose
fig 3.37, where it is compared with representative life cycles year. Its environmental and hormonal control is described
of other plant groups. There is still an alternation of in section 15.5.
generations in the life cycle, as is shown more simply in fig The parts of the flower are arranged spirally (in a few
20.13D. Since the gametophyte generation is virtually primitive flowers) or in whorls around the upper part
non-existent and is not free-living, it would be difficult to (receptacle) of a flower stalk (pedicel). A generalised
realise that alternation of generations occurs were it not for flower and the arrangement of flowers in groups (inflores-
the comparison that can be made with more primitive cences) are shown in fig 20.14, and some of the terms used
ancestors. The life cycle of flowering plants is described in to describe the flower parts are explained below.
more detail below and, strictly speaking, involves asexual
Parts of a flower
reproduction of the dominant sporophyte generation (the
flowering plant) and sexual reproduction of the game- The inflorescence is a collection of flowers borne on the
tophyte generation. same stalk, the peduncle (see fig 20.146). A collection of
flowers may be more attractive to pollinating insects than a
small solitary flower.
20.2.2 The structure and functions of the The receptacle is the end of the flower stalk (pedicel)
flower
from which the perianth, gynaecium and androecium arise.
The common use of the term ‘flowering plants’ The receptacle and flower are described as hypogynous if
to describe the angiosperms is a reference to the unique- the stamens and perianth are inserted below the gynaecium
ness of this group in producing flowers. Flowers are as in fig 20.14a, epigynous if stamens and perianth are
reproductive structures whose evolutionary origins are inserted above the ovary, and perigynous if the receptacle
unclear, but which are sometimes regarded as collections of is flattened or cup-shaped with the gynaecium at the centre
highly specialised leaves, analogous to the cones of and the stamens and perianth attached round the rim. (See
gymnosperms in that they carry the spore-producing also inferior and superior ovaries under gynaecium below.)
structures. They can be regarded as organs of both asexual The perianth consists of two whorls of leaf-like parts
and sexual reproduction, asexual because they produce called perianth segments. In monocotyledons the two
spores (pollen grains and embryo sacs) and sexual because whorls are usually similar, for instance daffodil (Narcis-
gametes are later produced within the spores. They are sus), tulip (Tulipa) and bluebell (Endymion non-scripta).
commonly referred to simply as organs of sexual reproduc- In dicotyledons the two whorls are usually different,
tion. In areas where there are distinct seasons, flowering is consisting of an outer whorl of sepals called the calyx and
usually synchronised with a particular season and time of an inner whorl of petals called the corolla.
733
The calyx is the collection of sepals. Sepals are usually Monoecious plants — separate male and female flowers
green and leaf-like structures that enclose and protect the borne on the same plant, such as oak (Quercus), hazel
flower buds. Occasionally they are brightly coloured and (Corylus), beech (Fagus) and sycamore (Acer
petal-like, serving to attract insects for pollination. pseudoplatanus)
Polysepalous — having free (unfused) sepals Dioecious (unisexual) plants — male and female sex
Gamosepalous — having sepals that are at least partly organs borne on separate plants, that is the plants are
fused into a tube. either male or female, for example yew (Taxus),
The corolla is the collection of petals. In insect- willow (Salix), poplar (Populus) and holly (Ilex)
pollinated flowers the petals are usually large and brightly Hermaphrodite (bisexual) flowers — male and female sex
coloured, serving to attract insects. In wind-pollinated organs in the same flower, such as in buttercup, white
flowers the petals are usually reduced in size and green, or deadnettle, bluebell and pea
may be entirely absent. Unisexual flowers — separate male and female flowers,
Polypetalous — having free (unfused) petals, for ex- for example oak, hazel, yew, willow, poplar and holly
ample pea (Pisum), rose (Rosa) and buttercup Symmetry of flowers
(Ranunculus)
Gamopetalous or Sympetalous — having petals that are If the flower parts are arranged in radial symmetry around
at least partly fused into a tube, for example foxglove the receptacle, the flower is said to be regular or
(Digitalis), primrose (Primula), dandelion (Tarax- actinomorphic, for instance buttercup and bluebell. If the
acum) and white deadnettle (Lamium album) flower shows bilateral symmetry only, it is said to be
The androecium is the collection of stamens, forming the irregular or zygomorphic, as in white deadnettle and pea.
male reproductive organs of the flower. Each stamen Representative flowers. Some representa-
consists of an anther and a filament. The anther contains tive insect-pollinated flowers are illustrated in figs 20.15—
the pollen sacs, in which pollen is made. The filament 20.18. Fig 3.38 illustrates the floral morphology of a typical
contains a vascular bundle that carries food and water to grass, a wind-pollinated monocotyledon.
the anther.
The gynaecium or pistil is the collection of carpels, stamen
forming the female reproductive organs of the flower. A numerous and spirally arranged on the receptacle;
collection of stamens forms the androecium
carpel consists of a stigma, style and ovary. The stigma
filament anther
receives the pollen grains during pollination and the style white, short yellow
bears the stigma in a suitable position in the flower to
receive the pollen. The ovary is the swollen, hollow base of
the carpel and contains one or more ovules. Ovules are the
structures in which the embryo sacs develop and which,
stigma
after fertilisation, become seeds. Each is attached to the curved
ovary wall by a short stalk called the funicle and the point of
attachment is called the placenta. style A(X)
The carpels of a flower may be separate and free, the very short a Q \\ep
apocarpous condition, as in buttercup, or fused to form a .
ey
Naa?
\ 3 ) WAL
single structure, the syncarpous condition, as in white

sepal
SEN"
SS
"ss |_
g
deadnettle. The ovary formed by fusion of carpels may five form calyx; green, hairy, petal
have a single chamber or loculus (unilocular) or several curved round petal, free five form corolla;
ovar bright yellow,
loculi (multilocular) with one loculus formed by each y |
apocarpous, superior, one BLOssyUPPEL
constituent carpel. The styles of syncarpous flowers may be carpel, one locule, one ovule;
surface, free
perianth
(polypetalous)
fused or separate. collection of carpels forms the
gynaecium; carpels are spirally nectary
Superior ovary — an ovary inserted above the other arranged on the receptacle covered by a
flower parts on the receptacle, that is the ovary of a small scale at
base of petal
hypogynous or perigynous flower
Division: Spermatophyta
Inferior ovary — an ovary inserted below the other Class: Angiospermae
receptacle
flower parts on the receptacle, that is the ovary of an Subclass: Dicotyledonae pedicel
epigynous flower Family: Ranunculaceae
The nectaries are glandular structures that secrete Fig 20.15 Ha/f-flower of meadow buttercup (Ranunculus
nectar, a sugary fluid that attracts animals for pollination, acris), a polypetalous dicotyledon. It is a herbaceous
usually insects, but also birds and bats in the tropics. perennial common in damp meadows and pastures. It
The following terms are applied to whole plants and perennates by means of a rhizome. Flower: actinomorphic
flowers. and hypogynous. Flowers appear April to September.
Pollination: insects such as flies and small bees. Fruit: each
Hermaphrodite (bisexual) plants — male and female sex carpel contains one seed and forms afruit called an achene.
organs borne on the same plant No special dispersal mechanism
734
The structure of a flower is best illustrated by means ofa _ plane, so as to split the pedicel in two longitudinally) and
half-flower, a view of the flower obtained by cutting the drawing one half, showing the cut surface as a continuous
flower vertically into two equal halves (along the median _line.
Half-flower
standard petal (one)
one free stamen standard petal
wing petal (two)
keel petal (two fused petals) landing

sepal
platform
stigma
for bees
receptacle style
anther
stamen
pedicel filament | (ten present)
i (i) whole flower
ovule
nectary ; ovary (one carpel) becomes a
locule pod after fertilisation (see fig
trough formed 20.28)
by nine fused I! faced i
filaments
stamens,
of nine ovat wie stamens
surrounds ovary aren
e free stamen
Division: Spermatophyta
Class: Angiospermae Q
Subclass: Dicotyledonae Croeciu® (iii) half keel removed (iv) stamen trough removed
Family: Leguminosae nine fused filaments Progressive dissection of pea flower, removing shaded parts
Fig 20.16 (above) Structure of flower of sweet pea (Lathyrus odoratus), a polypetalous dicotyledon. Flower: actinomorphic
and polypetalous. Flowers appear in July. Calyx: five sepals. Corolla: five petals, one standard petal, two wing petals, two fused
keel petals interlocking with wing petals. May be white or coloured. Pollination: bees are attracted by colour, scent and nectar.
The standard petal is especially conspicuous. The two wing petals act as a landing platform. When a bee lands, its weight pulls
them down together with the keel to which they are linked. The style and stigma emerge, striking the undersurface of the bee
that may be carrying pollen from another flower. As the bee searches for nectar at the base of the ovary with its long proboscis,
the anthers may rub pollen directly on to the undersurface of the style, from where it may be passed to the bee. Self-pollination
may also occur. The garden pea (Pisum sativum) is similar, but more commonly self-pollinated. Fruit: a pod consisting of one
carpel with many seeds. (See fig 20.28 for self-dispersal.)
Fig 20.17 (below) Half-flower of white deadnettle (Lamium album), a sympetalous dicotyledon. The plant is a herbaceous
perennial common in hedgerows and on waste ground. It perennates by means of a rhizome. Flower: zygomorphic (irregular)
and hypogynous. Flowers appear in April to June and autumn. Pollination: mainly bumblebees. A bee lands on the lower lip
and as it enters the flower its back, which may be carrying pollen, touches the stigma first, thus favouring cross-pollination,
although self-pollination is possible. Fruit: four nutlets, each with a hard wall and containing one seed. The fruit is called a
carcerulus
anther of anterior stamen, stamen,
upper lip of corolla | four form androecium,
black with yellow along line
shelters stamens and style which are attached to petals,
of dehiscence, hairy
positioned to touch the bee’s back as filament anterior two have
it enters the flower; two petals white longer filaments
stigma
petal,
bilobed, projects below anthers
five form corolla, fused
(gamopetalous), white, corolla has
style
two lips (bilabiate)
long and thin
sepal, lower lip of corolla
five form calyx, fused (gamosepalous), acts as a landing platform for insects,
green, hairy one petal
ring of hairs
at narrowing of corolla tube prevents
receptacle access to nectaries by small insects
ovary
pedicel syncarpous, superior, two carpels,
very short four locules (original two locules
further divided by ‘false’ septum),
one ovule p per locule
Division: Spermatophyta nectary
Class: Angiospermae two present, swellings at base of
Subclass: Dicotyledonae anterior side of ovary
Family: Labiatae
735
perianth segments
six present in two whorls ofthree, blue
or violet (rarely pink or white), fused
into a tubular structure at their bases
anther
small in outer whorl {] stigma
small and three-lobed
androecium stamen
i i of outer whorl | filament ; aie
anaes a attached to middle of
three, attached poe ae and thin
to perianth anther gynaecium
segments, large in inner whorl
anthers cream
in colour stamen ovary '
of inner whorl | filament
attached near base of syncarpous, superior, three
perianth carpels, three locules, many
ovules on axile placentae
ovules
loculepag. a) speak ieee eae
YG receptacle
Division: Spermatophyta placenta
Class: Angiospermae if
Subclass: Monocotyledonae | pedicel
Family: Liliaceae
Fig 20.18 Half-flower of bluebell (Endymion non-scripta), a monocotyledon. The plant is an herbaceous perennial, common
in woods and hedgerows, favouring shady positions and light, acid soils. It perennates by means of a bulb. Flower:
actinomorphic and hypogynous. Parts are in threes, as is typical of monocotyledons. Flowers appear in April to May. :
Pollination: insects such as the honey bee. The flower is slightly scented and nectar is produced at the top of the ovary. Fruit: a
capsule dehiscing into three valves
pollen mother cell connective, contains vascular

dividing by meiosis bundle from filament
anther fibrous layers outer wallrs. ee

cell walls develop one layer of large cells
thickened strips
filament concerned with __) inner wall
dehiscence of several layers of flattened
anthers cells
stamen
tapetum
cells provide nourishment for pollen mother
cells; this layer gradually breaks down as
pollen grains form
epidermis
line of dehiscence
pollen sacs
four present in anther before dehiscence after dehiscence
Fig 20.19 7S mature anther before and after dehiscence
Development of pollen grains
OO Ge
Fig 20.20 Development of pollen grains Each stamen consists of an anther in which four pollen sacs
meiosis I meiosis I] <> secretion of walls produce pollen, and a filament which contains a vascular
bundle supplying food and water to the anther. Fig 20.19
<e> © illustrates the internal structure of an anther, with its four
pollen mother tetrad of four haploid separate pollen
cell (diploid) pollen cells e
grains B pollen sacs containing spore mother cells. Each spore
mother cell undergoes meiosis to form four pollen grains as
exine thick, outer shown in fig 20.20.
generative nucleus sculptured wall
later forms two male Immediately after meiosis the young pollen grains are
intine thin inner wall
gametes seen in tetrads (groups of four). Each grain develops a thick
pollen tube nucleus pit wall, often with an elaborate sculptured pattern character-
one mature pollen grain istic of the species or genus. The outer wall, or exine, is
736
made of a substance related to cutin and suberin (sporopol- anatropous; if the funicle and ovule remain straight, then
lenin) but is more durable than either. It is one of the most the ovule is described as orthotropous.
resistant substances in nature and allows grain coats to The main body of the ovule is the nucellus, enclosed and
survive unchanged over long periods of time, sometimes protected by two sheaths or integuments. A small pore is
millions of years. This fact, together with the ease of left at one end of the ovule, the micropyle. The other end of
identifying the parent genus or species which produced the the ovule, where the funicle joins to the nucellus and
grain, has given rise to the science of palynology or pollen integuments, is called the chalaza.
analysis. By studying pollen grains from a particular time Within the nucellus, at the end nearest the micropyle,
and place it is possible to determine what plants were one spore mother cell develops, known as the embryo sac
growing and thus to gain information about, for example, mother cell. This diploid cell undergoes meiosis and gives
the ecosystems (including animals) and climate of that rise to a haploid megaspore or embryo sac as shown in fig
time. A particularly abundant source of pollen grains is 20.21. The embryo sac grows and as soon as its nucleus
peat, which accumulates to great depths over long periods divides by mitosis its contents can be regarded as a female
of time in peat bogs. Cores of peat can be extracted using gametophyte. Continued mitotic divisions result in the
special peat borers. production of eight nuclei, one of which is the nucleus of
the female gamete.
Two polar nuclei migrate to the centre of the embryo sac
and fuse to become asingle diploid nucleus (in some cases
they fuse later at fertilisation). The remaining six nuclei,
three at each end, become separated by thin cell walls and
The pollen grain at this stage is equivalent to a only one of these, the female gamete, appears to serve any
microspore, as discussed in chapter 3. Its nucleus divides further function.
into two by mitosis to form a generative nucleus and a The final appearance of the mature carpel at fertilisation
pollen tube nucleus (fig 20.20). As soon as this happens, the is shown in fig 20.22.
contents of the pollen grain can be regarded as equivalent
to a male gametophyte, because male gametes will be 20.2.4 Pollination
formed from the generative nucleus. After formation of pollen grains in the pollen
sacs, the cells in the walls of the anther begin to dry and
20.2.3 Development of the embryo sac and shrink, setting up tensions that eventually result in splitting
female gamete (dehiscing) of the anther down the sides along two lines of
Each carpel consists of a stigma, style and weakness (fig 20.19). The pollen grains are thus released.
ovary. Within the ovary one or more ovules develop, each The transfer of pollen grains from an anther to a stigma is
attached to the ovary wall at a point called the placenta bya called pollination. This must be achieved if the male
short stalk or funicle through which food and water pass to gametes, which develop inside the pollen grains, are to
the developing ovule. Often, as it develops, the ovule reach the female gamete, and special, often elaborate,
bends over so that its tip is facing downwards near the base mechanisms have evolved that ensure successful pollina-
of the funicle. If this occurs, the ovule is described as tion.
stigma nucellus
style MiteSa embryo sac
see below
ovary
Y
oung ovule micropyle
aoe funicle
carpel young ovule
inner cell grows three

antipodal cells
mitosis mitosis mitosis _ nuclear
eee = > —_> two polar nuclei en diploid nucleus
degenerate growth usion
ovum
embryo sac (e) aren (female gamete)
motheret cell pounce’
y nucellus , mature embryo
h sac
eae(female
(diploid) ss megaspore endnearest female gametophyte two synergids Seabees ee tre
Bit See isas Eiickop ile sos cinbiyiosac six are haploid, one is diploid
Fig 20.21 Development of embryo sac and female gamete (in an anatropous ovule)
ia
Key germinating pollen grain
stigma
fe diploid tissue
style
[| haploid tissue:
pollen tube
contents of pollen_
grain and pollen tube is ovary wall
equivalent too”
gametophyte. Embryo ocelot in
‘animals than
an :
sac 1s equivalent
to
Dichogamy. Sometimes anthers mature and

integuments stigmas become receptive at different times, a condition
__nucellus known as dichogamy. If the anthers mature first, this is
three antipodal cells described as protandry, if the stigmas mature first it is
ovule diploid nucleus protogyny. Protandrous flowers are much more common,
formed by fusion —| embryo for example white deadnettle, dandelion, rosebay willow-
of polar nuclei sac
herb (Epilobium angustifolium); see also sage (Salvia) (fig
20:23):
Protogyny occurs in bluebell and figwort (Scrophularia).
male nuclei Penden In most cases of dichogamy there is an overlapping period
tube nucleus { pollen tube when both anthers and stigmas are ripe, thus allowing
selfing if crossing has been unsuccessful. A similar
Fig 20.22 LS carpel at fertilisation. The ovule illustrated is mechanism for ensuring cross-fertilisation exists in some
anatropous. Note that the ovule, which becomes the seed hermaphrodite animals, for instance Hydra, where the
after fertilisation, contains both diploid parent tissue and
haploid embryo sac tissue testes ripen before the ovary.
Self-incompatibility (self-sterility). Even if
Transfer from an anther to a stigma of the same flower, self-pollination does occur, the pollen grain often does not
or a flower on the same plant, is called self-pollination. develop, or develops very slowly, so preventing or
Transfer of pollen from the anther of one plant to the discouraging self-fertilisation. In all such cases there is a
stigma of another plant is called cross-pollination. Cross- specific inhibition of pollen penetration of the stigma, or of
pollination leads to cross-fertilisation and has the advan- pollen tube growth down the style, and this is genetically
tage of increasing the chances of variation. It is thus a form determined by self-incompatibility genes.*
of outbreeding. There are often special features to When self-incompatibility occurs, the extent of compa-
encourage it, some of which are described below. tible cross-pollination is variable and again is genetically
Self-pollination leading to self-fertilisation has the determined. For most efficient use of the pollen, a high
advantage of greater reliability, particularly where mem- proportion of crosses should be compatible. An extreme
bers of the species are uncommon and are separated by example is clover, where all plants are self-incompatible,
large distances. This is because it is not dependent on an but cross-incompatibility occurs between less than one in
external agency, such as wind or insects, to deliver the 22 000 pairs. A less efficient system occurs where the
pollen. However, self-fertilisation is the extreme form of compatible types are characterised by differences in floral
inbreeding and can result in less vigorous offspring as morphology. An example is the primrose (Primula),
described in section 25.4. Self-pollination is used in discussed below.
groundsel (Senecio) and chickweed (Stellaria), which
produce no nectar or scent. S| Self-incompatibility iscontrolled
Both cross- and self-pollination have advantages and
disadvantages and many plants balance the advantages by ] alleles, - S,and S,, and (b) that self
devices which favour cross-pollination but allow selfing to _ occurs ifthe pollen grain and the sty
occur if crossing fails. For example, some of the buds - allele in common, what proportic
produced by violet (Viola) and wood sorrel (Oxalis) never _ grains from a plant with the ger
open, so that self-pollination inside these is inevitable. _ capable of successfully ge '
the acronre S2So?
Features favouring cross-pollination
Dioecious plants. Self-pollination in dioe-

cious plants is impossible. Monoecious species, where
separate male and female flowers occur on the same * The genetics of self-incompatibility and its commercial implications are
outside the scope of this book, but are dealt with in Sexual Incompatibility
hermaphrodite plant, also favour cross-pollination but in Plants, Institute of Biology, Studies in Biology, No. 110; Lewis, Dan;
selfing may also occur. Arnold, (1979).
738
and, as the head of the bee pushes
Fig 20.23 A bee entering a flower of meadow sage. The stamens are hinged to a plate
bee’s abdomen. The stigma of the flower increases in
against this, the stamens are lowered. This brushes pollen onto the
into contact with the stigma and the pollen will be
length as the flower ages. If a bee now enters an older flower, it will come
cross-pol lination to take place
transferred from its abdomen to the stigma. This series of activities causes
Special floral structures. In most her- Wind pollination and insect pollination
maphrodite flowers there are structural features that Pollen grains are spores, but unlike the spores of the
favour cross-pollination. non-seed bearing plants they cannot germinate on land and
In the case of insect-pollinated flowers the stigma is must be transferred to the female parts of either cones, in
usually borne above the anthers, thus removing the the case of gymnosperms, or flowers in the case of
possibility of pollen falling on to the stigma of the same angiosperms. The original agent of spore dispersal was
flower. A visiting insect, possibly carrying pollen from wind, but this is very inefficient in pollen transfer because
another plant, will touch the stigma first as it enters the the mechanism relies solely on chance interception of the
flower. Later, while the insect is seeking nectar, pollen is pollen grains by the cones or the flowers. All conifers, and
either brushed against it or falls on to it before it leaves the many flowering plants such as grasses and most temperate
flower. This occurs in white deadnettle (fig 20.17). A more trees, such as oak and hazel, still rely on wind, but it is at
primitive mechanism may ensure that the stigma brushes the expense of producing enormous quantities of pollen, a
against the insect as it lands, as in pea (see fig 20.16 for drain on the plant’s materials and energy. Insect pollina-
details). Such mechanisms are generally reinforced by tion was rapidly exploited once flowers evolved because the
dichogamy and the flowers are often complex and insect is a much more precise agent of dispersal. It can carry
zygomorphic in shape. a small amount of pollen from the anthers of one flower and
Flowers attract insects by providing a source of food deposit it precisely on the stigma of another flower. As a
(nectar or pollen) and stimulating the senses of sight and result, special relationships between flowers and insects
smell of the insects. Those characteristics that enable have evolved, the reward the insects receive from the
flowers to do this are discussed below. flowers being food in the form of nectar, and for some,
In the case of wind-pollinated flowers the stamens, the pollen. The insects specialised for flower-feeding appeared
whole flower, or the inflorescence may hang downwards so at the same time as the flowering plants, and include bees,
that falling pollen will drop clear of the plant before being
blown away, for example hazel catkins. Table 20.2. Summary of typical differences between wind-
and insect-pollinated flowers
20.6 Fig 20.24 shows two types ofprim-
_ rose flower. They occur naturally in roughly equal Typical wind-pollinated flower Typical insect-pollinated flower
_ numbers anddiffer in length of style (heterostyly) and_ Small petals not brightly Large coloured petals; flowers
_ position of anthers. (a) Given that bees colle coloured (usually green), or therefore conspicuous. If flowers
rom the base of the corolla tube, exp petals absent; flowers there- relatively inconspicuous they
ross-pollination between pin-eyed and th fore inconspicuous may be gathered together in
inflorescences
lowers rather than between flowers of 1 ame
type, is as ()Whatis theenovanioge ofsucha Not scented Scented
: system? Nectaries absent Nectaries present
Large branched and feathery Small stigma, sticky to hold
Although the ie described in ars 20.6 and stigma hanging outside flower pollen and enclosed within
to trap pollen flower
fig 20.24 apparently favours outbreeding, a much more
important difference between pin-eyed and thrum-eyed Pendulous stamens hanging Stamens enclosed within flower
outside flower to release
primroses is a self-incompatibility mechanism which is pollen
more effective in restricting cross-fertilisation to pin-eye/
Anthers versatile, i.e. attached Anthers fixed at their bases
thrum-eye crosses. The genes that control incompatibility, only at midpoints to tip of (basifixed) or fused along their
style length and anther height lie close together on the same filament so that they swing backs to the filaments
chromosome and behave asa single inheritable unit. freely in air currents (dorsifixed) so that they are
immovable
Large quantities of pollen Less pollen produced
owing to high wastage
Pollen grains relatively light Pollen grains relatively heavy
and small; dry, often smooth, and large. Spiny walls and sticki-
walls ness help attachment to insect
anthers body (fig 20.26)
stigma
Flower structure relatively Elaborate structural modifica-
stigma simple
anthers tions for particular insects often
occur
ny
) nectary
Y nectary Flowers borne well above Position and time of appearance
foliage on long stalks (e.g. variable in relation to foliage,
grasses) or appear before though often borne above it for
(a) pin-eyed (b) thrum-eyed leaves (e.g. many British increased conspicuousness
trees)
Fig 20.24 Heterostyly in primrose (Primula) flowers moo
aea_=<_—>*_$[————————maoana(#agpaeqonaeoaeeaoooaoaonwwx~—~—_—_S
DD
740
wasps, butterflies and moths. In a few particular cases the More specific than colours are the scents produced by
insect and the plant it pollinates are so interdependent that flowers some of which, like lavender and rose, are used by
neither species can survive without the other, such as the Man in perfumes. Smells of rotting flesh that attract
yucca plant and its associated moth. carrion-eating insects are also produced by some plants:
Insect pollination has the important additional advan- the arum lily (Arum maculatum) attracts dung flies.
tage that it encourages cross-pollination and hence cross- Specificity for recognition is also provided by flower
fertilisation, so the modifications of flowers to encourage shape.
insect pollination described below could be added to the list One of the most complex and bizarre mechanisms for
of features favouring cross-pollination. ensuring cross-pollination is the sexual impersonation of
In order to attract insects, flowers generally are large, female wasps by certain orchids. The flower parts mimic
with brightly coloured petals or, if small, are grouped into the shape, colourings and even the odour of the female
inflorescences. Insects can see ultra-violet wavelengths that wasp, and the impersonation is so convincing that the male
are invisible to Man and so flowers that appear white to wasps attempt to copulate with the flower (fig 20.25). While
Man may, in fact, appear coloured to insects. Often there doing so they deposit pollen and, on leaving the flower,
are markings on the petals such as lines, spots or an collect fresh pollen to take to the next flower.
increased intensity of colour that guide insects to the A summary of the typical differences between wind-
nectaries, as in violet and pansy (Viola), orchids (Orchis pollinated and insect-pollinated flowers is given in table
and other genera) and foxgloves (Digitalis). DOE2.
Fig 20.25 A digger wasp

(Argogorytes mystaceus) —
copulating with the fly orchid
(Ophrys insectifera)
741
surface of stigma
PONS etM TEE Mmaniceecuity
spiny pollen grains
A ees
Fig 20.26 bonne een electron
Scanning ors micrograph
graph of
of pollen
polle grains
] on the stigma
i i
of a flower. The spiked surface of the grains is typical
742
20.2.5 Fertilisation. with the surface of a dehisced anther. Observe the slide at
Once a pollen grain has landed on the stigma, intervals over a period of 1-2 hours. The nuclei at the tip of
a sucrose solution secreted by epidermal cells of the stigma growing tubes may be stained by irrigating with a drop of
stimulates germination of the grain and possibly supplies acetocarmine or neutral red.
food (fig 20.26). A pollen tube emerges from one of the
pores in the wall of the pollen grain and grows rapidly down 20.2.6 Development of the seed to dormancy
the style to the ovary. Its growth involves secretion of
digestive enzymes and is controlled by the tube nucleus of Immediately after fertilisation, the ovule
the pollen grain, which is found at the growing tip of the becomes known as the seed and the ovary the fruit.
tube. Growth is stimulated by auxins produced by the The zygote grows by mitotic divisions to become a
gynaecium, and the pollen tube is directed towards the multicellular embryo which consists of a first shoot, the
ovary by certain chemicals, an example of chemotropism. plumule, a first root, the radicle, and either one or two
It is probably also negatively aerotropic, that is it grows seed-leaves called cotyledons (one in monocotyledons and
away from air. Growth depends on compatibility between two in dicotyledons). These cotyledons are simpler in
the pollen and the style tissue as already described. structure than the first true foliage leaves and may become
During growth of the pollen tube the generative nucleus swollen with food to act as storage tissue, as in the pea and
of the pollen grain divides by mitosis to produce two male broad bean (Vicia faba). The plumule consists of a stem,
nuclei that represent the male gametes (fig 20.22). They are the first pair of true foliage leaves and a terminal bud. The
non-motile, unlike the sperms of lower plants, and depend triploid primary endosperm nucleus undergoes repeated
on the pollen tube to reach the female gamete which is divisions (mitotic) to form the endosperm, a mass of
triploid nuclei which are separated from one another by
located in the embryo sac of the ovule. The pollen tube
thin cell walls. In some seeds this remains as the food store,
enters the ovule through the micropyle, the tube nucleus
as in maize (Zea mays).
degenerates and the tip of the tube bursts, releasing the
If the cotyledons act as a food store they grow at the
male gametes in the vicinity of the embryo sac which they
expense of the endosperm, which may disappear
enter. One nucleus fuses with the female gamete, forming a
altogether. Some seeds store food in both endosperm and
diploid zygote, and the other fuses with the two polar nuclei
cotyledons.
(or diploid nucleus if the latter have already fused) forming
Thus within the developing seed, both embryo and in
a triploid nucleus known as the primary endosperm
some cases the endosperm, grow within the embryo sac. As
nucleus. This double fertilisation is unique to flowering
their growth continues the surrounding nucellus becomes
plants.
disorganised and breaks down, supplying nutrients for
If, as is often the case, more than one ovule is present in
growth. Further nutrients are supplied by the vascular
the gynaecium, each must be fertilised by a separate pollen
bundle in the stalk (funicle) of the ovule. There are close
grain if it is to become a seed. Thus each seed may have
analogies here with the mammal, where food passes from
been fertilised by a pollen grain from a different plant.
the parent to the developing embryo via the placenta. As
already noted, the term placenta is also used for the point
Experiment 20.1: To investigate the growth of of attachment of the funicle to the ovary wall.
pollen tubes The micropyle remains a small pore in the testa through
which oxygen and water will enter when the seed
Stigmas secrete a solution containing sucrose ranging in germinates. The testa is a thin but tough protective layer
concentration from about 2 to 45%. This helps to stick derived from the integuments. The final stages in seed
pollen grains to the stigma and to promote their germina- maturation involve a reduction in the water content of the
tion. The addition of borate to the experimental solution seed from the normal levels for plant tissues of about 90%
helps to prevent osmotic bursting of pollen tube tips and by mass to about 10-15% by mass. This markedly
stimulates growth. reduces the potential for metabolic activity and is an
Materials essential step in ensuring seed dormancy.
While the seeds develop, the ovary becomes a mature
microscope
fruit, its wall being known as the pericarp. The changes that
cavity slide
occur vary with species, but generally the fruit is adapted to
flowers containing dehiscing anthers, such as dead-
protect the seeds and to aid in their dispersal as discussed in
nettle or wallflower
section 20.2.8.
10-20% (w/v) sucrose solution also containing sodium The remaining flower parts wither and die and are
borate to a concentration of 0.01% abscissed in a controlled manner, just as leaves are in
acetocarmine or neutral red deciduous plants. In a few cases structures such as the
Method receptacle, style or sepals are retained and contribute to
dispersal. If the receptacle is involved, as in strawberry
Place a drop of sucrose solution in the central depression
(Fragaria), the structure is called a false fruit.
of acavity slide and add pollen grains by touching the drop
743
Table 20.3. Summary of the changes that occur after fertilisation in flowering plants
SS
After fertilisation
plumule
zygote > embryo radicle
one or two
cotyledons
(may or may
not store food)
primary ———— endosperm (may or may seed — a fertilised ovule fruit —a fertilised ovary. It contains one or
ovule endosperm not persist as a food store) (one scar — funicle stalk) moreseeds(twoscars—receptacle and style)
nucleus
nucellus ————> disappears
integuments
—> testa
ovary wall ——~ pericarp
expanding embryo sac Advantages

endosperm (1) The plant is independent of water for sexual reproduc-
tion and therefore better adapted for a land environ-
lil embryo cell first
division of ment.
suspensor cell
Gg zygote by S (2) The seed protects the embryo.
ao mitosis
i l (3) The seed contains food for the embryo (either in
food from aca, cotyledons or in the endosperm).
parent plant
cotyledons (4) The seed is usually adapted for dispersal.
YW ;
embryo i]
i developing (5) The seed can remain dormant and survive adverse
\ plumule conditions.
and radicle
(6).The seed is physiologically sensitive to favourable
conditions and sometimes must undergo a period of
after-ripening so that it will not germinate immediately
cotyledons (see chapter 15).
testa growing
(7) The seed is a product of sexual reproduction and
cotyledons a plumule
swollen with > A therefore has the attendant advantages of genetic
; ~——
food | variation.
remains of . radicle
endosperm GF Disadvantages
micropyle
(1) Seeds are relatively large structures because of the
Fig 20.27 Growth of an embryo in a non-endospermous extensive food reserves. This makes dispersal more
dicotyledon seed, such as shepherd's purse (Capsella bursa- difficult than by spores.
pastoris)
(2) Seeds are often eaten by animals for their food
reserves.
The hormonal control of fruit development is discussed (3) There is a reliance on external agents such as wind,
in section 15.3.5.
insects and water for pollination. This makes pollina-
Some of the changes that occur after fertilisation are
tion (and hence fertilisation) risky, particularly wind
summarised in table 20.3. pollination.
Some stages in the development of the embryo are
(4) There is a large wastage of seeds because the chances of
shown in fig 20.27. survival of a given seed are limited. The parent
sporophyte must therefore invest large quantities of
20.2.7 Advantages and disadvantages of material and energy in seed production to ensure
reproduction by seed
success.
The seed is the characteristic product of sexual (S) The food supply in a seed is limited, whereas in
reproduction in the spermatophytes, being defined as a vegetative reproduction food is available from the
fertilised ovule. It contains an embryo plant with one or parent plant until the daughter plant is fully estab-
more cotyledons, sometimes an endosperm, and is sur- lished.
rounded by a protective testa. (6) Two individuals are required in dioecious species
744
making the process more risky than reproduction in dehiscent or indehiscent according to whether they do or
which only one parent is involved. However, dioecious do not release seeds by splitting. Alternatively dry fruits
plants are relatively rare. may split into a number of one-seeded portions and are
The information provided above can be used to compare then described as schizocarpic.
the advantages and disadvantages that seed-bearing plants The further the seeds are dispersed, the less chance there
have compared with non-seed-bearing plants, or to is of competition from the parent plant. There is also more
compare the relative merits of sexual reproduction and chance of finding a fresh area to colonise, thus increasing in
vegetative propagation within the seed-bearing plants. time the overall population size. However, there is an
attendant risk of the seed not finding a suitable place for
germination if it is dispersed some distance from the
20.2.8 Fruit and seed dispersal
parent.
After seed development either the entire fruit There are three major external agents of dispersal,
or the seed(s) contained within it are dispersed from the namely wind, animals and water. In addition self-dispersal
parent sporophyte. If the pericarp (wall) of the fruit mechanisms exist, often involving an explosive release of
becomes hard and dry it is called a dry fruit and if it seeds from the fruit. Examples of all these types of
becomes fleshy, a succulent fruit. Dry fruits may be dispersal are given below, fig 20.28, pages 745-8.
Fig 20.28 Examples of different types of fruit and seed and their methods of dispersal
ANIMAL DISPERSAL Animal dispersal is in general more reliable than wind dispersal because animals tend to frequent fertile places suitable for
seed germination
(a) Hooked fruits e.g. goosegrass (Galium), avens Geum, burdock (Arctium), carrot (Daucus), buttercup (Ranunculus),
Goosegrass or cleavers bur-marigold (Bidens), agrimony (Agrimonia)
(Galium aparine)
Co hooked pericarp
PICS hard hooked spines on
Ar? ety <2 receptacle for animal
two one-seeded SSUES at dispersal
AK
portions of fruit BY
pedicel receptacle covered

with fine hairs ;inside
Hooked or spiny fruits become attached to the skin. fur or wool of passing are two carpels but
animals and may be carried some distance before dropping off or being Agrimony only one develops a
scratched off. The hooks or spines develop from various parts ofthe fruit seed
or surrounding structures. In Geum the style is hooked. In burdock a
collection of small fruits is surrounded by hooked bracts (modified leaves) Agrimony has hard hooked
forming one structure. In addition, small stiff hairs break off and spines on its receptacle.
penetrate the skin, causing irritation and scratching with consequent
removal of the fruits. Goosegrass, carrot and buttercup all have hooked
pericarps. In bur-marigold the fruit has a pappus like the dandelion, but
with strong barbs.
Scanning electron micrograph of the

fruit of goosegrass, showing the
numerous small hooks which
develop from the pericarp. The
hooks cling very effectively to the
skin, fur or wool of passing animals
745
ANIMAL DISPERSAL
(b) Succulent fruits e.g. plum (Prunus), blackberry (Rubus), tomato (Lycopersicum), apple (Malus), strawberry (Fragaria)
Succulent fruits have fleshy parts that provide food for animals, including
birds. They usually attract animals by becoming brightly coloured and (c) False succulent fruits
scented as they ripen. The fruit is eaten and digested but the seeds are Sometimes the receptacle becomes swollen and fleshy and itself resembles
resistant to digestive enzymes and pass unharmed through the gut of the a fruit. Such fruits are called false fruits. Examples are apple and
animal, to be deposited in the faeces, often on fertile soil. Nutrients from the strawberry.
decomposition of the faeces may increase the fertility around the seed. Apple
Representative examples of different types of fruit structure are illustrated The apple is an example of a false fruit called a pome. In pomes the flesh
below. ofthe fruit is formed from the hollow receptacle which surrounds and
encloses the carpels. The pericarpofthe true fruit becomes the ‘core’
Plum which contains the seeds (‘pips’). Other examples are pear (Pyrus) and
The plum is an example of a fruit called a drupe in which the pericarp has hawthorn (Crataegus).
three layers, an epicarp (protective skin), a mesocarp (succulent) and an
LS apple remains of sepals (originally at
inner woody endocarp, (‘stone’) that protects the seed and resists digestion.
rim of cup-shaped receptacle)
LS plum (swollen and fleshy)
remains of style Nee
pericarp (hard outer layer of
locule epicarp (skin) ‘core’)
seed (‘pip’)
funicle mesocarp (flesh) \ pericarp from one
carpel
testa endocarp (stone).
one seed
a sepal
carpels 3-5 and fused, two seeds per carpel
pedicel
Other examples of drupes are peach, cherry and almond (also Prunus
spp.), elder (Sambucus) and coconut (Cocos nucifera) (fibrous mesocarp). Strawberry
Almonds sold in shops have normally had the epicarp and mesocarp The receptacle becomes swollen and bears on its surface small green fruits
removed. called achenes (‘pips’), each of which contains a single seed.
Section through blackberry — a collection of drupes formed from one flower
separate fruits each from one
carpel LS strawberry
receptacle individual fruits
receptacle (swollen and fleshy)
withered stamens
sepal
remains of style
sepal
epicarp ; LS one fruit
mesocarp | pedicel
remains of style
endocarp (‘pip’)
pericarp locule
LS one fruit (drupe) seed
testa of seed
placenta
Tomato
The tomato is an example ofa fruit called a berry. It resembles a drupe,
but the endocarp is fleshy not stony. (d) Nuts e.g. oak (Quercus), beech (Fagus), chestnut (Castanea) Nuts are
relatively large, dry fruits that do not split open to allow seed dispersal
LS tomato (indehiscent). The whole fruit is dispersed, often as a result of being
fusion of two, three or four carpels collected for food stores by animals, particularly by rodents such as
squirrels.
remains of style
Oak The nut is called an acorn.
epicarp (skin)
mesocarp (flesh) remains of style
endocarp (softer and juicier LS acorn
| than mesocarp) three fused carpels
seed but only one seed
<u funicle
placenta
cupule — cup-
develops
woody pericarp
shaped structure
sepal made from bracts
(modified leaves)
pedicel
Other examples of berries are blackcurrant and gooseberry (Ribes spp.), pedicel
marrow (Cucurbita), orange and lemon (Citrus spp.), banana (Musa),
grape (Vitis), date (Phoenix dactylifera) (in date the single seed is woody).
746
WIND DISPERSAL (a) Parachute mechanism e.g. seeds of willowherb (Epilobium), willow (Salix), cotton (Gossypium hirsutum), fruit of dandelion
(Taraxacum), ‘old man’s beard’ (Clematis)
Dandelion
flower
corolla
stigma Wy appus
structures above 3 } WY rite hairs formed from sepals
white
pappus drop off anthers \ ' BZ after fertilisation, large surface
after fertilisation area aids wind dispersal
pappus | LS fruit
See
ovary
stalk
grows after fertilisation
placenta funicle
small hooks
fruit i Ffusi
(two fused carpels) from inferior Feibl fusion of two carpels
ovary, small and light i eines bse sins ;
scar of attachment to receptacle
(b) Wings e.g. seeds of Pinus (a gymnosperm), fruit of elm (U/mus), ash (Fraxinus),
sycamore (Acer), hornbeam (Carpinus)
long hairs Sycamore (Acer pseudoplatanus)
y seed
two wings formed
from one pericarp
(two carpels) dry and
membranous
Willowherb
scar of style
locule
hairy seeds stacked
in four vertical rows seed inside
prige e LS of right
line of fusion of
line of fusion of testa side of fruit
parpels capsule a fruit carpels finicle ane
splitting into four
pericarp males scar of flower parts
placenta
pedicel pedicel pericarp
One fruit of sycamore Either before or after wind dispersal the fruit separates into two
halves along the line of fusion of the carpels (the fruit is described as schizocarpic); as
(c) Censer mechanism* e.g. poppy (Papaver), love-in-a-mist the fruit falls it rotates and is carried by the wind
(Nigella), foxglove (Digitalis), campion (Lychnis) (all
these examples have fruits called capsules)
__stigmas
Poppy raised ridges above stigmatic
disc (no styles), 4-20 present,
one per carpel pore
stigmatic disc
\ ee _imrele
roof of ovary
pore
placenta
lines of fusion of carpels
pericarp
scars of flower parts
pedicel
LS capsule
which sways in the
The fruit, called a capsule, is borne at the end ofa long pedicel (d) Some plants have extremely small seeds that are light enough to be
at the top of the
wind, causing the numerous small seeds to escape from the pores to distributed by wind without the aid of special appendages to
from side
capsule (* A censer is a vessel in which incense is burnt and is swayed increase surface area, e.g. orchids
side asit is carried)
747
eCiimcrrns AL The most common self-dispersal mechanism involves dehiscent fruits.
Dehiscent fruits ) ;
These dry and break open, often by splitting lines of weakness, e.g. pods. Seeds may be forcibly ejected with
varying degrees of violence or may simply drop out. The seeds released may also be modified for wind
dispersal (e.g. willowherb). Capsules are also dehiscent (see poppy and willowherb under wind dispersal) and
some release seeds violently, e.g. violet (Viola).
Pods e.g. pea (Pisum), broad bean (Vicia), runner bean (Phaseolus), gorse (Ulex) broom and laburnum (Cytisus spp.)
Pods are the characteristic fruits of the legume family (Leguminosae). Pods consist of one carpel containing
many seeds. The pericarp splits along two sides.
Broom, laburnum or pea
line of dehiscence
pericarp
locule two lines of dehiscence

along single carpel
seed
; depression where seed

funicle opposite fits
placenta
remains of style
(see also fig 20. 16 for earlier stages in development of pea pod)
The pericarp twists on drying due to oblique layers of fibrous tissue. As the twisting tension builds up, the
pod may split suddenly along lines of dehiscence, forcibly ejecting some of the seeds. Others are released less
forcibly as twisting continues.
WATER DISPERSAL
Few fruits and seeds are specialised for water dispersal, though water may act as a chance agent of dispersal.
Those that are specialised are made buoyant by structures possessing air cavities. The coconut
(Cocos nucifera) is a drupe (compare plum) whose mesocarp ts fibrous and contains numerous air spaces.
The water lily (Vymphaea) has a spongy outgrowth of its seed (the aril) derived from the funicle.
Water lily
testa
funicle
CHANCE DISPERSAL
The categories of wind, animal and self-dispersal are by no means rigid and the degree of specialisation of
fruits and seeds for particular methods of dispersal is very variable. An element of chance is therefore bound
to be involved in many cases, and more than one ofthe three dispersal methods could be used by a given fruit
or seed. Man is one ofthe chief agents of chance dispersal, with the possibility of seeds clinging to clothing
etc. or being carried in cargoes or by machinery. Contamination of grain crops by weed seeds is a common
phenomenon worldwide. Caches of nuts stored by rodents may be forgotten and germinate the following
spring. Chance floods, hurricanes, etc. may carry seeds further than usual.
748
20.3 Sexual reproduction in Man condition called cryptorchidism) become permanently
sterile and men persistently wearing tight underpants or
20.3.1 The human male reproductive system taking very hot baths may have such a reduced sperm count
The male reproductive system is composed of that it leads to infertility. Whales and elephants are two of
a pair of testes, genital ducts, accessory glands and the the few mammalian species with testes retained in the
penis. The testis is an ovoid-shaped compound tubular abdominal cavity.
gland surrounded by a capsule, the tunica albuginea, The seminiferous tubules are approximately 50 cm long
enclosing about 1 000 highly coiled seminiferous tubules and 200 um in diameter, situated in regions of the testis
embedded in connective tissue containing Leydig cells called testicular lobules. Both ends of the tubule are
(interstitial cells). The tubules of the testis produce the connected to the central region of the testis, the rete testis,
male gametes, spermatozoa (commonly referred to as by short tubuli recti. From here 10-20 vasa efferentia
sperm), and Leydig cells produce the male sex hormone collect sperm and transfer them to the head of the
testosterone. The testes are situated outside the abdominal epididymis where they are concentrated by reabsorption of
cavity in the scrotal sac and, as aresult, temperatures at fluid secreted from the seminiferous tubule. Sperm mature
which sperm develop are 2-3 °C lower than the core in this region of the epididymis before passing 5 m along
temperature. The lower temperature in the scrotal sac is the coiled tubule to the base of the epididymis where they
partly due to its position and partly due to the vascular are stored for a short period before entering the vas
plexus formed by the spermatic artery and vein acting as a deferens. The vas deferens is a straight tube about 40 cm
countercurrent heat exchange system. The dartos muscle of long forming, with the spermatic artery and vein, the
the scrotal sac moves the testes towards or away from the “spermatic cord’ and conveying sperm to the urethra which
body according to the outside temperature in order to traverses the penis. The relationship between these
maintain sperm production at an optimum level. Males structures and the male accessory glands and penis is shown
passing through puberty with undescended testes (a in figs 20.29 and 20.30.
bladder
ejaculatory duct
prostate gland HL y I |
external re) a ee ta
bulbo-urethral gland
penis
vas deferens
internal urethra
epididymis
erectile tissue
| testis
prepuce
scrotal sac
glans be ea Thy
N
Fig 20.29 (above) Diagram of the human male reproductive system
Fig 20.30 (below) Simplified diagram showing the structure

of the human testis and ducts conveying sperm from Gamete formation (gametogenesis)
seminiferous tubules to the urethra 'torarerhes Cells of the germinal epithelium in both male and female
gonads undergo a sequence of mitotic and meiotic
head of epididymis divisions, called gametogenesis to produce mature male
Se
gametes (spermatogenesis) and female © gametes
testicular lobule
(oogenesis). In both cases the process involves three
J’ ) s ast
stages, a multiplication stage, a growth stage, and a
rete testis -- (s XK i vasa efferentia
4 Ly i NAGTB ({ maturation stage. The multiplication stage involves re-
peated mitotic divisions producing many spermatogonia
tubuli recti
wy and oogonia. Each undergoes a period of growth in
preparation for the first meiotic division and subsequent
seminiferous a base of epididymis cytokinesis. This marks the beginning of the maturation
tubule
| stage during which the first and second meiotic divisions
= vas deferens occur followed by differentiation of the haploid cells and
tunica albuginea
the formation of mature gametes.
749
spermatids Leydig cells spermatids
Development of spermatozoa spermatogonia
(spermatogenesis) basement primary
blood capillary
Spermatozoa are produced by a sequence of cell divisions membrane spermatocyte | spermatozoa
called spermatogenesis followed by a complex process of

differentiation called spermiogenesis (fig 20.31). Spermato-
zoa production takes approximately 70 days and 10’
spermatozoa are produced per gram of testis per day. The
epithelium of the seminiferous tubule consists of an outer
layer of germinal epithelial cells and about six layers of cells
produced by repeated cell divisions of this layer (figs 20.32
and 20.33). These represent successive stages in the
development of spermatozoa. Initial divisions of the
germinal epithelial cells give rise to many spermatogonia
which increase in size to form primary spermatocytes.
These undergo the first meiotic division to form haploid
secondary spermatocytes and the second meiotic division to
form spermatids. Between these ‘strands’ of developing
cells are large Sertoli or nurse cells stretching from the
outer layer of the tubule to the lumen.
Fig 20.32 (right) Photomicrographs of a section through

the human testis showing seminiferous tubules and Leydig
(interstitial) cells
Fig 20.31 (below) Summary diagram of the process of
spermatogenesis and spermiogenesis in Man, not drawn to
scale.
Phase of Mechanism of division Type of germ cell Chromosome

development number
germinal epithelium cell 46
mitosis and cell cleavage @) @ @

x©
spermatogonia 46
6
¢ XN
multiplication // repeated divisions *\ SS
@
7
a
growth
©\ primary
spermatocyte
46
@
meiosis I and cell cleavage m
secondary 23
spermatocytes
Hwa
meiosis II and cell cleavage
@ © spermatids 23
BP
: ' (undergoing
spermiogenesis)
maturation
=
spermiogenesis
spermatozoa 23
750
acrosome
head
spermatid (Sym) nucleus
undergoing
spermiogenesis neck = 5
age centriole aa
early spermatid middle alle
(4) piece i mitochondria iL
(5 pm) (arranged as a (b) VS
secondary spiral) sperm
spermatocyte (3)
cell
principal
primary
piece
(45 am) Ht
M1
spermatocyte (2) coke
periphera |
microtubules
spermatogonium spermatogonia central
undergoing mitosis 1) microtubules
end Ee (a) LS human
basement
(5 pm) sperm (c) TS principal piece
membrane
fibrous Fig 20.34 Diagram showing the structure of a mature human

connective tissue spermatozoon
capillary
Fig 20.33 Diagram showing the structure of part of the wall

of a seminiferous tubule and interstitial cells. Cells in various
stages of spermatogenesis and spermiogenesis are shown
and run the entire length of the rest of the spermatozoon

forming the axial filament of the flagellum. The middle
piece is enlarged by the presence of many mitochondria
arranged in a spiral surrounding the flagellum. The energy
released by the mitochondria is used in the contractile
mechanisms which bring about movement of the flagellum.
The principal and end pieces of the spermatozoon are
Spermatocytes become embedded in the many invagina- flagellate and in transverse section show the characteristic
tions in the lateral margins of the Sertoli cells where they arrangement of nine pairs of peripheral microtubules
develop into spermatids before passing to the edge of the surrounding a central pair of microtubules.
cell bordering the lumen where they mature as spermato- Human sperm have a rounded head in plan view and are
zoa. The Sertoli cells are thought to provide mechanical flattened in a lateral view. Activation of the flagellum is
support, protection and nourishment during maturation of described in section 20.3.2, but it is known that flagellar
the spermatozoa. All nutrients, oxygen and waste sub- movement is insufficient to cover the distance from the
stances exchanged between the developing gametes and vagina to the site of fertilisation. The principal locomotory
the blood vessels surrounding the tubules pass through the function of sperm is to cluster around the oocyte and to
Sertoli cells. The fluid carrying spermatozoa through the orientate themselves prior to penetration of the oocyte
tubules is secreted by the Sertoli cells. membranes.
Spermatozoa Endocrine function of the human testis
Spermatozoa are minute, motile male gametes produced The growth, development and maintenance of the testis is
by the male gonads, the testes, and released in millions. controlled by the pituitary gonadotrophins, follicle stimu-
They have various shapes according to species but share a lating hormone (FSH) and luteinising hormone (LH). FSH
common structure. Each spermatozoon is composed of five stimulates the development of spermatozoa and LH
regions as shown in fig 20.34. The head consists of a stimulates the synthesis of the steroid hormone testoster-
nucleus, containing the haploid number of chromosomes, one by the Leydig cells of the testis. LH exerts its influence
covered by a membrane-enclosed structure, the acrosome. on the Leydig cells by releasing membrane-bound cyclic
This contains hydrolytic enzymes which will be involved in AMP (adenosine monophosphate) into the cytoplasm
the penetration of the oocyte by the sperm immediately which then passes to the nucleus where it stimulates the
prior to fertilisation. Functionally, therefore, it may be synthesis of enzymes involved in the synthesis of testoster-
thought of as an enlarged lysosome. The short neck region one from cholesterol. (LH is sometimes referred to as
contains a pair of centrioles lying at right-angles to each interstitial cell stimulating hormone (ICSH) in males,
other. The microtubules of one of the centrioles elongate because of its site of action.)
751
Testosterone is the principal androgenic hormone and it and the medulla is composed of stroma, containing
affects both primary and secondary sexual characteristics. connective tissue, blood vessels and mature follicles (fig
Both testosterone and FSH are required for the successful 20.38).
production of sperm whereas testosterone alone controls The fallopian tube is a muscular tube about 12 cm long
the development of the secondary sexual characteristics and conveys female gametes from the ovary to the uterus.
during puberty and maintains these throughout adult life. The opening of the fallopian tube is expanded and split into
These characteristics include the development of the male fringes or fimbriae which move nearer to the ovary at
external genitalia and the accessory glands of the reproduc- ovulation. The lumen of the fallopian tube is lined with
tive tract, increased muscle development, enlargement of ciliated epithelium and female gametes move towards the
the larynx producing deepening of the voice, the growth uterus aided by peristaltic movements of the muscle wall of
and distribution of hair and behavioural activities associ- the fallopian tube.
ated with mating and parental concern. The uterus is a thick-walled organ about 7.5 cm long and
5 cm wide and composed of three main layers. The outer
covering is called the serous coat and this encloses the
20.3.2 The human female reproductive middle layer, the myometrium, which forms the bulk of the
system wall. The myometrium is composed of bundles of smooth
Female involvement in reproduction is greater muscle cells which are sensitive to oxytocin during birth.
than that of the male and it involves reciprocal interrela- The inner layer, the endometrium, is soft and smooth and
tionships between the pituitary gland, ovary, uterus and composed of epithelial cells, simple tubular glands and
foetus. The female reproductive system is composed of spiral arterioles supplying the cells. The cavity of the uterus
paired ovaries and fallopian tubes, the uterus, vagina and is capable of extending 500 times during pregnancy, that is
external genitalia (fig 20.35). from 10 cm’ to 5 000 cm’. The lower entrance to the uterus
The ovaries are attached to the wall of the body cavity by is the cervix which separates the uterus from the vagina.
a fold in the peritoneum and have the dual function of The vaginal orifice, the urethral orifice and the clitoris are
producing female gametes and secreting female sex protected by two folds of tissue called the vulva composed
hormones. They are about the size and shape of an almond of the labia majora and labia minora. The clitoris is a small
and consist of an outer cortex and inner medulla sur- erectile structure which is homologous with the male penis.
rounded by a connective tissue sheath, the tunica albu- Within the walls of the vulva are the vestibular glands
ginea. The outer layer of cells of the cortex is composed of which release mucus when the female is sexually aroused
germinal epithelial cells from which gamete cells are and this helps to lubricate the penis during intercourse.
produced. The cortex is composed of developing follicles
Development of human ova (oogenesis)

fallopian a
Unlike the production of spermatozoa in males which only
tube
fimbriae
begins at puberty, the production of ova in females begins
uterine cavity
before birth and is completed only after fertilisation. The
stages in oogenesis are shown in fig 20.36. During foetal
development primordial germ cells undergo repetitive
opening of mitotic division and produce many larger cells called
fallopian tube oogonia. These undergo mitosis and form primary oocytes
right ovary oe which remain at prophase of this stage until just before
peritoneal fold ovulation. Primary oocytes are enclosed bya single layer of
body of uterus cells, the membrana granulosa, and form structures known
as primordial follicles. Approximately 2 x 10° of these
labium majorum follicles exist in the foetal female just before birth but only
vulva ; s
labium minorum urethral
opening
about 450 ever develop into secondary oocytes which are
released from the ovary during the oestrous cycle. Prior to
vaginal opening ovulation the primary oocyte undergoes the first meiotic
anus
division to form the haploid secondary oocyte and the first
polar body. The second meiotic division proceeds as far as
metaphase but does not continue until a sperm fuses with
the oocyte. At fertilisation the secondary oocyte undergoes
the second meiotic division producing a large cell, the
Fig 20.35 Simplified diagram showing the human female
reproductive system. The uterus and vagina are shown in ovum, and a second polar body. All polar bodies are small
section, and the external genitalia, urethral and anal openings cells. They have no role in oogenesis and they eventually
are shown in surface view with the labia parted degenerate.
TS2
Phase of Mechanism
development of division
Type of
germ cell
Stage in Chromosome phases as shown in fig 20.37. The events of the menstrual
life number
cycle cycle involve the ovaries (the ovarian cycle) and the
(e) germinal 46 uterus (the uterine cycle) and these are regulated by
~ epithelium
hormones secreted by the ovary which in turn is regulated
by pituitary gonadotrophins.
(e) primordial germ
§ |~ cell Ovarian cycle
:
ies mitosis and (e) ©) In an adult female the ovarian cycle, which is summarised
= cell cleavage | Se in fig 20.38, begins with the development of several primary
S| foetal
E (e) (-) y development follicles (containing primary oocytes) induced by the
} oogonia
release of follicle stimulating hormone (FSH) from the
ae S anterior pituitary gland. Only one of these follicles
continues to grow whilst the rest break down by a
degenerative process known as follicular atresia. The cells
= oe in C) of the membrana granulosa of the follicle proliferate to
5 DEO DOaSe primary oocyte in he produce an outer fibrous layer several cells thick, known as
bb
q rimary follicle
Bas birth the theca externa, and an inner vascular layer, the theca
meiosis | and-—=>== -\~ =-
cell cleavage first polar body
interna. The granulosa cells secrete a fluid which collects
prior to
ovulation
and forms a space, the antrum, within the follicle.
secondary Luteinising hormone (LH) released from the pituitary
oocyte 23
S :
gland stimulates the cells of the thecae to produce steroids,
oS | ovulation
the principal one being 17f-oestradiol. Increasing levels of
S sperm second meiotic division 23
& is initiated by stimulus fertilisation oestradiol during the follicular phase feed back negatively
E meiosis I1_~\“* ee on the pituitary gland causing a decrease in FSH levels in
and 1 second polar body the blood (days 4-11); LH levels remain unchanged.
cell cleavage
Oestrogen levels reach a maximum about three days before
ovum (containing
male and female
ovulation and at this time have a positive feedback action
pronuclei) on the pituitary gland causing the release of both FSH and
LH.
Fig 20.36 Summary diagram of the process of oogenesis in
It is thought that FSH is necessary to stimulate the
Man
growth of follicles but that the continued follicular
development is controlled mainly by LH. The granulosa
Menstrual cycle cells line the periphery with the ovum displaced to one side
of the follicle but still surrounded bya layer of granulosa
In males, gamete production and release is a continuous
cells. This mature follicle, known as the Graafian follicle, is
process beginning at puberty and lasting throughout life. In
about 1 cm in diameter and protrudes from the surface of
females, it is a cyclical activity with a periodicity of
the ovary giving it a warty appearance (fig 20.39). The exact
approximately 28 days and involves changes in the
mechanism of ovulation is unknown but LH, FSH and
structure and function of the whole reproductive system. It
prostaglandins are thought to be involved.
is called the menstrual cycle and can be divided into four
At ovulation the secondary oocyte detaches from the
wall of the follicle, is released into the peritoneal cavity and
(14) ovulation passes into the fallopian tube. Usually only one oocyte is
(LH and FSH)
oocyte released
released each month by one of the ovaries so that ovulation
alternates between the pair of ovaries. The ovulated oocyte
consists of a cell whose nucleus is in metaphase I of meiosis
surrounded by a cell layer known as the zona pellucida and
(7) follicular (FSH, LH and (progesterone) (21)luteal a layer of granulosa cells known as the corona radiata which
phase oestrogen) follicle corpus luteum phase
development develops protects the oocyte up to fertilisation.
Following ovulation, LH levels fall to follicular levels,
(progesterone and in the presence of another gonadotrophin, prolactin,
(5) withdrawn) luteal
menstruation regression
the cells of the ruptured follicle change to form the corpus
luteum (yellow body). This begins to secrete and release
(1) | (28) another female hormone, progesterone, and smaller
amounts of oestrogen. These two hormones maintain the
Fig 20.37 Summary of the main phases of the human _
structure of the endometrium of the uterus and inhibit FSH
menstrual cycle. (Figures in brackets indicate days coinciding
with phases of the cycle) and LH release by negative feedback on the hypothalamus.
753
(aypreas follicle secondary _ thecae antrum
follicle
SSS See granulosa cells
primordial salek2
(Qarkas, follicle
germinal a
Zs iene ae Shee theca
tunica albuginea = SS \\0 membrana granulosa
antrum
st corona radiata
eS
= .
: i zona pellucida
; - 4 we
stroma with blood vessels eX Vo
and connective tissue LZ :
corona radiata
2 ae y & pee
(9) (7) mptured zona pellucida

regressing corpus luteum \O secondary oocyte
active corpus luteum follicle n
Graafia
Fig 20.38 (above) Diagrammatic representation of a section through a human ovary showing the stages in the development
of a Graafian follicle, ovulation and the formation and regression of the corpus luteum. Not all these stages would be seen
together. The numbers indicate the sequence of the stages
Fig 20.39 (below) Photomicrograph of mature human Graafian follicle

4 menstruation
new cycle
at
ay\e
a s
4 ‘ LH
e %
¢ oS
Ae ‘.
Concentrations of ’ ~
FSH and LH in blood a sy
oe Ay
Ae A ‘
FSH—— ca
?
oo ne ‘,
pe =
!
ea
a Sat fiat icesses ‘
sSrasereniis
eo \ pees
Sey ,
_—-— I I I
() A
Time/days
178 oestradiol ~\ progesterone

(oestrogen) 7 ~
Concentrations of 178
oestradiol and
progesterone in blood
4 4 4 4 1 4 {— a 1 4 1 — 4 4 1
eZee
eke 1
St
4 ull
on
a
ae
=
<8 5 9 OND 13 14S tee 17 1S 19-0021 yn, DBS ee OTs) 5 8
| .
Time/days
cece
(-) ovary
y
primary follicle
. .
thecae developing ovulation |
corpus luteum
(a) ares sé V7 OESTROCEN | PROGESTERONE ||
a's
os |
= |
zs | !
1 menstrual 5 proliferative 14 secretory ' 28
Time/days
Fig 20.40 Summary diagram showing the relationship between (a) pituitary gonadotrophins, (b) ovarian steroids, (c) follicle
and corpus luteal development and (d) the thickness of the endometrium during the human oestrous cycle
If fertilisation does not occur factors described later cause levels fall FSH release is no longer inhibited, and as the
the regression of the corpus luteum, which persists as a FSH level rises a new cycle of follicle development occurs.
‘scarred’ area, the corpus albicantus, and the consequent The relationship between the activity of the pituitary gland,
decrease in progesterone and oestrogen levels. As these ovary and uterus is shown in fig 20.40.
tained. two. oocytes each and 2% of the ‘oocytes -

contained two nuclei each. eo thateach
follicle produces a corpus luteum,
7c) (i) at what age did the woman commence
ovulation?
of for approximately ae many years+ would ;
ovulation have continued? —
Gi)how many potential sets of twins could this
- woman have produced and pee a oftwins
_ would they be? —
1What process accounts for the presence of two
nuclei in some of the oocytes (section 2.0)?
755
st
Uterine cycle absence of menstrual bleeding (the ‘period’) is the earlie
sign of pregnancy. (Chorionic gonadotrophin has an
The uterine cycle consists of three phases represented by. interstitial cell stimulating effect on the male foetus
and
structural and functional changes of the endometrium. causes the secretion of testosterone by the foetal testes
These are as follows.
which induces growth of the male sex organs.) The placenta
Menstrual phase. This is the shedding of the begins to assume greater importance in about week 10 of
epithelial lining of the endometrium. Just prior to this pregnancy when it begins to secrete most of the progester-
phase the blood supply to this region is reduced by one and oestrogen essential for a normal pregnancy.
constriction of spiral arterioles in the wall of the uterus as a Premature failure of the corpus luteum before the
result of the fall in the progesterone level in the blood secretory ability of the placenta is established fully is a
following the regression of the corpus luteum. This leads to common cause of miscarriage at about 10-12 weeks of
the death of the epithelial cells. Following the period of pregnancy.
constriction the spiral arterioles then dilate and the During pregnancy HCG may be detected in the urine
increased blood flow detaches the lining of the uterus and and this forms the basis of pregnancy testing. The current
it is shed together with a variable amount of blood in the test used is called the agglutination-inhibition test and
menstrual flow. involves the addition of chorionic gonadotrophin-coated
latex particles to a mixture of urine and an antiserum that
Proliferative phase. This coincides with the will agglutinate with chorionic gonadotrophin. If chorionic
follicular phase of the oestrous cycle and involves the rapid gonadotrophin is present in the urine it will react with the
proliferation of endometrial cells causing the thickening of agglutinating antiserum and not with the latex particles.
the endometrium under the control of oestrogen from the The absence of agglutination of the latex particles indicates
developing follicle. pregnancy and can be used from 14 days after the missed
Secretory phase. During this phase period.
progesterone from the corpus luteum stimulate s the
secretion of mucus from tubular glands and this maintains 20.3.3 Copulation
the lining of the uterus in a receptive state for the
Internal fertilisation is an essential part of
implantation of a fertilised ovum.
reproductive cycles in terrestrial organisms and this is
The effect of failure of fertilisation on the facilitated in many organisms, including Man, by the
oestrous cycle development of an intromittent organ, the penis, which is
inserted into the vagina and releases gametes as high as
If fertilisation does not occur within 24 h of ovulation the
possible within the female reproductive tract. Erection of
secondary oocyte undergoes autolysis in the fallopian tube,
the penis occurs as a result of a local increase in blood
as do any spermatozoa remaining in the female genital
pressure in erectile tissue of the penis, due to para-
tract. The corpus luteum persists for 10-14 days following
sympathetic nervous activity producing constriction of the
ovulation (usually up to day 26 of the cycle) but then it
veins and dilation of the arterioles following some form of
stops secreting progesterone and oestrogen, as a result of
inadequate LH circulating in the blood, and undergoes
sexual excitement. In this state the penis can be inserted
into the vagina where the friction, produced by the
autolysis. Recent evidence has suggested that in some
rhythmic movements of sexual intercourse, increases the
species the wall of the uterus not containing afertilised
ovum produces a factor known as luteolysin which is a tactile stimulation of sensory cells at the tip of the penis.
prostaglandin, prostaglandin F2a. This is thought to pass This activates sympathetic neurones which lead to closure
via the bloodstream to the ovary where it causes regression of the internal sphincter of the bladder and contraction of
of the corpus luteum, by rupturing lysosomes within the the smooth muscle of the epididymis, vas deferens, and the
granulosa cells of the corpus luteum, which then undergoes male accessory glands, the seminal vesicle, prostate and
autolysis. bulbo-urethral glands. This action discharges sperm and
seminal fluids into the proximal (internal) urethra where
The effect of fertilisation they mix to form semen. The increased pressure of these
If fertilisation occurs the zygote develops into a blastocyst fluids in the proximal urethra leads to reflex activity in the
which embeds itself into the wall of the uterus within eight motor neurones supplying the muscles at the base of the
days of ovulation. The outer cells of the blastocyst, the penis. Rhythmic wave-like contractions of these muscles
trophoblastic cells, then begin to secrete a hormone, force semen out through the distal (external) urethra
human chorionic gonadotrophin (HCG), which has a during ejaculation which marks the climax of copulation.
similar function to LH. This function includes prevention The other physiological and psychological sensations
of autolysis of the corpus luteum and the secretion by it of associated with this climax in both males and females are
increased amounts of progesterone and oestrogens which called orgasm. Lubrication is provided during intercourse
cause increased growth of the endometrium of the uterus. partly by a clear mucus secreted by the male bulbo-urethral
Loss of the lining of the endometrium is inhibited and the glands following erection but mainly by glands in the vagina
756
and vulva. The secretions of the male accessory glands
are oocyte after spending several hours in the female genital
alkaline and contain mucus, fructose, vitamin C, citric
acid, tract, usually seven hours, during which time they undergo
prostaglandins and various clotting enzymes, and these
a process known as capacitation. This involves a change in
increase the normally more acidic pH of the vagina to 6-6.5
the properties of the membrane covering the acrosome and
which is the optimum pH for sperm motility following enables fertilisation to proceed. Fertilisation usually occurs
ejaculation. Approximately 3 cm? of semen is discharged
high up the fallopian tube.
during ejaculation of which only 10% comprises sperm. When a sperm reaches the oocyte (fig 20.41), the outer
Despite this low percentage, semen contains about 10° membrane of the sperm covering the acrosomal region and
sperms cm’, the membranes of the acrosome rupture enabling hyalur-
onidase and protease enzymes stored in the acrosome to
20.3.4 Fertilisation ‘digest’ away the cell layers, including the zona pellucida,
surrounding the oocyte. These changes in the sperm head
Sperm are deposited high up the vagina close are known as the acrosome reaction. Subsequent changes in
to the cervix. Investigations have shown that sperm pass the sperm head evert the inner membrane of the acrosome
from the vagina through the uterus and to the top of the allowing penetration of the zona pellucida and the plasma
fallopian tube within five minutes as a result of contractions membrane of the oocyte and, in Man, entry of the entire
of the uterus and fallopian tubes. These contractions are sperm. Once one sperm has entered the oocyte, cortical
thought to be initiated by the release of oxytocin during granules beneath the plasma membrane, beginning at the
sexual intercourse and the local action of prostaglandins, point of sperm entry, rupture, releasing a substance which
present in semen, on the uterus and fallopian tubes. Sperm causes the zona pellucida to thicken and separate from the
are viable in the female genital tract for 24-72 h but are plasma membrane. This is called the cortical reaction and
only highly fertile for 12-24 h. Sperm can only fertilise the spreads over the entire surface of the oocyte causing the
ali
_ ¥ a
Fig 20.41 ning electron micrograph of human sperms clustered around secondary oocyte
(ob) The NE fertilisation ee see penetrates the membrane of a sea-urchin's egg. Nes an Wet Seat
head of the sperm, which contains the genetic code. The grey shape behind itis where energy is release se preY spokeee
for the tail. The sperm has digested the egg’s surface coating of sugary protein and entered. Now the egg s ubat ge
itself to the outside of the sperm, and draws it in to complete the mating. In some unknown way, the entry of one sp
prevents any others getting in, probably because of rapid changes in the egg’s surface coating
TST
corona radiata
chromatin of second meiotic

zona pellucida division
fertilisation membrane
cortical granules
plasma membrane
‘conliie= dheeae oocyte
spermatozoon
maternal chromosomes
second polar body
spindle fibres
female pronucleus
male pronucleus
paternal chromosomes
Fig 20.42 Simplified diagrams illustrating fertilisation. (a) A spermatozoon having undergone capacitation is able to penetrate
the zona pellucida of the secondary oocyte and cause the breakdown of cortical granules. (0) The cortical reaction has
converted the zona pellucida into the fertilisation membrane which moves off the plasma membrane creating the perivitelline
space. (c) The second meiotic division produces the second polar body and is followed by the formation of the female
pronucleus and male pronucleus. (d) The nuclear envelopes of the pronuclei break down and maternal and paternal
chromosomes align themselves along the equator of the ovum attached to spindle fibres. This is metaphase of the first mitotic
division of the zygote
zona pellucida to form an impenetrable barrier called the the cells continue to be retained within the zona pellucida.
fertilisation membrane which prevents the entry of further The cleaved cells are called blastomeres and they form a
sperm, that is preventing polyspermy. wall of cells enclosing a central cavity in the morula, called
The entry of a sperm acts as the stimulus for completion the blastocoel, which fills with liquid from the oviduct. The
of the second meiotic division of the secondary oocyte outer layer of blastomeres is called the trophoblast and this
which produces the ovum and the second polar body. The differentiates at one point to form a thickened mass of cells,
second polar body immediately degenerates, and the tail of the inner cell mass. This stage is called the blastocyst and is
the sperm is lost within the cytoplasm of the ovum. Sperm reached about 4-5 days following ovulation. The structure
and ovum nuclei form pronuclei which are drawn together. of the blastocyst is shown in fig 20.43.
The membranes of the pronuclei break down and the When the blastocyst arrives in the uterus it spends about
paternal and maternal chromosomes attach to spindle two days in the lumen during which the zona pellucida
fibres which have formed. Both haploid sets of chromo- gradually disappears enabling the cells of the trophoblast to
somes by this stage have undergone replication and align make contact with the cells of the endometrium. The
themselves as 46 pairs of chromatids along the equator of trophoblast cells multiply in the presence of nourishment
the spindle as in metaphase of mitosis. This fusion of from the endometrium and between the sixth and ninth
pronuclear chromosomes is termed fertilisation. At this days after ovulation the blastocyst becomes embedded
point the diploid number of chromosomes is restored and
the fertilised ovum is now called a zygote.
Anaphase and telophase of the zygote cell follow and
complete the first mitotic division of the zygote. The zygote trophoblast
then undergoes cytokinesis and produces two diploid (each cell is a blastomere)
daughter cells. The process of fertilisation is summarised in
fig 20.42. blastocoel
20.3.5 Implantation
inner cell mass
As the zygote passes down the fallopian tube it
cleaves by successive nuclear and cell divisions to produce a
collection of cells called the morula. Cleavage at this stage Fig 20.43 Simplified diagram of a human blastocyst four
does not result in an increase in size of the morula because days after ovulation
758
> uterine venule
eroded uterine venule

trophoblastic villi
uterine wall
(formed from outer layer of trophoblast)
Re = y—\\ lacunae (blood spaces)
—_= embryonic disc
eos)ee gf nn cavity
blastocoel
inner layer of trophoblast

cavity of uterus
outer epithelium of uterus wall
within cells of the endometrium. This process is called Fig 20.44 (above) Simplified diagram showing a recently
implantation. The cells of the trophoblast differentiate into implanted human blastocyst in the endometrium of the uterus.
two layers and the cell membranes of cells of the outer layer Enzymes produced by the syncitium of the outer trophoblast
of the trophoblast break down to form the trophoblastic break down the blood vessels of the endometrium producing
villi which grow into the endometrium (fig 20.44). The lacunae containing blood which are used in the nourishment
and excretion of the blastocyst
areas of the endometrium between these villi form
interconnecting cavities, called lacunae, which give this
region of the endometrium a spongy appearance. Hydro-
lytic enzymes released by this multinucleate structure cause trophoblastic villi
the arterial and venous blood vessels in the endometrium to
break down and blood from them fills the lacunae. In the
early stages of development of the blastocyst, exchange of chorion bees chorionic
villi
nourishment, oxygen and excretory materials between the
cells of the blastocyst and the maternal blood in the uterus amnion
wall occurs through the trophoblastic villi. Later in
development this function is taken over by the placenta. amniotic cavity placenta
will
develop
embryo in this
20.3.6 Early embryonic development region
The outer cells of the blastocyst, the tropho- allantois

blast, grow and develop into an outer membrane called the yolk sac 4
chorion which plays a major role in nourishing and chorion

removing waste from the developing embryo. Two cavities
Fig 20.45 Simplified diagram showing the relationship
appear within the inner cell mass and the cells lining these between the human embryo and extra-embryonic membranes
give rise to two membranes, the amnion and the yolk sac. about five weeks after ovulation. The area of the allanto-
The amnion is a thin membrane covering the embryo and chorion becomes the placenta
has a protective function. Secretion of amniotic fluid by the
cells of the amnion fill the amniotic cavity between the stage the mesoderm is formed and these three germ layers
amnion and the embryo. As the embryo increases in size give rise to all the tissues of the developing embryo as
the amnion expands so that it is always pressed up against described in section 21.8.
the uterus wall. The amniotic fluid supports the embryo During the early stages of embryonic development
and protects it from mechanical shock. The yolk sac has no exchange of materials between embryo and mother across
significant function in Man but is important in reptiles and the trophoblastic villi is adequate, but soon a fourth
birds for the absorption of food from the yolk and its membrane, the allantois, develops from the embryonic
transfer to the midgut of the developing embryo. hindgut. The chorion, amnion, yolk sac and allantois are
The cells of the inner cell mass, beneath the early called extra-embryonic membranes, or foetal membranes
amnion, and the yolk sac form a structure called the (fig 20.45). The allantois grows outwards until it comes into
embryonic disc, which gives rise to the embryo. The cells of contact with the chorion where it forms a richly vascular-
the disc differentiate at an early stage (when the diameter is ised structure, the allanto-chorion, which contributes
less than 2 mm) and form an outer layer of cells, the towards the development of a more efficient and effective
ectoderm, and an inner layer, the endoderm. At a later exchange structure, the placenta.
139
venous
Placenta by the difference in pressure between arterial and
vessels.
The placenta is a temporary organ found only in eutherian
The cell membranes in the wall of the chorionic villi
mammals and is the only organ in animals composed of
bathed by maternal blood bear microvilli, which increase
cells derived from two different organisms, the foetus and
their surface area, for the exchange of substances by
the mother. It is a point of close association between
diffusion and other methods of transport. Numerous
maternal and foetal circulations and facilitates the transfer
mitochondria are found in these cells whose membranes
of nutrients, oxygen and metabolic waste products between
contain carrier molecules used in the uptake of materials
foetus and mother. The placenta is a discrete disc-shaped
into the villi by active transport. The presence of numerous
structure localised in one region of the uterus wall and as it
small vesicles within the cells of the villi suggests that
develops it takes over from the trophoblastic villi as the
materials are taken up from the maternal body by
principal site of exchange of materials after 12 weeks.
pinocytocis (section 7.2.2). A number of mechanisms of
The foetal part of the placenta consists of connective
uptake are necessary since the distance between foetal and
tissue cells of the chorion which invade the trophoblastic
maternal blood is large, for example ten times that across
villi in one region of the uterus wall and produce larger
the alveolar membranes of the lung. Water, glucose, amino
projections called chorionic villi. The inner regions of the lipids, mineral salts, vitamins,
acids, simple proteins,
chorionic villi become invaded with looped capillary
hormones, antibodies and oxygen pass from mother to
networks derived from two blood vessels of the foetus, the other nitrogenous waste
foetus, and water, urea and
umbilical artery and umbilical vein. These blood vessels are
materials, hormones and carbon dioxide pass from foetus
derived from the allantois and run between foetus and
to mother across the ‘placental barrier’. Potentially
uterus wall in the umbilical cord which is a tough structure
harmful substances such as bacteria, viruses, toxins and
about 40 cm long covered by cells derived from the amnion
and chorion (fig 20.46). drugs can pass to the foetus, but this is offset by certain
antibodies, globulins, antibiotics and antitoxins passing in
The maternal part of the placenta is composed of
the same direction. This ensures that the baby is born with
outward projections of the outer layers of the endomet-
passive immunity (section 14.14.5) to certain diseases.
rium, the decidua. Between these and the chorionic villi are
The placental barrier not only protects the foetus from
lacunae supplied with arterial blood from the uterine
many harmful situations which may occur to the mother but
arterioles and drained by venules of the uterine vein. The
direction of blood flow through the lacunae is determined also shields the foetal circulation from the high blood
deciduous endometrium
placenta
permanent endometrium
myometrium
maternal blood ihe S|
( : — IB chorionic villus
é TO FOETUS
os ae i Sou eed eagl
us
ePies
umbilical arteries
IG) =: “PS
lacuna’ : eb chorionic capillaries
D D ate
TE A OS. es
: Ik
Ga a Se ae lacuna hel fe
=n =\ B sills
PORPIDSy) epee:
umbilical vein acune :
amniotic cavity —
v “lacuna ?=)° '
nae ee val te ao
amnion a ig) EL uterine venule
oc OM Arce oo uterine arteriole
foetal arteries . Me
Se) ;
point of detachment
foetal veins chorion of endometrium
after birth
Fig 20.46 Simplified diagram showing the relationship !between

E umbilical blood vessels, : capillari es of the chorion
ionic vil
blood spaces of the lacunae of the human placenta. This structure forms the link between the circulatory Risiaite rfthafeet
and the mother
760
pressure of the maternal circulation. It cannot function, genetic sex Xx Y
however, as an immunological barrier, and since the foetus yXx Xx
i
carries paternal genes it will produce antigens foreign to the
mother who will produce antibodies against them. The gene Tfm Y-linked testis- / Tfm
mechanisms accounting for the remarkable ability of the determining gene ye
Vee
/
foetus (or, in immunological terms, the homograft) to resist
rejection for the 40 weeks of gestation is not known but is synthesised androgen H-Y antigen / androgen
protein receptor Le receptor
thought to involve the production of immune suppressive protein protein
substances which circulate in the maternal plasma.
embryonic
The continual passage of oxygen from mother to foetus is genital ridge x
vital to the life and development of the foetus and this is
ensured by the difference in affinity for oxygen between x
<
foetal and maternal haemoglobins as described in section embryonic
14.13.1. gonad
The placenta is an endocrine organ whose major /
secretions are chorionic gonadotrophin, oestrogens, testosterone
progesterone and human placental lactogen. The latter

androgen receptor/
hormone stimulates mammary development in preparation testosterone complex
for lactation. The site of secretion of all these hormones is
the connective tissue of the chorion. rudimentary male/female male/female
reproductive
Sexual development in the embryo system
The genetic sex of the embryo is determined at fertilisation

foetus male female
by the sex chromosomes carried by the father’s sperm, X in
the case of a female and Y in the case of a male. Despite Fig 20.47 Summary diagram showing the events involved in
this, it would appear that the basic disposition of the human the differentiation of the embryonic genital ridge and
body is towards being female, largely as a result of the rudimentary reproductive system into the specific gonad and
reproductive system of the foetus
presence of an X chromosome in both sexes. In the early
stages of embryonic development a pair of undifferentiated
embryonic gonads, the genital ridges, and both rudi- and do not develop. In an XX embryo, the absence of
mentary female and male reproductive systems develop in testosterone allows the reproductive system to develop in
the embryo. As a result of this, all embryos are potentially its inherent direction towards that of female.
bisexual up to the sixth week of development. Thus it may be concluded that placental influences will
Recent investigations have revealed a possible mechan- direct the development of the embryo in the direction of a
ism whereby the sex chromosomes determine which of female unless diverted by a mechanism initiated by the Y
these systems is activated and lead to the phenotypic chromosome. A summary of these events is shown in fig
expression of the embryo’s sex.
20.47.
The X chromosome carries a gene, the Tfm gene
20.3.7 Birth
(testicular feminisation gene) which specifies the produc-
tion of an androgen-receptor protein molecule in the cells of From the beginning of the third month of
the developing reproductive system. Since both male and pregnancy the human embryo is referred to as the foetus
female embryos carry at least one X chromosome, this and it normally completes a total of 40 weeks of
molecule is present in both sexes. development, the gestation period, before birth occurs.
The Y chromosome carries a gene called the Y-linked Most of the major organs are formed by the twelfth week of
testis-determining gene specifying the production of a pregnancy and the remainder of the gestation period is
protein molecule, the H-Y antigen which stimulates the taken up by growth.
cells of the embryonic genital ridges to differentiate into Throughout pregnancy oestrogen and progesterone are
seminiferous tubules and interstitial cells. Testosterone secreted in progressively greater amounts, first by the
released into the embryonic circulatory system reacts with corpus luteum and then principally by the placenta. In the
the androgen-receptor molecules in the target cells of the last three months of pregnancy oestrogen secretion
potential reproductive system. The androgen-receptor/ increases faster than progesterone secretion and, im-
testosterone complex formed passes to the nuclei where it mediately prior to birth, the progesterone level declines
activates genes associated with the development of the and the oestrogen level increases. The functions of these
tissues. Testosterone will activate only those tissues which hormones in pregnancy are summarised in table 20.4.
give rise to the male reproductive system and therefore an It was thought that hormonal activities within the mother
XY embryo will develop into a male foetus. The tissues of controlled the timing of birth but recent evidence obtained
the potential female reproductive system are not activated from research on several mammals has suggested there is a
761
Table 20.4. Summary of major functions of human oestrogen progesterone production and an increase in secretion of
and progesterone during pregnancy prostaglandins. The reduction in progesterone level allows
eee the maternal posterior pituitary gland to release the octa-
Oestrogen Progesterone peptide hormone, oxytocin, and removes the inhibitory
effect on contraction of the myometrium. Whilst oxytocin
Growth of mammary glands Growth of mammary glands causes contraction of the smooth muscle of the myomet-
Inhibits FSH release
Inhibits FSH release
Inhibits prolactin release rium, prostaglandins increase the power of the contrac-
Inhibits prolactin release
Prevents infection in uterus Inhibits contraction of tions. The release of oxytocin occurs in ‘waves’ during
Increases size of uterine muscle myometrium ‘labour’ and provides the force to expel the foetus from the
cells uterus. The onset of contractions of the myometrium,
Increases ATP and creatine so-called ‘labour pains’, are accompanied by the dilation of
phosphate formation the cervix, the rupture of the amnion and chorion releasing
Increases sensitivity of of
myometrium to oxytocin amniotic fluid from the cervix, and the stimulation
Nee
———— eee ee ee eee eee eee stretch receptors in the walls of the uterus and cervix. The
latter activate the autonomic nervous system and auton-
high degree of foetal involvement in the timing of birth. omic reflexes induce contraction of the uterus wall. Other
This would seem to have profound adaptive survival value impulses pass up the spinal cord to stimulate the hypothala-
in ensuring birth of the foetus at a stage in development at mus to release oxytocin from the posterior pituitary gland.
which it can lead a relatively independent existence. The The pressure of the head of the foetus ‘engaged’ in the
initial stages of birth are believed to result from stimuli, as pelvis pressing against the cervix with the face towards the
yet undefined, influencing the foetal hypothalamus to mother’s anus, irritates the cervix and leads to stronger
release ACTH from the foetal pituitary. One area of contractions of the myometrium.
current thought concerning the initial birth stimuli is that of Uterine contractions spread down over the uterus and
‘foetal stress’ brought about by an immunological rejection are strongest from top to bottom, thus pushing the baby
of the mature foetus by the tissues of the mother. As foetal downwards. Throughout these contractions the cervix
ACTH is released it stimulates the foetal adrenal gland to gradually dilates and the time between bouts of contrac-
release corticosteroids which cross the placental barrier tions decreases. This is the first stage of labour and it ends
and enter the maternal circulation causing a decrease in when the cervix has the same diameter as the head. The
maternal
hypothalamus and
posterior pituitary
oxytocin (6)
decreased increased adrenal cortex

progesterone prostaglandin
output (5) output (7)
(9a)
autonomic (9a)
) sensory spinal cord
and motor
placenta
Unknown initial stimuli (1)

‘foetal stress’?
immunological rejection?
mature foetal hypothalamus?
mature foetal adrenal gland?
dilation of cervix
Fig 20.48 Possible mechanisms associated with birth. : Of all the hormonal and nervous mechani isms shown above, onl
effect of oxytocin on the uterus wall has been clearly demonstrated. Numbers 1—7 show the possible sequence of Berta i
inducing contraction of the uterus wall and 8, 9a and 9b show the possible reflex pathways involved in the control of the
contractions
762
9
Fig 20.49 (a) (above) The baby’s head has been guided
through the vagina by the doctor attending the birth and it is
now fully emerged. (b) (right) The body of the baby is now
partially emerged. The legs are still inside and the umbilical
cord remains attached. (c) (below) The baby has been
cleaned and is resting on the mother. At this stage the
umbilical cord is intact and the baby is still attached to the
placenta
SAS
NOR
S
Fig 20.49 (d) The doctor is now cutting the umbilical cord and the process of birth will then be complete
second stage of labour involves the passage of the head and

body through the vagina and the delivery of the baby. The pectoralis muscle
umbilical cord is ligatured in two places close to the baby
and a cut made between the ligatures allowing the baby to
be now totally separated from any immediate physiological alveoli of mammary gland
reliance on the mother. Within 10-45 minutes after birth
the uterus contracts dramatically and separates the
placenta from the wall of the uterus and the placenta then mammary duct
passes out through the vagina. This is the third stage of
labour. Bleeding, throughout this period, is limited by mammary sinuses
contraction of smooth muscle fibres which completely
surround all uterine blood vessels supplying the placenta.
Average blood loss is kept to about 350 cm’. + openings of ducts on nipple
20.3.8 Lactation pigmented areola
The mammary glands, or breasts, consist of supporting stroma

two types of tissue, glandular tissue and supporting tissue
or stroma. Glandular epithelial cells line small sacs called adipose tissue
alveoli arranged in 15 lobules within the breast. The alveoli
are surrounded bya layer of myoepithelial (contractile)
tissue involved in the release of milk. Milk is produced by i
the glandular cells and passes, via a series of ducts and Fig 20.50 Simplified diagram of the human female breast
sinuses which store milk, to separate openings in the showing the glandular alveoli where milk is secreted and the
nipple. ducts and sinuses conveying milk to the nipple.
764
During puberty the breasts enlarge due to development ductus arteriosus
of the stroma under the influence of oestrogen and
progesterone. Internal changes in the nature of the stroma | \
and secretory ducts, and in the amount of fat in the breasts, | |
occur throughout the menstrual cycle, enlarging slightly | |
Lecgtiecoi Pro vcccere +) LUNGS — Jesseerens,,
during the luteal phase, but during pregnancy there is |
:__| pulmonary
greater increase in size and activity of the breasts. foramen ovale
The increase in the size of the breasts during pregnancy is

pulmonary
due to the development of the secretory areas under the artery
influence of oestrogen, progesterone, corticosteroids,
growth hormone, placental lactogen and _ prolactin.
However, throughout ‘pregnancy the presence of
progesterone inhibits the formation of milk (lactogenesis). inferior vena
cava
At birth, when the progesterone level falls, prolactin is no
longer inhibited and it stimulates the alveoli to secrete
milk. TISSUES
Human milk contains fat, lactose (milk sugar) and the
proteins lactalbumin and casein which are all easily
digestible. The milk is synthesised from metabolites
ductus venosus {
circulating in the blood, such as lactose from glucose under
A;___hepatic portal vein
the influence of the enzyme lactose synthetase, protein
from amino acids, and fats from fatty acids, glycerol and XN 73
> Guile codecs
acetates. This food alone is adequate to produce weight | |
gains in the baby of 25-30 g per day. : }—_umbilical
umbilical vein | artery
In between breast feeds prolactin stimulates milk \ ]
production for the next feed. Oestrogen is necessary for the \—4 PLACENTA }——~—-~7
——— ante-natal only
continued production of milk, but artificially induced high
ccccccces post-natal only
levels of oestrogen have been used to inhibit lactogenesis in
ante- and post-natal
mothers not wishing to breast feed. (It is common practice
now to use a drug called bromocriptine instead of Fig 20.51 Simplified diagram showing ante-natal (foetal) and
oestrogen.) post-natal circulatory systems. (The periods when the blood
The ejection of milk from the nipple involves a simple vessels are functional are indicated on the diagram.)
reflex action, the milk ejection reflex. The sucking of the
baby on the breast stimulates sensory receptors in the here most of the blood passes through an opening in the
nipple to set up impulses passing via the spinal cord to the atrial septum, the foramen ovale, into the left atrium. Some
hypothalamus which releases oxytocin from the posterior blood passes from the right atrium into the right ventricle
pituitary gland. This causes contraction of the myoepithe- and into the pulmonary artery but does not pass to the
lial tissue surrounding the alveoli and forces milk through lungs. Instead it is shunted via the ductus arteriosus directly
the ducts and sinuses and out of the nipples. to the aorta thereby by-passing lungs, pulmonary vein and
The initial secretion of the breasts, following birth, is not the atrium and ventricle of the left side of the heart. Blood
milk but colostrum. This has a yellow colour and contains from the left atrium passes into the left ventricle and into
cells from the alveoli and is rich in the protein, globulin, but the aorta which supplies blood to the head, upper limbs,
low in fat. It is believed to be a means of passing antibodies, trunk and lower limbs and the umbilical artery. Pressure in
particularly IgA (section 14.14.3), from mother to baby. the foetal circulatory system is greatest in the pulmonary
artery and this determines the direction of blood flow
through the foetus and placenta.
20.3.9 Changes in foetal circulation at birth
Throughout development in the uterus the eee
foetal lungs and digestive tract do not function, since gas- the foetal circulationifbloodpressure were
eous exchange and nutrition are provided by the mother
via the placenta. Most of the oxygenated blood returning to
the foetus via the umbilical vein by-passes its liver in a At birth the sudden inflation of the lungs reduces the
vessel, the ductus venosus, which shunts blood into the in- resistance to blood flow through the pulmonary capillaries
ferior vena cava and passes it to the right atrium (fig 20.51). and blood flows through them in preference to the ductus
Some blood from the umbilical vein flows directly to the arteriosus; this reduces the pressure in the pulmonary
liver; blood entering the right atrium, therefore, contains a artery. Simultaneously the tying of the umbilical cord
mixture of oxygenated and deoxygenated blood. From prevents blood from flowing through the placenta, and this
765
increases the volume of blood flowing through the body of 20.4 Phylogenetic review of sexual
the baby and leads to a sudden increase in blood pressure in reproduction in vertebrates
the aorta, left ventricle and left atrium. This pressure
change causes the small valves guarding the foramen ovale, Sexual reproduction occurs in every animal
which open to the left atrium, to close, preventing the phylum and, whilst the basic processes of gamete forma-
short-circuiting of blood from right to left atrium. Within a tion, fertilisation and zygote formation are common to all
few months these valves fuse to the atrial septum and close groups, there is considerable variation in the anatomical,
the foramen ovale completely. If this does not occur the physiological and behavioural aspects of sexual reproduc-
baby is left with a ‘hole in the heart’ and will require surgery tion. Many examples of the variations shown by the
to correct the defect. invertebrate phyla are described in chapter 4.
The increased pressure in the aorta and decreased In vertebrates, too, tremendous variation exists in many
pressure in the pulmonary artery forces blood backwards of the aspects of sexual reproduction, and some of these,
along the ductus arteriosus into the pulmonary artery and together with selected examples, are summarised in table
hence to the lungs, thereby boosting its supply. After a few 20.5. The phylogeny of the vertebrates shows a gradual
hours muscles in the wall of the ductus arteriosus constrict adaptation to life on land. One of the major problems to
under the influence of the rising partial pressure of oxygen overcome in making the transition from an aquatic
in the blood and close off this blood vessel. A similar existence to a terrestrial existence involved reproduction.
mechanism of muscular contraction closes off the ductus The majority of fish shed their gametes directly into water,
venosus and increases blood flow through the liver. The fertilisation is external, eggs contain a considerable amount
mechanism of closing down the ductus venosus is not of yolk, larval stages are common and any degree of
known but is essential in transforming the ante-natal parental care is rare. In the amphibia there are several
(before birth) circulation into the post-natal (after birth) examples of adaptations to terrestrial life but few of these
condition. involve mechanisms of reproduction. Amphibia have to
Table 20.5. Summary table showing variations in the mechanisms of sexual reproduction in selected vertebrates
ee
Organism Number of female Diameter Site of Site of embryonic Degree of parental care
gametes released of egg/mm fertilisation development
per year nneee EEE En nnEEEnnn SSE
Cod 3-7 x 10° 0.13 external Sea as larvae, None, very little yolk in egg
oviparous*
Stickleback 60 2.0 external Nest, built by male, Male occupies territory, courts
oviparous female; eggs laid and fertilised in
nest; young protected in nest
Spotted 144 15.0 internal Egg case, ‘Mermaids Mating occurs near surface of water,
dogfish purse’; no larval large supply of yolk in egg
(Scyliorhinus stage, Oviparous
canaliculus)
Frog 2 000 Z external Water, collectively Simple courtship, mating occurs;
(Rana as ‘frog spawn’, Moderate amount of yolk in egg;
temporaria) oviparous; larval no after-care
(tadpole) stage under-
goes metamorphosis
Sand lizard 6 8.0 internal Amniote egg in Simple courtship, mating occurs;
(Lacerta holes in sand or soil, large amount of yolk; no after care
agilis) Oviparous
Robin > 20.0 internal Amniote egg in nest Male occupies territory, elaborate
(Erithacus built by female, courtship, mating occurs, female
rubecula) Oviparous incubates eggs for 14 days, fledgling
fed by both parents, offspring
independent after six weeks
*Offspring /aid or spawned as eggs.
766
return to water to mate, and the early stages of their
development take place there also. There are, however, co oT
RLS
shell
many amphibian species that show elaborate behavioural WA
patterns associated with parental care. For example, the XY chorion
iA
0
male Pipa toad spreads the fertilised eggs over the back of
4 amnion
the female where they stick, become ‘embedded’ in the E]
skin and develop into tadpoles. After about three weeks a

4 i amniotic cavity
they escape from the mother’s back and lead an indepen- Lf E
dent existence. embryo

Reptiles were the earliest group of vertebrates to
overcome the problems of fertilisation and development on allantois
land. Clearly shedding of gametes ina terrestrial situation
is impossible, so the first requirement of totally land- Lt yolk sac
dwelling organisms must have been the introduction of
male gametes into the female body, that is internal
fertilisation. Internal fertilisation occurs in reptiles and the Fig 20.52 Simplified diagram of the amniote egg
increased chances of fertilisation reduces the numbers of
gametes which it is necessary to produce. Once fertilised,
the zygote develops within a specialised structure, the embryo can develop. All reptiles, birds and mammals have
amniote (cleidoic) egg, which provides’the embryo with a an amnion and are called amniotes. As the embryo grows
fluid-filled cavity in which it can develop on land. The outer the amnion is pushed out until it fuses with the third
shell provides immediate protection from mechanical embryonic membrane, the chorion, which lies immediately
damage and envelops the four membranes which surround within the shell and prevents excessive water loss from the
the embryo. These four extra-embryonic membranes, the amnion. The allantois is an outgrowth of the embryonic
yolk sac, amnion, chorion and allantois are derived from hindgut and rapidly expands, in reptiles and birds, to
ectoderm, endoderm and mesoderm and provide the underlie the chorion. Here it functions primarily as a
embryo with protection and facilitate many of its metabolic ‘bladder’ for storing excretory products and as the gas
activities including nutrition, respiration and excretion. exchange organ of the embryo, facilitating the transfer of
The yolk sac develops as an outgrowth of the embryonic gut respiratory gases between the environmental atmosphere
and encloses the yolk which is gradually absorbed by the and the amniotic fluid via the porous shell. The structure of
blood vessels of the yolk sac. When the yolk has been used the amniote egg is shown in fig 20.52.
up the yolk sac is withdrawn into the gut. The amnion forms Birds and primitive egg-laying mammals called mono-
from an upgrowth of the cells beneath the embryo and tremes, such as the duck-billed platypus, Ornithorhyncus,
completely encloses the embryo in the amniotic cavity all produce an amniotic egg and, whilst the shell of the egg
which becomes filled with amniotic fluid secreted by the is lost in higher mammals, the four extra-embryonic
cells of the amnion. This provides the embryo with an membranes are retained; two of them, the chorion and
immediate fluid environment (a replica of the ancestral allantois, give rise to the placenta in eutherian (placental)
aquatic environment of the amphibian) in which the mammals (section 20.3.6).
767
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Chapter Twenty-one
Growth and development
Growth is a fundamental characteristic of all living growth. Growth can therefore be defined as an irreversible
organisms. It is often thought of simply as an increase in increase in dry mass of protoplasm. This in turn reflects an
size, but careful consideration shows this to be an increase in the amount of protein which has been
inadequate definition. For example, the size of a plant cell synthesised, and the fact that the process of protein
may increase as it takes up water by osmosis, but this synthesis forms the basis of growth.
process may be reversible and cannot then be thought of as Growth may be positive or negative. Positive growth
genuine growth. Also, during cleavage of the zygote and in occurs when anabolism exceeds catabolism, whereas
the early embryo there is an increase in cell numbers negative growth occurs when catabolism exceeds anabol-
without an increase in size (volume or mass). This is the ism (chapter 11). For example, in the course of germination
result of cell division without subsequent increase in size of of a seed and the production of a seedling various physical
daughter cells. The process is a developmental one and so parameters increase in magnitude, such as cell number, cell
perhaps should be regarded as growth despite the fact that size, fresh mass, length, volume and complexity of form,
no increase in size occurs. whilst others such as dry mass may actually decrease. From
The process of development is so closely linked with the definition, germination in the latter case is therefore
growth that the phrase ‘growth and development’ is strictly a time of negative growth.
commonly used to describe the processes which are
normally thought of as growth. Starting with an individual
cell, growth of a multicellular organism can be divided into 21.1 Measurement of growth
three phases:
(1) cell division (hyperplasia) — an increase in cell number Growth occurs at many levels of biological
as a result of mitotic division and cell division; organisation from the community level down to the
(2) cell expansion (hypertrophy) — an irreversible increase molecular level. In all cases if the increase in measurable
in cell size as a result of the uptake of water or the parameter is plotted against time an S-shaped growth curve
synthesis of protoplasm; is obtained. Fig 21.1 shows a variety of growth curves
(3) cell differentiation — the specialisation of cells; in its produced by plotting different parameters such as length,
broad sense, growth also includes this phase of cell height, mass and surface area, volume and numbers against
development. time. The shape of these curves is described as sigmoid,
However, each of these processes can occur at separate meaning S-shaped.
points in time. The example of cleavage has already been A sigmoid curve can be divided into four parts. The
mentioned above. An increase in volume without change in initial phase is the lag phase during which little growth
cell numbers may also occur, as in the region of cell occurs. This leads into the second phase, the log phase
elongation behind the root and shoot tips of higher plants. (grand period of growth) during which growth proceeds
In the case of single-celled organisms such as bacteria, cell exponentially. During this phase the rate of growth is at its
division results in reproduction (not growth) of the maximum and at any point the rate of growth is
individual and growth of the population. proportional to the amount of material or numbers of cells
All stages of growth involve biochemical activity. or organisms already present.
Protein synthesis is particularly important since it is the In all cases of growth the exponential increase declines
means by which the DNA message is expressed in terms of and the rate of growth begins to decrease. The point at
enzymes synthesised by the cell. Enzymes control cell which this occurs is known as the inflexion point. The third
activities. Changes at the cell level bring about changes in phase is the decelerating phase (self-retarding phase)
overall form and structure, both of individual organs and of during which time growth becomes limited as a result of the
the organism as a whole, and this process is known as effect of some internal or external factor, or the interaction
morphogenesis. of both. The final phase is the plateau phase or stationary
A definition of growth should satisfy the criterion of phase. This usually marks the period where overall growth
increase in size that occurs in all organisms from single- has ceased and the parameter under consideration remains
celled organisms to the most advanced plants and animals, constant. The precise nature of the curve during this phase
as well as reflecting the metabolic activity associated with may vary depending on the nature of the parameter, the
769
female Daphnia
D
oO
- S S
s
CS
o.
Length/mm
S«3 /
/ us
Zea mais
a 7&
2.0
aan
1.0 aes x
i asl a a el | |i | |
| L | ee | | | 100 120
30 40 50 60 70 (b) 20 40 60 80
(a) 10 20
Time/days Time/days
1.2b
S 0 Pinus sylvestris
oS nN
Sap
Sp
volume/m?
Mean
organisms
of
Number
ie —L i 0 BRE ee |
50 100 150 200 (a) 0 4 8 12 16
(¢)
Time/years Time/h
Fig 21.1 (above) Growth curves obtained using six different parameters and four different species. In all cases the curves are
sigmoid. ((b) After Kreusler. (c) Indicates growth of trees, after Tischendorf. (d) After Knopf.)
Fig 21.2 (below) A typical sigmoid growth curve showing the

four characteristic growth phases and the inflexion point. (a) 21aisl Methods of measuring growth
Lag phase; (b) log phase; (c) inflexion point; (d) decelerating
phase; (e) plateau or stationary phase Growth can be measured at various levels of
biological organisation, such as growth of a cell, organism
or population. The numbers of organisms in a population at
different times can be counted and plotted against time to
produce a population growth curve as shown in fig 21.1d
| (section 12.6.3). At the level of the organism there are a
|
|
|
variety of parameters which may be measured; length,
|
|
area, volume and mass are commonly used. In plants,
Growth
parameter |
growth curves for roots, stems, internodes and leaf area are
often required, and length and area are the parameters
@ O) C.@.,... i.
+»
4W______»-4+______»><t> chosen. In the case of growth in animals and entire plants
Time
length and mass are two commonly measured parameters.
species and internal factors. In some cases the curve may With regard to mass there are two values that can be used,
continue to increase slightly until the organism dies as is the namely fresh (wet) mass and dry mass. Of the two, fresh
case with monocotyledonous leaves, many invertebrates, mass is the easier parameter to measure since it requires
fish and certain reptiles. This indicates positive growth. In less preparation of the sample and has the advantage of not
the case of certain coelenterates the curve flattens out causing any injury to the organism, so that repeated
indicating no change in growth whilst other growth curves measurements of the same organism may be taken over a
may tail off indicating a period of negative growth. The period of time.
latter pattern is characteristic of many mammals, including The major disadvantage of using fresh mass as a growth
Man, and is a sign of physical senescence associated with parameter is that it may give inconsistent readings due to
increasing age. fluctuations in water content. True growth is reflected by
770
changes in the amounts ’of constituents other than water could be obtained by calculating the slope of the absolute
and the only valid way to measure these is to obtain the dry growth curve ($) at various points and plotting these
mass. This is done by killing the organism and placing it in
against time).
an oven at 110 °C to drive off all the water. The specimen is This curve shows how the rate of growth changes during
cooled in a desiccator and weighed. This procedure is the time of the study. In particular it shows the period when
repeated until a constant mass is recorded. This is the dry growth is most rapid and this corresponds to the steepest
mass. In all cases it is more accurate to obtain the dry mass part of the absolute growth curve. The peak of the absolute
of as large a number of specimens as is practicable and from growth curve marks the point of inflexion after which the
this calculate the mean dry mass. This value will be more rate of growth decreases as the adult size is attained.
representative than that obtained from asingle specimen. Dividing each of the above values for the absolute rate of
growth by the amount of growth at the beginning of each
21.1.2 Types of growth curve : : dm . : panier
time period fe —) and plotting these values against time
Plotting data obtained from any one of the produces a relative rate of growth curve (or specific rate of
physical parameters described above, such as dry mass growth curve) (fig 21.5). This is a measure of the efficiency
(m), against time (t) produces a growth curve which is of growth.
known as the actual or absolute growth curve (fig 21.3). The A comparison of relative rate of growth curves for
usefulness of this curve is that it shows the overall growth organisms grown or reared under different conditions
pattern and the extent of growth. Data from this graph shows clearly the most favourable conditions for rapid
enable the growth curves in figs 21.3—-5 to be constructed. growth and for growth over an extended period. In the case
Plotting the change in parameter against time produces of mammals the absolute growth curve is sigmoid but the
an absolute growth rate curve (fig 21.4). (The same curve exact shape of the curve appears to be related to the time
taken to reach sexual maturity. In the rat the curve is steep
and truly sigmoid since sexual maturity is reached quickly
(within 12 months), whereas in Man the absolute growth
curve shows four distinct phases of increased growth (fig
DARO Ys
Live
mass
Fig 21.3 Actual or absolute growth curve obtained by

plotting live mass against age for sheep. (Data from L. R.
Wallace (1948) J. Agric. Sci., 38, 93 and H. Palsson & J. B.
embryo infancy /childhood adolescence adulthood
Vergés (1952) J. Agric. Sci., 42, 93)
400 = _@e-—----—-—
an 30 foetalgrowth
26 300 = 40
s, 200 Z 30 Juvenile
growth
S 20
ay
= 100 10
A 0
Ort 207 30 40 50 60 ibirth Age/years (1)

i h Age/weeks conception
6 |
= \
Fig 21.4 Absolute growth rate curve plotted from data Set a
Sra a
shown in fig 21.3 zZ |
= on 3 |
1 200 s¢ !
1 000 ee (Oe 4 6 8 10 12 14 16 18 AX 2)
800 | Age/years (1)
& 1.0
600 = |
a=
400 I
200
oS 0.5 |
sf
Gain
of mass 0
as previous
percentage oo :
| 10 20, 30 40 50 60 >= |
a
bh Age/weeks a re ae ae ee ee CC
os
ms Age/years (1)
Fig 21.5 Relative growth rate curve plotted from data shown
in fig. 21.3 Fig 21.6 Three types of growth curve for Man
771
with time therefore show a marked change in mass: an
21.2 Patterns of growth increase in length of only 10% is accompanied by a 33%
increase in mass.
occur among
Various patterns of growth Allometric (allos, other; metron, measure) growth
organisms. occurs when an organ grows at a different rate from the rest
of the body. This produces a change in size of the organism
21.2.1 Isometric and allometric growth which is accompanied by a change in shape of the organism.
Isometric (isos, same; metron, measure) This pattern of growth is characteristic of mammals and
growth occurs when an organ grows at the same mean rate illustrates the relationship between growth and develop-
as the rest of the body. In this situation change in size of the ment. Fig 21.8 shows how the relative proportions of
organism is not accompanied by a change in shape or various structures in Man change as a result of simul-
external form of the organism. The proportions of the two taneous changes in patterns of growth and development. In
structures remain the same. This type of growth pattern is almost all animals the last organs to develop and
seen in fish and exopterygote insects, such as locusts differentiate are the reproductive organs. These show
(except for wings and genitalia) (fig 21.7). In such cases as allometric growth and can be observed only in those
these there is a simple relationship between linear organisms with external genital organs, hence they are not
dimension, area, volume and mass. The area increases as seen in many species of fish where growth appears to be
the square of linear dimension (A ~ I?) whereas volume purely isometric. Fig 21.9 shows the degree of variation in
and mass increase as the cube of linear dimension (V « F patterns of growth of different organs of Man. Again it can
and M « /°), Animals showing little change in overall shape be seen that the last organs to develop are the reproductive
organs.
The shapes of absolute growth curves for whole
organisms as represented by length or mass show remar-
kable similarities and generally conform to the sigmoid
shape described in section 21.1. However, several groups
of organisms show variations on the general pattern which
reflect adaptations to particular modes of life and environ-
ments as described in sections 21.2.2—21.2.3.
200 (b)
Fig. 21.7 (above) Development in fish — an example of S=

isometric growth. The external structures retain their shape
and spatial relationships as a result of a proportional growth
rate. (After Batt (1980) Influences on animal growth and growth/%
Relative
development, Studies in Biology, No. 116, Arnold.)
10 20
Fig 21.8 (below) Development in Man — an example of Age/years
allometric growth. To show the relative rates of growth from
the age of two months to 25 years each stage has been given Fig 21.9 (above) Relative growth rates of (a) brain, (b)
a constant height. (After Stratz, cited in J. Hammond (ed.) thymus and (c) reproductive organs of Man. Each curve is
(1955) Progress in the physiology of farm animals, 2, 437, drawn relative to the absolute growth curve of the whole body
Butterworths.) (d). (After Scammon.)
772
Vi
2 mth
(foetal)
u newborn 2yr 6 yr 12 yr
21.2.2 Limited and unlimited growth
Studies of the duration of growth in plants and
animals show that there are two basic patterns, called E
limited (definite or determinate) growth and unlimited =
Ss
“80
(indefinite or indeterminate) growth.
Growth in annual plants is limited and after a period of
maximum growth, during which the plant matures and
reproduces, there is a period of negative growth or
senescence before the death of the plant. If the dry mass of
the annual plant is plotted against time then an interesting
variation on the sigmoid curve of fig 21.2 is seen, as shown
Time/years
in fig 21.10.
Fig 21.11 Growth curve of a birch tree — a woody perennial
1 | Bile SnSl | | aaa jee |

6 8 IO 1 Te iG ie 20) ae oA
onset of germination Time/weeks 21.2.3 Growth in arthropods
Fig 21.10 Growth curve of a broad bean (Vicia faba) A striking and characteristic growth pattern is
planted in March — an annual plant associated with crustacea and other arthropods, such as
hemimetabolous insects and the larvae of holometabolous
insects. Due to the inelastic nature of their exoskeletons
they appear to grow only in spurts interrupted by a series of
moults (discontinuous growth). A typical hemimetabolous
growth pattern is shown in fig 21.12. It is growth curves
such as these, based on length, which do not give a true
reflection of growth. If a growth curve is plotted for the
same insect, using dry mass as the growth parameter, a
normal sigmoid curve is produced demonstrating that true
growth, as represented by increase in protoplasm, is
continuous.
adult
4.5-
Several plant organs show limited growth but do not a S T
undergo a period of negative growth, for example fruits, Sth

ao Nn
T
instar
organs of vegetative propagation, dicotyledonous leaves 4th
and stem internodes. Animals showing limited growth ee S T instar
3rd
include insects, birds and mammals. Is) n T instar
Woody perennial plants on the other hand show Length/cm 2nd
unlimited growth and have a characteristic growth curve
2.0} instar
Ist
which is a cumulative series of sigmoid curves (fig 21.11), F instar
each of which represents one year’s growth. With unlimited 1 | | | | | | ! fi | | | ! | | | 1 | | J
growth, some slight net growth continues until death. 2 4 6 8 1012 14 16 18 20 22:24 26 28 30 32 34 36 38 40 4244
Other examples of unlimited growth are found among Time/days
fungi, lower plants, particularly algae, and many animals, Fig 21.12 Growth curve showing increase in length of the
particularly invertebrates, fishes and reptiles. Monocotyle- short-horned grasshopper. (After R. Soper & T. Smith (1979)
donous leaves show unlimited growth. Modern Biology, Macmillan.)
ie
21.3 Control of growth and development Recent experiments involving transplantation of tissues
from species with different patterns of growth have
The process of growth and development in an revealed the occurrence of a form of control of growth
organism is controlled ultimately by the information resulting from tissue interaction. If each tissue were to
contained in its DNA. Growth, however, is the result of the grow at its normal rate discrepancies would occur and the
interaction between the DNA and the internal and external organ would show malformations. Instead the growth rates
environments of the organism. Important external in- of both tissues complement each other and growth of the
fluences include availability of food, light, heat and water. organ is normal. The conclusion drawn from these studies
The internal environment includes chemicals such as is that one of the tissues determines the growth rate of the
hormones and recently discovered cytoplasmic proteins other.
(‘transcription factors’) that influence gene expression Some of the factors affecting growth and development
either directly or indirectly. External factors can influence are summarised in table 21.1 (see also fig 21.13).
internal factors, for example lack of iodine in the diet of
Man leads to an inability to make the hormone thyroxine 21.3 From the data shown in fig 21.13
with a consequent reduction in growth rate. explain fully why body mass is at its lowest level at
of factors inflencing growth and times of maximum food intake. |
Table 21.1. Summary
development
30) :
External factors Process affected
Mass/kg a nh
Light (intensity, Photosynthesis — energy source (ch. 9)
quality and Photomorphogenesis (ch. 15)
Food intake/kJ
duration) Phototropism (ch. 15) kg"! day“'
Greening (ch. 15)
Photoperiodism (chs 15 and 16)
Synthesis of vitamin D in Man Mean monthly
temperature/°C
Short wavelength Mutagenesis (ch. 23)
electromagnetic
Nov Jan Mar May Jul Sep Nov
radiation
Month
(X-rays, y-rays etc.)
Nutrients Autotrophic nutrition — carbon Fig 21.13 Graphs showing the effects of food intake and
dioxide, water, inorganic salts temperature on body mass of a husky dog. (Data from J. L.
required — deficiency diseases in Durrer & J. P. Hannon (1962) Am. J. Physiol., 202, 375.)
absence (ch. 9)
Heterotrophic nutrition — organic
compounds, water and inorganic salts
required (ch. 10); deficiency 21.4 Development
diseases in absence (chs 9 and 10)
Temperature Affects growth via effect on enzymes In its broadest sense, growth includes not only
(ch. 6 and fig 21.13): thermo- an irreversible increase in dry mass as a result of cell
periodism (ch. 15) division and increase in cell size, but also the subsequent
Oxygen Needed by aerobic organisms for process of development. During development cells become
respiration: particularly important specialised for particular functions within the organism, a
for active uptake of ions by plant roots
and foetal development (ch. 12)
process known as differentiation. In other words, division
of labour occurs among the cells. Morphogenesis also takes
Water Essential for many processes (ch. 5)
place, as mentioned earlier. The extent of cell differentia-
Seasonal influences Dormancy of buds, seeds and
perennating organs (ch. 15) tion is linked with the level of phylogenetic organisation as
Leaf fall (ch. 15) shown in table 21.2.
Reproduction (chs 15, 16 and 20) One of the intriguing problems of differentiation is the
Photoperiodism and mechanism by which it occurs. Cell division is a major
thermoperiodism (chs 15 and 16) contributory factor in growth, and in all multicellular
Metabolic waste Not normally a problem, but may be organisms this occurs by mitosis. The implication of this is
products inhibitory, e.g. in bacterial colonies
that every cell derived from the original zygote or spore has
Internal factors Process affected an identical genetic composition (genotype) and therefore
ought to have identical structure and function. That the
Genes Protein synthesis (ch. 23), hence latter is not the case in mature organisms is self-evident.
enzyme synthesis and control of cell
chemistry What then are the mechanisms bringing about differences
between cells, tissues, organs and organ systems? Cell
Hormones and Many processes (chs 15 and 16)
growth substances structure and function are determined by the expressed
activity of genes, and if cells differ in their structure and
774
Table 21.2. The approximate numbers of cells and cell types course of development. By transplanting tissue from a
reflecting the degree of differentiation seen at different levels donor embryo into a host embryo they were able to
of phylogenetic organisation demonstrate that the donor tissue was able to change the
TT,
course of development of the host. This process is known as
Organism Approximate number Approximate number induction and the areas of the donor which bring about
of cells of cell types induction are known as organisers. A fuller account of the
Pion 107 107 process, and of Spemann and Mangold’s experiments, is
Porifera +10° $10 given in section 22.8.2. The exact way in which the
Hydra 10° 10-20 influence is exerted is not entirely clear but it is thought to
Annelids 10° 10° involve the differential repression and activation of genes
cue 10" 10° in different cells. This is discussed in detail in section
a
22.8.3.
ooo
Another factor affecting differentiation is hormones.
function this would appear to be due to a differential Specific examples of growth hormones in plants and a range
expression of genes. There are two possible reasons for of animals are described later in this chapter and in
this. Either cells specialise by losing certain genes, chapters 15 and 16. Whilst the action of growth hormones
retaining only those needed for the specialised function, or in plants may be variable, there is evidence that certain
specialisation involves ‘switching’ on and off different aspects of growth hormone activity are common to all
genes in different cells. It became obvious in the early 1960s organisms. Hormones may act directly or indirectly on the
that the latter was the case, at least in plants. Professor genes and exert their influence by ‘switching’ genes ‘on’ or
Steward of Cornell University showed that when differenti- ‘off’ in a sequential manner which determines the pattern
ated cells of a carrot plant, such as phloem cells, were of development. Studies carried out on the giant chromo-
placed in a suitable culture medium they were capable of somes from the salivary glands of dipteran fly larvae show
growing into new carrot plants and therefore still possessed ‘puffing’ in various regions of the chomosomes (sections
all the relevant information. In 1967 Gurdon, working at 22.8.5 and 23.5.1). The positions of these puffs correspond
Oxford University, showed the same to be true of animals to certain stages in the development of the fly. These
when he transplanted nuclei from the intestinal epithelium regions are genes and are the sites of active messenger
of the African clawed toad, Xenopus, into eggs of the same RNA synthesis which results in the synthesis of substances
species whose own nuclei had been destroyed by ultra- vital to certain stages in the life cycle of the fly. Injections of
violet radiation. He found that normal development insect growth hormones can induce ‘puffing’ in larvae
through the tadpole to the adult stage occurred in a small which have had their growth-hormone-producing glands
number of these transplants, suggesting that reactivation of removed. Alternatively the injection of supra-optimal
genes is possible even in differentiated cells, provided the levels of growth hormone can alter the normal sequence of
genes are placed in a suitable environment. Less differenti- ‘puffing’ and hasten the development of the adult fly form.
ated cells from an early embryonic stage of development, These studies suggest that hormones sometimes act by
such as the blastula, gave a higher success rate, and any ‘switching on’ genes associated with development.
number of genetically identical frogs can be produced using
this technique. Identical offspring from a single parent are
referred to as clones, and the technique described above is 21.5 Morphogenesis
called cloning (section 20.1.1).
It is difficult to grow isolated mammalian cells in culture, During the course of development differenti-
and even more difficult to induce differentiation. This ated cells come to occupy specific regions within the
reflects the highly specialised nature of mammalian cells. organism where they undergo further growth and cell
Differentiation may reach such a level of complexity that division. This process is part of morphogenesis because it
even growth is prevented. This situation is found in results in cells, tissues and organs, which give form and
neurones. On the other hand, liver cells remain relatively structure to the mature organism, coming to occupy their
unspecialised and are able to perform a great many characteristic positions. The exact nature of the mor-
functions (section 18.5). An adult mammalian liver, which phogenic movements is species-specific, but some charac-
has had two-thirds of its mass removed, will regenerate in teristic examples of morphogenesis in plants and animals
three weeks to its original size and shape. This evidence not are described in the next two sections. The major
only demonstrates the extent of regenerative growth, but morphogenic events in plants occur during the develop-
also indicates that the growth of organs and tissues is to a ment of the stem, root, leaves and flower. In animals there
predetermined size and shape. are many spectacular examples of morphogenesis, but
If nuclei lose none of their genetic potential as a result of those described here are limited to showing the striking
differentiation that would suggest that the cytoplasm has a changes in form associated with metamorphosis and the
role in regulating differentiation. Spemann and Mangold increasing tissue complexity which accompanies vertebrate
investigated the influence of cytoplasmic factors on the development.
HIS
21.6 Growth and development in the 21.5 The light which passes through -
flowering plant leaves is enriched in green and far-red light relative —
to the light which strikes the leaf surface. -
21.6.1 Seed dormancy (a) Why is this?
Certain environmental conditions, namely (b) What ecological significance might this have in
availability of water, optimum temperature and oxygen, relation to seeds like lettuce, where germination
must be present before the embryo of a seed will grow. is a phytochrome-controlled response? (Read
However, in the presence of these factors, some apparently section 15.4.2 if necessary. }
mature seeds will not germinate and must undergo certain
internal changes which can generally be described as 21.6.2 Germination
after-ripening. These changes ensure that premature
Germination is the onset of growth of the
germination does not occur. For example, seedlings
embryo, usually after a period of dormancy. The structure
produced immediately from seeds shed in summer or
of the seed at germination has been described in section
autumn would probably not survive winter. In other words,
mechanisms exist which ensure that germination is syn-
ee:
chronised with the onset of a season favourable for growth. Environmental conditions needed for
These mechanisms often involve the outer layers of germination
seeds, which may contain growth inhibitors, being imper-
vious to water or the passage of oxygen, or being physically Water. The initial uptake of water by a seed
strong enough to prevent growth of the embryo as in many is by a process called imbibition. It takes place through the
legumes. Sometimes physical damage (scarification) to the micropyle and testa and is purely a physical process caused
seed coat can remove this restriction, a process which can by the adsorption of water by colloidal substances within
be induced artificially by removing the testa or simply the seed. These include proteins, starch and cell wall
pricking it with a pin. Under natural circumstances, materials such as hemicelluloses and pectic substances. The
bacteria may have the same effect. More usually, however, swelling of these substances can lead to strong imbibitional
the restriction is removed by some physiological change, forces sufficient to rupture the testa or pericarp surround-
involving the following factors. ing the seed. Water subsequently moves from cell to cell by
osmosis. It is required to activate the biochemical reactions
Growth inhibitors. Many fruits or seeds con- associated with germination, because these take place in
tain chemical growth inhibitors which prevent germina- aqueous solution. Water is also an important reagent at this
tion. Abscisic acid often has this role, for instance in ash stage in the hydrolysis (digestion) of food stores.
seeds. Thorough soaking of the seeds might remove the
inhibitor or its effect may be overridden by an increase in a Minimum or optimum temperature. There is
growth promoter such as gibberellin. usually a characteristic temperature range outside which a
given type of seed will not germinate. This will be related to
Light. The dormancy of some seeds is broken the normal environment of the plant concerned and will be
by light after water uptake, a phytochrome-controlled within the range 5—40 °C. Temperature influences the rate
response. This is associated with a rise in gibberellin levels of enzyme-controlled reactions as described in section6.4.3.
within the seed. Less commonly, germination is inhibited
by light, such as in Phacelia and Nigella. (For a relevant Oxygen. This is required for aerobic respira-
experiment, refer to the foot of Table 15.5, Section tion, although such respiration can be supplemented with
15.4.2.) anaerobic respiration if necessary.
Temperature. As noted in section 15.5.1, Physiology of germination

seeds commonly require a cold period, or stratification, A typical seed stores carbohydrates, lipids and proteins,
before germination will occur. This is common among either in its endosperm or in the cotyledons of the embryo.
members of the rose family (Rosaceae) and cereals. It is Usually lipids in the form of oils form the major food
associated with a rise in gibberellin activity and sometimes reserves of the seed, though notable exceptions are the
a reduction in growth inhibitors. Leguminosae (legumes) and Gramineae (grasses, includ-
Exactly how light and cold treatments affect seeds is not ing cereals) where starch is the major food reserve. These
clear, but increased permeability of the seed coat, as well as two groups form the principal crops of Man and thus he
changes in levels of growth substances, may be involved. gets the bulk of his carbohydrates from them. Legumes are
also especially rich in proteins, particularly soya bean;
21.4 Seeds which require a stimulus of hence the use of the latter as a source of protein in new
light or germination are usually relatively er foods. In addition, seeds contain high levels of minerals,
What could be re significance of this? notably phosphorus, as well as normal cytoplasmic consti-
tuents such as nucleic acids and vitamins.
776
As a result of imbibition and osmosis the embryo using samples of barley grains at different times from
becomes hydrated, and this activates enzymes such as the germination, increase in amylase activity per grain over a
enzymes of respiration. Other enzymes have to be period of a week can be determined.
synthesised, possibly using amino acids provided by the
digestion of stored proteins. 21.6 Explain the results shown in fig
Broadly speaking, there are two centres of activity in the 2114. ....
germinating seed, the storage centre (food reserve) and the
growth centre (embryo). The main events in the storage
centre, with the exception of enzyme synthesis, are total mass
catabolic, that is concerned with breakdown.
Digestion of the food reserves proceeds mainly by
hydrolysis as below:
Dry
/g
mass
proteases
proteins ———~ amino acids
carbohydrases
polysaccharides ——————> sugars 0 2 4 6 8 10
Time from sowing/days
amy lase maltase
for example starch maltose ———> glucose Fig 21.14 Relative changes in dry mass of endosperm and
embryo during germination of barley
lipids “PSS fatty acids + glycerol
Those seeds which store lipids convert them to fatty acids
and glycerol. Each molecule of lipid yields three molecules
The soluble products of digestion are then translocated of fatty acid and one of glycerol (section 5.3.1). Fatty acids
to the growth regions of the embryo. The sugars, fatty acids are either oxidised directly in respiration or converted to
and glycerol may be used to provide substrates for sucrose, which is then translocated to the embryo. The
respiration in both the storage and the growth centres. latter involves theee cycle (section 11.5.5).
They may also be used for anabolic reactions in the growth
centre, that is, reactions concerned with synthesis. Of - (This question tests” some basic
particular importance in these reactions are glucose and s aeee. che nistry and of the chemistry oO
amino acids. A major use of glucose is for the synthesis of lipids. The latteris covered insection 5.3.)
cellulose and other cell wall materials. Amino acids are _ Suppose 51.29 dry mass of seeds ‘containing
used mainly for protein synthesis, proteins being important 50% fatty acid by mass, converteddallthefeya
acid_ ;
as enzymes and structural components of protoplasm. In _ sugar in the following reaction: .
addition, mineral salts are required for the many reasons _ CigH.O, + 110;——> C,,H,,0,, + 4004.+ Ho . energy
given in table 9.10. fatty acid | ‘sugar _ _
Both storage and growth centres obtain the energy for ff) Assuming that no other ee bectied whch. -
their activities from respiration. This involves oxidation of might affect dry mass, Ce thegene lossin
a substrate, usually sugar, to carbon dioxide and water. A _ dry mass ofthe seeds. -
net loss in dry mass of the seed therefore occurs, since
carbon dioxide is lost as a gas, and water does not eee atomic masses: oS H ah:
| 0-16) —
contribute to dry mass. This loss will continue until the
seedling produces green leaves and starts to make its own (b)a other importantchangemightaffectdy.
food (section 21.2.2). macs?
A well-studied example of germination of a polysacchar-
i“
c) Calculate the volume of carbon diowde evolved :
ide-rich seed is the barley grain, where it has been shown
from the seeds at STP cme fompa latte /
and Pressure). — |
that the synthesis of a-amylase and other enzymes takes
place in the outer layers of the endosperm in response to a moleof gas att STP occupies 22.4am?) _
gibberellin secreted by the embryo. These outer layers . (@)How can fatty acid be obtained from a lipid,and :
contain stored protein which is the source of amino acids what would be the other component¢of the lipid?
for protein synthesis. The process is described and (e)How many carbon atoms would one molecule of ©
experimentally investigated in section 15.2.6. Fig 15.21 _ the parent lipid have contained if CrcHasOe was
shows an example of the role of hormones in early _ the only,fatty acid produced?
germination. The appearance of amylase in germinating What is the identity of the sugar formed inthe
barley grains can also be investigated by grinding them in reaction shown? —
water, filtering and centrifuging to obtain a clear extract (9)t
How does the oxygen reach thestorage tissue? :
and testing the activity of the extract on starch solution. By
ad
Respiration in germinating seeds 21.9 The RQ of peasis normally between_
Respiratory rates in both storage tissues and embryo are 2.8 and 4 during the first seven days of germination, —
high owing to the intense metabolic activity in both regions. but is 1.5-2.4 ifthe testas are removed. Inbothcases _
Substrates for respiration may differ in each region and ethanol accumulates in the seeds, but in much
may also change during germination. This is revealed by smaller amounts when the testas are removed.
changes in the respiratory quotient (section 11.7.7). Account for these observations.
21.8 When castor oil seeds were ana-

lysed for lipid and sugar content during germination
_ in darkness, the results shown in fig 21.15 were Growth of the embryo
obtained. . Within the embryo growth occurs by cell division,
The RQ of the seedlings was measured at day 5 enlargement and differentiation. Amounts of proteins,
and the embryo was found to have an RQ of about cellulose, nucleic acids and so on increase steadily in the
1.0, while the remaining cotyledons had an RQ of growing regions while dry mass of the food store decreases.
about 0.4—0.5. The first visible sign of growth is the emergence of the
(a) Suggest as full an explanation of these results as embryonic root, the radicle. This is positively geotropic
you can (refer to section 21.6.2 for relevant and will grow down and anchor the seed. Subsequently, the
information). embryonic shoot, the plumule, emerges and being nega-
(b) What would you expect the RQ of the whole tively geotropic (and positively phototropic if above
seedling to be on day 11? Explain very briefly. ground) will grow upwards.
There are two types of germination according to whether
or not the cotyledons grow above ground or remain below
50 total dry mass
it. In dicotyledons, if that part of the shoot axis, or
internode, just below the cotyledons (the hypocotyl)
elongates, then the cotyledons are carried above ground.
This is epigeal germination. If the internode just above the
cotyledons (the epicotyl) elongates, then the cotyledons
remain below ground. This is hypogeal germination.
In epigeal germination, the hypocotyl remains hooked as
seedlings/g
100
Mass
per
0 | alt 1 1 L j it grows through the soil, as shown in fig 21.16 (b), thus
2-4 6 One Oe:
Time from sowing/days
meeting the resistance of the soil rather than the delicate
plumule tip, which is further protected by being enclosed
Fig 21.15 Changes in lipid and sugar content of castor oil by the cotyledons. In hypogeal germination of dicotyledons
seeds during germination in the dark. (Based on data from the epicotyl is hooked, again protecting the plumule tip, as
R. Desveaux & M. Kogane-Charles (1952) Annis. Inst. natn.
shown in fig 21.16c. In both cases the hooked structure
Rech. Agron., Paris, 3, 385-416; cited by H. S. Street &
H. Opik (1976) The physiology of flowering plants 2nd ed., immediately straightens on exposure to light, a phyto-
Arnold.) chrome-controlled response.
endosperm
epicotyl___ (@)
plumule
cotyledon___ eee y
radicle hypocotyl
epigeal germination hypogeal germination

(hypocotyl elongates) (epicotyl elongates)
(b) plumule (¢)

two cotyledons hooked \ hooked first green leaves
hypocotyl plumule phytochrome-controlled
photosynthetic a
unhooking,
‘testa phytochrome-controlled
SS
\\ cotyledons
L NNO Aye eae
Bey Ys sae 4e:
ee
e.g. broad bean
lateral roots e.g. sunflower (non-endospermous)
(non-endospermous),
castor oil (endospermous) lateral roots
Fig 21.16 (a) Structure of a seed, (b) Epigeal germination, (c) Hypogeal germination
778
Table 21.3. Types of meristem and their functions
a
ee
Type of meristem Location Role Effect

Apical Root and shoot apex Responsible for primary growth, giving rise to Increase in length
primary plant body
Lateral Laterally situated in older parts of the Responsible for secondary growth. Vascular Increase in girth
(cambium) plant parallel with the long axis of cambium gives rise to secondary vascular tissue;
organs, e.g. cork cambium (phellogen), phellogen gives rise to the periderm, which
vascular cambium replaces the epidermis and includes cork
Intercalary Between regions of permanent tissue, Allows growth in length to occur in regions other Increase in length
e.g. at nodes of many monocotyledons, than tips. This is useful if the tips are susceptible
such as bases of grass leaves to damage or destruction, e.g. eating by herbivores
(grasses), wave action (kelps). Branching from the
main axis is not then necessary
Se
ORe—eeleoqeueesS$$99m9amw—e—weaeqws
=saoa=$eoqo
In the grasses, which are monocotyledons, the plumule is namely herbaceous plants or herbs. A few herbaceous
protected by a sheath called the coleoptile, which is plants show restricted amounts of secondary thickening, as
positively phototropic and negatively geotropic as de- in the development of additional vascular bundles in
scribed in sectionn 15.1.1. The first leaf grows out through Helianthus (sunflower).
the coleoptile and unrolls in response to light. On emerging
Apical meristems and primary growth
into light a number of phytochrome-controlled responses
rapidly occur, collectively known as photomorphogenesis. A typical apical meristem cell is relatively small, cuboid,
The overall effect is a change from etiolation (section with a thin cellulose cell wall and dense cytoplasmic
15.4.1) to normal growth. The major changes involved are contents. It has a few small vacuoles rather than the large
summarised in table 15.5 and include expansion of the vacuoles characteristic of parenchyma cells, and the
cotyledons or first true foliage leaves, as well as formation cytoplasm contains small, undifferentiated plastids called
of chlorophyll (‘greening’). At this point photosynthesis proplastids. Meristematic cells are packed tightly together
begins and net dry mass of the seedling starts to increase as with no obvious air spaces between the cells.
it finally becomes independent of its food reserves and The cells are called initials. When they divide by mitosis
assumes an autotrophic existence. Once exposed to light, one daughter cell remains in the meristem while the other
the shoot also shows phototropic responses although these increases in size and differentiates to become part of the
are not phytochrome controlled. permanent plant body.
21.6.4 Primary growth of the shoot

21.6.3 Growth of the primary plant body
The structure of a typical apical shoot meri-
Meristems stem is illustrated in figs 21.17 and 21.18. Fig 21.18 shows
In contrast to animals, growth in multicellular plants, with the approximate division of the shoot apex into regions of
the exception of the young embryo, is confined to certain cell division, cell expansion and cell differentiation. There
regions known as meristems. A meristem is a group of cells is more overlap in these zones than in the root. Passing
which retain the ability to divide by mitosis, producing back from the dome-shaped apical meristem, the cells get
daughter cells which grow and form the rest of the plant progressively older, so that different stages of growth can
body. The daughter cells form the permanent tissue, that be observed simultaneously in the same apex. Thus it is
is, cells which have lost the ability to divide. There are relatively easy to study developmental sequences of plant
three types of meristem, described in table 21.3. Two types tissue.
of growth are mentioned in table 21.3, namely primary and Three basic types of meristematic tissue are recognised,
secondary growth. namely the protoderm, which gives rise to the epidermis,
Primary growth is the first form of growth to occur. A the procambium, giving rise to the vascular tissues, in-
whole plant can be built up by primary growth, and in most cluding pericycle, phloem, vascular cambium and xylem;
monocotyledonous plants and herbaceous dicotyledons it and the ground meristem, producing the parenchyma
is the only type of growth. It is a result of the activity of the ground tissues, which in the dicotyledons are the cortex and
apical, and sometimes intercalary, meristems. The ana- pith. These meristematic types are laid down by division of
tomy of mature primary roots and stems is dealt with in the initials in the apex. In the zone of expansion, the
section 14.3.9. daughter cells produced by the initials increase in size,
Some plants continue with secondary growth from lateral mainly by osmotic uptake of water into the cytoplasm and
meristems. This is most notable in shrubs and trees, which then into the vacuoles. Increase in the length of stems and
are sometimes described as arborescent to distinguish them roots is mainly brought about by elongation of cells during
from plants which lack extensive secondary growth, this stage. The process is illustrated in fig 21.19.
I79
cytoplasm nucleus
cell wall
epidermal hair
leaf
a. :
ae apical meristem
* zone of cell division
large
leaf primordium central
vacuole
protoderm developing vacuoles
procambial strand
< — elongated cells
Fig 21.19 Expansion phase of growth of a meristematic cell
that develop into
xylem and phloem
still growing. In contrast, developing sclerenchyma cells

ek Ze
ae a deposit thick layers of lignin on their walls and soon die.
SAN rae Thus their differentiation does not start until expansion is
Ni
we
oe ey
7
D
yA.
] ss virtually completed.
Mes FY, The procambium forms aseries of longitudinally running
AS
rea * oO
2 ~ oe 7 oxke iP strands whose cells are narrower and longer than those of
the ground meristem. The first cells to differentiate in the
CAS
ay Pe baat ry
Perkin
+
ge s
AE
( k
procambium are those of the protoxylem to the inside, and

Pan eA OT Uf protophloem to the outside. These are the parts of the
Fig 21.17 (above) The apical meristem primary xylem and phloem respectively which form before
elongation is complete. The protoxylem typically has only
annular or spiral thickenings of lignin on tracheids (section
Fig 21.18 (below) LS shoot tip of a dicotyledon showing
8.2.1) which, being discontinuous, allow extension and
apical meristem and regions of primary growth. For simplicity,
vascular tissue to leaves and buds has been omitted stretching of the cellulose between the thickenings as the
surrounding tissue elongates. Both protoxylem and proto-
leaf primordium forming at node
phloem elements soon die and generally get crushed and
apical meristem
leaf protoderm—» epidermis stretched to the point of collapse as growth continues
ZONE OF
ground meristem —pground tissues around them. Their function is taken over by later-
CELL DIVISION procambium—+» primary
vascular developing xylem and phloem in the zone of differentia-
eee ee eS a
Voip tissues tion.
ZONE OF protophloem) beginnings of
CELL DIVISION protoxylem Jf vascular tissue In the zone of differentiation each cell becomes fully
AND CELL
EXPANSION
internode — region between specialised for its own particular function, according to its
two nodes
lateral or axillary bud position in the organ with respect to other cells. The
node
ZONE OF greatest changes occur in the procambial strands, which
=)
CELL DIFFER- epidermis
ENTIATION pericycle — sclerenchyma differentiate into vascular bundles. This involves lignifica-
cambiuium phloem tion of the walls of sclerenchyma fibres and xylem
PERMANENT
TISSUES elements, as well as development of the tubes characteristic
of xylem vessels and phloem sieve tubes. The final forms of
these tissues are described in section 8.2. Sclerenchyma
The small vacuoles increase in size, eventually fusing to and xylem now supplement the support previously given by
form a single large vacuole. The turgor pressure developed collenchyma and turgid parenchyma. Between the xylem
inside the cells stretches their thin walls and the orientation and phloem there are cells which retain the ability to
of cellulose microfibrils in the walls helps to determine divide. They form the vascular cambium, whose activities
the final shape assumed by the cells. The final volume of are described later, with secondary thickening.
cytoplasm may not be significantly greater than in the
original meristematic cell, but is now confined to the cell Leaf primordia and lateral buds
periphery by the vacuole. As expansion nears completion, Development of the shoot also includes growth of leaves
many cells develop additional thickening of the cell walls, and lateral buds. Leaves arise as small swellings or ridges
either of cellulose or lignin, depending on the type of cell called leaf primordia, shown particularly clearly in fig
being formed. This may restrict further expansion, but 21.17. The swellings contain groups of meristematic cells
does not necessarily prevent it. Collenchyma cells in the and appear at regular intervals, their sites of origin being
cortex, for example, can continue elongating while extra called nodes and the regions between internodes. The
cellulose is laid down in columns on the inside of the pattern of leaf arrangement on the stem varies and is called
original walls. Thus they can give support to the plant while phyllotaxis. Leaves may arise in whorls with two or more
780
leaves at each node, or singly, either in two opposite ranks grains which act as statoliths, sedimenting to the bottoms of
or in a spiral pattern. Generally, however, they are cells in response to gravity. Their role is described in more
arranged to minimise overlapping, and hence shading, detail in section 15.2.2.
when fully grown so that they form a mosaic. Behind the quiescent centre, orderly rows of cells can be
The primordia elongate rapidly, so they soon enclose seen and the meristematic regions already described in the
and protect the apical meristem, both physically and by the shoot, namely protoderm, ground meristem and procam-
heat they generate in respiration. Later they grow and bium, can be distinguished (fig 21.20). In the root the term
increase in area to form the leaf blades. Cell division procambium is used to describe the whole central cylinder
gradually ceases but may continue until they are about half of the root, even though at maturity this contains the
their mature size. non-vascular tissues of the pericycle and the pith, if
Soon after the leaves start to grow, buds develop in the present.
axils between them and the stem. These are small groups of The zone of cell division typically extends 1-2 mm back
meristematic cells which normally remain dormant, but from the root tip, and overlaps slightly with the zone of cell
retain the capacity to divide and grow at a later stage. They elongation. Root tips are convenient material for observa-
form branches or specialised structures such as flowers and tion of mitosis and a procedure for this is described in
underground structures such as rhizomes and tubers. They section 22.2. Behind this zone, growth is mainly by cell
are thought to be under the control of the apical meristem elongation, cells increasing in size in the manner described
(see apical dominance, section 15.3.3). for the shoot and shown in fig 21.19. The zone of elongating
cells extends to a point about 10 mm behind the root tip
21.6.5 Primary growth of the root and their increase in length forces the root tip down
through the soil.
The structure of the typical apical root system
is illustrated in fig 21.20. Some cell differentiation begins in the zone of cell
division, with the development of the first phloem sieve
At the very tip of the apical meristem is a quiescent
tube elements (fig 21.20). In longitudinal sections, neat
centre, a group of initials (meristematic cells) from which
files of developing sieve tube elements can be seen, getting
all other cells in the root can be traced, but whose rate of
progressively more mature further back from the root tip,
cell division is much slower than their daughter cells in the
until they become mature sieve tubes. Development of
apical meristem around them. To the outside, the cells of
phloem is from the outside inwards.
the root cap are formed. These become large parenchyma
Further back in the zone of elongation, the xylem vessels
cells which protect the apical meristem as the root grows
start to differentiate, also from the outside inwards (exarch
through the soil. They are constantly being worn away and
xylem) in contrast to the stem (endarch xylem). The
replaced. They also have the important additional function
first-formed vessels are protoxylem vessels, as in the stem,
of acting as gravity sensors, since they contain large starch
and they show the same pattern of lignification and ability
lateral root primordium to stretch as cells around them grow. Their role is taken
beacaproovemersing
root eee 4 Q
arising from pericycle over by metaxylem, which develops later and matures in
PERMANENT TISSUES ; F
eyes sis eb RS 7 epidermis the zone of differentiation after elongation has ceased. The
mature root hair cortex xylem often spreads to the centre of the root, in which case
GUL
ZONE OF CELL
DIFFERENTIATION pericycle ; no pith develops.
young root hair Development is easier to examine in roots than in shoots.
I aa ees tie: protoxylem
metaxylem In the latter procambial strands to the leaves complicate
e xylem vessel elements begin to the distribution of developing tissues. Development of
Py differentiate, protoxylem to the xylem, in particular, is easily seen by squashing apical
ZONE OF CELL pe outside
ION ‘
pith — may be absent
portions of fine roots such as those of cress seedlings and
oe
first mature phloem sieve tubes staining appropriately.
phloem sieve elements begin to After all cells have stopped elongating, further dif-
differentiate
procambiumrpericycle, primary ferentiation is completed. This includes the development
phloem, vascular cambium, of root hairs from the epidermis.
primary xylem, pith (if present):
: ‘| forms a solid central cylinder
Lateral roots
ZONE OF CELL DIVISION oy ground meristem cortex,
Ee including endodermis Branching from the main root may occur but not by means
protodermp epidermis of buds, in contrast to the shoot. Instead, a small group of
root cap quiescent zone — apical initials pericycle cells in the zone of differentiation resumes
meristematic activity and forms a new root apical meri-
Fig 21.20 LS apical meristem of a typical root. Xylem _ stem. It then grows, forcing its way out through the
differentiation is shown to the right and phloem differentiation endodermis, cortex and epidermis as shown in fig 21.20.
to the left. In reality xylem and phloem alternate round the Such development is termed endarch, compared with
root and would be on different radii. (See fig 14.16). Also, in exarch development of lateral buds.
reality, the zone of elongation would be longer
781
plane of cell companion cell sieve plate of sieve
Adventitious growth division is tube
Adventitious structures are those growing in uncharac- tangential secondary phloem
teristic positions. Adventitious roots and buds may arise in ( : | eS > fusiform initial
a variety of situations as a result of certain cells resuming
meristematic activity. Adventitious roots develop indepen- fusiform initial ea) secondary xylem
dently of the original primary root and form the main (a) secondary
rooting system of monocotyledons, arising from nodes on xylem vessel
the stem. Rhizomes and runners are stem structures from (b) cell
which adventitious roots arise directly. Adventitious roots > ———_
are also important in propagation of plants by stem |
cuttings. Ivy clings by adventitious roots. one fusiform plane of cell two fusiform initials
division is radial
Adventitious buds may develop on roots, stems or leaves. initial
For example African violets can be propagated from leaf Fig 21.22 (a) Two successive divisions of a fusiform initial to
cuttings, which develop adventitious roots and buds. Trees form xylem and phloem, seen in TS. In reality, differentiation
may develop new branches adventitiously from buds that of xylem and phloem to the stages shown would take some
time, during which more cells would be produced. (b) Division
arise in the trunk.
of a fusiform initial to form a new fusiform initial, seen in TS
21.6.6 Lateral meristems and secondary
growth
Secondary growth is that growth which occurs
after primary growth as a result of the activity of lateral
meristems. It results in an increase in girth. It is usually
associated with deposition of large amounts of secondary Secondary growth in woody dicotyledon
xylem, called wood, which completely modifies the primary stems
structure and is a characteristic feature of trees and shrubs. The vascular cambium is originally located between the
There are two types of lateral meristem, the vascular primary xylem and primary phloem of the vascular
cambium which gives rise to new vascular tissue, and the bundles, its derivation from the apical meristem being
cork cambium or phellogen, which arises later to replace shown in fig 21.18. It becomes active very soon after
the ruptured epidermis of the expanding plant body. primary cell differentiation is complete. Fig 21.23 summa-
rises the early stages in secondary thickening of a typical
Vascular cambium
woody dicotyledon stem.
There are two types of cell in the vascular cambium, the Fig 21.23a shows the original primary stem structure,
fusiform initials and the ray initials, illustrated in fig 21.21. omitting the pericycle for simplicity. Fig 21.235 shows the
Fusiform initials are narrow, elongated cells which divide development of a complete cylinder of cambium. Fig
by mitosis to form secondary phloem to the outside or 21.23c shows a complete ring of secondary thickening.
secondary xylem to the inside. The amount of xylem Here, fusiform initials have produced large quantities of
produced normally exceeds the amount of phloem. secondary xylem, and lesser quantities of secondary
Successive divisions are shown in fig 21.22. Secondary phloem, while the ray initials have produced rays of
phloem contains sieve tubes, companion cells, scleren- parenchyma. As the stem increases in thickness, so the
chyma fibres and sclereids, and parenchyma. circumference of the cambium layers must increase. To
Ray initials are almost spherical and divide by mitosis to achieve this, radial divisions of the cambial cells occur, as
form parenchyma cells which accumulate to form rays shown in fig 21.22. The original ray initials produce primary
between the neighbouring xylem and phloem. medullary rays which run all the way from pith to cortex,
unlike the secondary medullary rays produced by later ray
initials. The rays maintain a living link between the pith and
LS two fusiform initials
cortex. They help to transmit water and mineral salts from
TS or LS ray initial, the xylem, and food substances from the phloem, radially
cell is isodiametric
(almost spherical) across the stem. Also, gaseous exchange can occur by
TS two fusiform initials diffusion through intercellular spaces. The rays may also be
used for food storage, an important function during periods
of dormancy, as in winter. In three dimensions they appear
thin cellulose as radially—longitudinally running sheets because the ray
cell walls
initials occur in stacks one above the other, as shown in fig
21.24. Fig 21.24 illustrates the appearance of wood
(secondary xylem) and the rays it contains in the three
Fig 21.21 Fusiform and ray initials planes TS, TLS and RLS.
782
(a)
epidermis Fig 21.25 shows part of the stem of a woody dicotyledon
in its third year of growth, revealing the large amounts of
D secondary xylem produced. Fig. 21.26 shows photographs
primary phloem
of a three-year-old and a five-year-old stem of Tilia, the
s intrafascicular cambium
=
lime tree.
xylem
Annual rings
Each year in temperate climates, growth resumes in the
primary medullary ray linking
cortex with pith spring. The first vessels formed are wide and thin-walled,
being suitable for the conduction of large quantities of
water. Water is required to initiate growth, particularly the
(b) expansion of new cells, as in developing leaves. Later in the
year, fewer vessels are produced and they are narrower
intrafascicular cambium (in vascular
bundle)
with thicker walls. During winter the cambium remains
aN interfascicular cambium (between dormant. The autumn wood produced at the end of one
M vascular bundles) year, as growth ceases, will therefore be immediately next
X = Jz
secondary xylem
phloem
to the spring wood of the following year and will differ
markedly in appearance. This contrast is seen as the annual
ring and is clearly visible in fig 21.26. Where vessels are
concentrated in the early wood it is said to be ring porous,
as opposed to diffuse porous wood, where they are evenly
primary phloem distributed, and where it is more difficult to see annual
secondary phloem rings. In tropical climates, seasonal droughts may induce
vascular cambium similar fluctuations in cambial activity.
secondary xylem The width of an annual ring will vary partly according to
primary xylem climate, a favourable climate resulting in production of
pith more wood and hence a greater distance between rings.
This has been used in two areas of science, namely
hy y primary medullary ray runs from dendroclimatology and dendrochronology. Dendroclima-
cortex to pith
See 4 secondary medullary ray
tology is the study of climate using tree ring data.
Applications vary from correlation of recent climatic
Fig 21.23 Early stages in secondary thickening of a typical
records with tree growth, of possible interest in a specific
woody dicotyledon stem. (a) Primary structure of stem. (b) locality, to investigations of more distant climatic events
Cambium forms a complete cylinder as parenchyma cells in several hundreds or even thousands of years in the past.
the medullary rays become meristematic, spreading outwards The oldest-known living trees, the bristlecone pines, are
from the vascular bundles. Meanwhile secondary xylem and about 5 000 years old, and fossil wood of even greater age
phloem are already being formed by the existing cambium. (c)
can be found.
A complete ring of secondary thickening has developed.
Thickening is most advanced at the sites of the original Dendrochronology is the dating of wood by recognition
vascular bundles where cambial activity first started of the pattern of annual rings. This pattern can act as a
primary medullary ray

TS Medullary ray appears as a radial row
secondary medullary ray
of ray parenchyma cells (extending
from pith to cortex if primary). Some
secondary phloem rays are more than one cell wide in TS.
vascular cambium [ ic
secondary xylem fe cortex
v=
aes
XZSBX
TLS Medullary ray appears edge on as a
column of ray parenchyma cells.
Pa Some rays are more than one cell
cortex fant wide in TLS.
Aaa
eeeTy
yy
VT
TELE
LL
ASE
SSE
YAAURRAARAUABA?”.
{
COLL
RLS Medullary ray appears as a sheet of
fusiform initials cells (extending from pith to cortex if
vascular cambium stack of ray primary).
ray initials initials (shaded)
woody dicotyledonous stem. A

Fig 21.24 Diagrammatic representation of primary and secondary medullary rays in a typical longitudinal
longitudinal section; RLS, radial
primary ray is shown to the right and a secondary ray to the left. (TLS, transverse
section.)
783
\
position of primary phloem (crushed beyond recognition) remains of epidermis

pericycle (Sclerenchym
= a) cork (phellem)
band of sieve tubes and companion cells cork cambium (phellogen) periderm
band of phloem fibres secondary cortex (phelloderm)
small autumn vessels (late wood) primary cortex (parenchyma)
large spring vessels (early wood) secondary phloem
secondary medullary rays
vascular cambium
annual ring
primary medullary rays
secondary xylem (wood) of second year’s growth

first year’s
metaxylem ,
growth primary xylem
protoxylem
pith (parenchyma) \) =
Fig 21.25 (above) TS of a typical woody dicotyledonous stem in the third year of growth (age two years), such as Tilia.
Details of secondary phloem, secondary xylem and secondary medullary rays are shown only in the left-hand sector
(a)
@© annual rings marking end of first year’s growth remains of epidermis
and second year’s growth respectively
medullary ray {ar 7 Bo}
secondary xylem (wood) of second
years growth secondary phloem ne oo CO” | LTS re 17 5 3i % primary medullary
»_
large spring vessels of band of fibres she
small
lenticel
af 4
primary
"Pty
y
xylem
a
w
’ 5 Ky
g@ :
small autumn vessels Age seconda
of second year’s growt ry
medullary rays
A ae b
vascular cambium primary medullary ray if3 : i he smallil vessels ads
ps
band of sieve tubes Ee of spring wood » %
and companion cells 1
Fig 21.26 (a) TS of a two-year-old (third year) twig of Tilia SA Vien Fons‘
joe ae
vulgaris (x 2.2). (b) Part of a TS of a five-year-old (sixth year) iA |
twig of Tilia vulgaris (x 11.5) “are
‘fingerprint’, pinpointing the time during which the wood _——whereas the outer, wetter conducting wood is called
was growing. Dating of timbers at archaeological sites, in | sapwood.
old buildings and ships and so on thus becomes feasible, ;
provided enough data are available. ae roteSe eres
s the secondary xylem grows outwards, so the tissues
Heartwood and sapwood outside it become teu compressed, as well as being
As a tree ages, the wood at the centre may cease toservea _ stretched sideways by the increasing circumference. This
conducting function and become blocked with darkly affects the epidermis, cortex, primary phloem and all but
staining deposits such as tannins. It is called heartwood, the most recent secondary phloem. The epidermis even-
784
tually ruptures and is replaced by cork as the result of the Secondary growth in roots of dicotyledons
activity of a second lateral meristem, the cork cambium or
Most dicotyledons that show secondary growth of the stem
phellogen. It generally arises immediately below the
also show secondary growth of the roots. In the case of
epidermis. Cork (or phellem) is produced to the outside of
plants with storage roots, such as carrots and turnip, it may
the cork cambium, while to the inside one or two layers of
be more conspicuous than in the stem, although paren-
parenchyma are produced. These are indistinguishable
chyma predominates in the xylem and phloem of the
from the primary cortex and form the phelloderm or secon-
examples given. The process of secondary growth is
dary cortex. The phellogen, cork and phelloderm together
summarised in fig 21.28 and is similar in principle to
comprise the periderm (fig 21.25).
secondary growth in stems.
As the cork cells mature, their walls become impreg-
nated with a fatty substance called suberin which is epidermis
impermeable to water and gases. The cells gradually die
PG, eee 5
cortex
and lose their living contents, becoming filled either with endodermis
air or with resin or tannins. The older, dead cork cells fit pericycle
together around the stem, preventing desiccation, infec- primary xylem
tion and mechanical injury. They become compressed as
the stem increases in girth and may eventually be lost and vascular cambium
replaced by younger cells from beneath. If the cork layer
primary phloem
were complete, the respiratory gases oxygen and carbon
dioxide could not be exchanged between the living cells of
the stem and the environment, and the cells would die. At () endodermis
random intervals, however, slit-like openings, or lenticels,
primary phloem and part of
develop in the cork containing a mass of loosely packed,
|
pericycle
thin-walled dead cells, lacking suberin. They are produced secondary phloem
by the cork cambium and have large intercellular air spaces wide primary medullary ray
allowing gaseous exchange. often formed opposite arms of
xylem
Fig 21.27 shows a diagram of cork and lenticels.
vascular cambium
air spaces allow lenticel remains of

gaseous exchange epidermis oy : primary xylem
radial row of loosely packed secondary medullary ray
compressed parenchyma
cork cells
() primary medullary ray
remains of primary phloem
cork (phellem)
secondary phloem
secondary medullary ray
[_~\ cork cambium
t- J (phellogen) secondary xylem
___ | phelloderm primary xylem
(secondary
cortex) vascular cambium
parenchyma of primary medullary ray
primary cortex
cork
Fig 21.27 VS lenticel (cell contents omitted) periderm
cork cambium
Bark Fig 21.28 Stages in the secondary growth of a

Eventually a woody stem becomes covered with a layer dicotyledonous root. (a) The vascular cambium develops from
commonly known as bark. The term bark is an imprecise cells of the procambium (fig 21.23) which have remained
meristematic. The cambium continues to grow to form a
one which is used to refer either to all the tissues outside the complete ring (see arrows) as cells of the pericycle outside
vascular system, or more strictly to those tissues outside the the xylem arms become meristematic. (b) The cambium
cork cambium. Peeling bark from a tree generally strips produces rows of secondary xylem vessels internally and
tissues down to the vascular cambium, athin layer of cells secondary phloem externally. Growth of secondary xylem
starts earlier and is more rapid between the arms of the
which is easily ruptured. xylem, until the cambium becomes circular, rather than wavy,
It is usual for the cork cambium to be renewed each year in outline. (c) As growth continues, the increasing girth
as the girth of the stem increases. Often a cork cambium stretches and tears the endodermis, cortex and epidermis,
arises in the secondary phloem, in which case the bark will, and these are sloughed off. A cork cambium develops from
over a number of years, build up a layered appearance due the pericycle. ((c) is drawn to a smaller scale than (a) and
to alternating layers of secondary phloem and bark. (b).)
785
position of vascular
e - cambium
cork cambium oN . secondary
periderm ee ee ee ee
x phloem
». secondary xylem
* (large vessels
and fibres)
primary 3~ .
xylem \ -
(4 arms present) ie
‘primary
medullary ray
Fig 21.29 Secondarily thickened root of Vicia faba (x 120)
Development of secondary xylem, secondary phloem Table 21.4. Some larval stages found in the animal kingdom
and medullary rays is basically similar to that in stems, and
annual rings are usually formed as in stems. A cork Animal group Larval stage
cambium develops from the pericycle and serves the same
function as in the stem. A phelloderm may be produced, coelenterates planula
trematodes miracidium/redia/cercaria
but is indistinguishable from the pericycle. As with the
cestodes hexacanth/cysticercus
stem, the original cork cambium may be replaced at polychaetes trochophore
intervals by cork cambium arising further inside the root. crustacea nauplius/cypris/zoaea
Fig 21.29 is a photograph of a transverse section through a insects caterpillar (maggot, grub) or nymph
root with secondary thickening. molluscs trochophore/Vveliger
echinoderms dipleurula/pluteus
urochordates ascidian tadpole
lamprey ammocoete
21.7 Metamorphosis amphibia tadpole
The term metamorphosis (meta, between;

morphe, form) applies to those rapid changes which occur 21.7.1 Adaptive significance of larval stages
during the transition from larval to adult form. This is the Larvae generally act as a dispersal phase for
process of postembryonic maturation and is found in the distribution of species. This is very important to sessile
many invertebrate groups, especially amongst insects, organisms as it provides a means of preventing overcrowd-
echinoderms, hemichordates, urochordates, cephalochor- ing of the parental organisms by their offspring. Over-
dates and some fish and amphibian species. Table 21.4 crowding would lead to increased competition for food and
lists various animal groups and names of their larval other resources and this could be harmful to the survival of
stages. the species. For example, many sessile marine organisms
786
such as barnacles and mussels produce
vast numbers of 21.7.3 Hormonal control of ecdysis and
larvae which are planktonic and are disp
ersed by ocean moulting in insects
currents. At a certain stage they settle on
solid objects, a
phenomenon known as ‘spat-fall’, and The involvement of hormones in ecdysis and
continue their
development. moulting was first described by Wigglesworth in Cam-
Larvae usually occupy a different habitat from bridge. In a series of experiments he was able to show that
the adult
forms and have different feeding habits, forms of moulting is controlled by a moulting hormone (MH) which
locomo-
tion and behavioural patterns which enable the is released in response to a specific stimulus. The result of
species to
exploit the potential of two ecological niches duri the moulting, in terms of stage in the life cycle, may be
ng the life
cycle. This increases the chances of survival of the modified by another hormone, juvenile hormone (JH),
species
between the egg and adult Stages. Many species, for which prevents the appearance of the adult form.
ex-
ample the dragonfly, only feed and grow during their larva Wigglesworth chose to investigate insect growth hor-
l
stages which form the longest period of their life. mone in Rhodnius prolixus, a blood-sucking hemipteran
Another feature of larvae is their ability to act as from South America. Rhodnius is an exopterygote insect
a
transition stage during which the species has time to adapt with five nymphal stages and no pupal stage. During each
to the new conditions that will be encountered in the adult nymphal stage only one large blood meal is required and
habitat. Physiological hardiness is another attribute which this acts as the stimulus for moulting. Wigglesworth was
allows larvae to act as a dormant Stage during adverse able to demonstrate that stretch receptors in the abdominal
conditions. For example, some insects overwinter in the wall, activated by distension, set up nerve impulses which
soil as larvae but most exist as another metamorphic stage, are transmitted to the brain. These impulses initiate a series
the pupa, and the physiological changes which accompany of coordinated endocrine secretions which result in
these stages are described in section 21.7.3. moulting. In the case of the moult from 4th to Sth instar this
A final advantage of larval stages is that they may occurs within 14 days of feeding at 28°C.
provide an opportunity for the numbers of larvae to be If the insect is decapitated within four days of a blood
increased. This is common in the lifecycles of certain meal and the wound plugged with wax it will live for 18
platyhelminths (section 4.5.3). months without showing any further growth or develop-
However, larvae should not be thought of as incom- ment. If decapitation occurs after this four-day period it
pletely developed forms. In many cases they are highly will continue to grow but no moulting will occur. These
developed, as shown by insect larval stages, where the only observations suggest that it takes four days for impulses
undeveloped structures are the reproductive organs. An from the gut to be passed to the brain and hormones to be
interesting example of a fully developed larval stage which released which affect the growth and differentiation of the
has attained sexual maturity, the axolotl, is described in target tissues, including the epidermal cells.
In his experiments Wigglesworth transplanted the brain
section 21.7.4.
from an insect which was just about to moult, into the
21.7.2 Characteristics of metamorphosis abdomen of a starved decapitated insect which then
proceeded to moult. Transplanting brain tissue from a
The structural and functional changes which moulting insect into an insect without head or thorax did
occur during metamorphosis prepare the organism for not result in moulting. Clearly then, a substance from the
adult life in a new habitat or environment. In some cases brain was influencing a region in the thorax which produced
these new adaptations may appear whilst the organism still the moulting response. Previous studies on silkworm
retains the larval form and occupies the larval habitat. In caterpillars had revealed the presence of a gland in the first
other words metamorphosis is not entirely a response to a thoracic segment, the prothoracic gland, which could
change in environment but a preparation for a change. For induce moulting if implanted into the posterior region of a
example, tadpoles, living in water where desiccation is not silkworm which was separated from the thorax by a
a problem, lose approximately 80% of their nitrogenous ligature. This suggested that moulting in Rhodnius is also
waste as ammonia. In the later stages of metamorphosis, controlled by a hormone from the prothoracic gland.
whilst still living in water, tadpoles begin to excrete urea. Moulting hormone was isolated in 1953 and is a steroid
By the time the young frogs are ready to leave the water with a similar structure to cholesterol (section 5.3.6). The
they are excreting 80% of their nitrogenous waste as urea. hormone was named ecdysone, and because of the presence
Whilst growth and differentiation may be influenced by of more hydroxyl groups than cholesterol it is water-
hormones, metamorphosis is entirely controlled and reg- soluble. Since then it has been synthesised and produces all
ulated by endocrine secretions. In some cases metamor- the responses normally associated with moulting when
phosis is a response to increasing hormone levels and in injected into insects.
other cases it is the result of different responses by different Ecdysone is thought to activate genes controlling the
target organs to the same level of hormone. These production of enzymes involved in growth. This response is
responses may involve either cell death and reduction in most marked in the epidermal cells, which undergo a series
size of an organ (involution) or cell division, differentiation of changes associated with ecdysis and which are described
and the growth of an organ. in fig 21.30.
787
(a) Sepicuticle
=) ee The corpus allatum produces a hormone called juvenile
in
hormone or neotonin. Growth and metamorphosis
ee
exocuticle
n
arthropods is now seen to be controlled by the interactio
a.
ane eae
is belie ved to
.____endocuticle of ecdysone and juvenile hormone. The latter
Ss 2
cause the retention of larval or nymphal characteristics by
LG
activating genes responsible for these characters and
thickening of suppressing genes responsible for producing adult cuticle
epidermal cells and structure.
The release of ecdysone and juvenile hormone is
(b) controlled by yet another hormone, which is produced by
i orgegen Tee Oa ..__epicuticle
the brain. The mechanism of release may be described with
exocuticle reference to the locust (fig 21.31).
endocuticle neurosecretory cells of the pars

intercerebralis
Z ae
<h sem new
“YS? epicuticle
nervi corpora cardiaca

(c) (d)
mid-dorsal line
epicuticle corpus cardiacum
exocuticle
a one
Se
corpus allatum
endocuticle
é ‘
a
SSeS
new epicuticle oesophagus
new exocuticle
E
SEEEE epidermis
, eee aS prothoracic gland
Fig 21.30 (a) TS Rhodnius cuticle. Without blood meal, Fig 21.31 Moulting glands of the locust
epidermis thin and flattened. Following blood meal the
epidermis starts to thicken out and separate from the
endocuticle above. (b) New epicuticle then forms over the Neurosecretory cell bodies situated in the pars inter-
epidermis. Chitinase and proteinase enzymes are secreted by cerebralis of the brain secrete a hormone, prothoracico-
epidermis and break down the old cuticle. New cuticle is trophic hormone (PTTH), which passes down the axons of
shielded from enzymes by the formation of a new epicuticle.
the neurosecretory cells through the nervi corpora car-
Old digested endocuticle is absorbed by the epidermis and
incorporated into new endocuticle. (c) Rhodnius takes in air, diaca possibly by axoplasmic streaming (section 16.6.2).
expands itself and breaks the old cuticle and moults. The new The hormone is stored in a pair of glands, the corpora
cuticle is soft and hardens by tanning. (d) Moulting occurs by cardiaca, which are situated immediately behind the brain.
splitting of the old cuticle along the mid-dorsal line which is The stimulus to moult, whatever it may be, sets up nerve
specially thinned
impulses in the brain which pass down the nervi corpora
cardiaca and cause the release of prothoracicotrophic
The addition of ecdysone to insect larvae with ‘giant hormone into the blood (fig 21.32). This passes to the
chromosomes’ produces ‘puffing’ in specific regions of the prothoracic gland and ecdysone is released which stimu-
chromosomes. These are sites of messenger RNA synthesis lates, in turn, the epidermal cells to produce a new cuticle.
and this evidence suggests that ecdysone works directly at A similar pattern of events is involved in the control of
the level of genetic transcription. ecdysis and moulting in Rhodnius but the corpora cardiaca
In a further series of experiments Wigglesworth obtained and corpora allata are not bilateral structures, having fused
data which suggested that the head region of Rhodnius together to form single structures situated in the mid-line of
larvae produces another hormone which promotes the the body.
retention of nymphal characteristics, thus preventing In endopterygote insects the formation of the pupal stage
development into the adult form. Investigations revealed occurs in the presence of a very low level of juvenile
that a region just behind the brain, the corpus allatum, hormone whereas the absence of juvenile hormone results
produced this response. Implantation of the corpus allatum in a direct metamorphosis into the adult form. If the corpus
from a third- or fourth-stage nymph into a fifth-stage allatum is removed from a second, third or fourth nymphal
produces a sixth-stage giant nymph and not an adult. stage of an exopterygote it will moult into a miniature
Implantation of the corpus allatum ofa fifth-stage nymph adult. Therefore the increase in size and the attainment of
does not have this effect. sexual maturity in insects may possibly result from falling
788
food light temperature
21.7.4 Metamorphosis in amphibia
Metamorphosis in amphibia involves the tran-

BRAIN sition of an aquatic larva into a semi-terrestrial adult. The
neurosecretory
mechanisms of metamorphosis are controlled by endocrine
secretions in an analogous way to those described in
insects. Gudernatsch, a German biologist, discovered in
1911 that tadpoles feeding on extracts of mammalian
thyroid glands would rapidly change into miniature frogs.
prothoracicotrophic hormone
This precocious metamorphosis, under the influence of
thyroid extract, implicated the hormone thyroxine as the
factor responsible for the many anatomical, physiological,
corpus cardiacum biochemical and behavioural changes in the life cycle of the
prothoracicotrophic hormone frog. Further investigations involved the removal of the
thyroid gland in some animals and the pituitary gland in
others. In both cases metamorphosis was prevented but
was restored by feeding with thyroxine.
The initial stimulus for metamorphosis is thought to be a
change in the environmental conditions such as increase in
juvenile hormone ecdysone
temperature or daylength. These external stimuli are
_transmitted to the brain and activate the hypothalamus
which secretes thyrotrophin releasing hormone (TRH).
This enters the blood vessels of the hypothalamico-
larval characteristics adult characteristics hypophyseal portal system (section 16.6.2) and passes to
Fig 21.32 The hormonal control of ecdysis and moulting in the anterior pituitary gland. Here thyroid stimulating
insects hormone or thyrotrophin (TSH) is released which enters
the general circulation and stimulates the thyroid gland to
levels of juvenile hormone. The control of growth and release thyroxine (tetraiodothyronine) and _triiodothyr-
metamorphosis in the life cycles of specific endopterygote onine. These hormones act directly on all metamorphos-
and exopterygote insects is summarised in fig 21.33. ing tissue and produce varying responses such as the
Once the adult form has been attained in pterygote atrophy of the larval tail and gills, accelerated growth of the
insects the prothoracic glands atrophy and disappear, limbs and tongue and differentiation of the skin to form
gland cells and chromatophores (pigment-containing
preventing any further ecdyses or moults. This does not
occur in apterygote insects. In all adult insects the corpus cells). The direct or local effects of thyroxine on meta-
allatum becomes active again and juvenile hormone is morphosis at the tissue and biochemical levels were demon-
strated by applying thyroxine ina fatty base to the left side
secreted. In females it is necessary for the deposition of
of the tail and the left eye of several tadpoles. Pure fatty
yolk in the egg and in males it is required for the normal
base was added to the right side to act as a control. After a
functioning of the accessory glands which produce the
few days, regression of the tails on the left side and a
spermatophore, the capsule used to tiansfer sperm to the
change from the larval visual pigment porphyropsin to the
female.
adult visual pigment rhodopsin had occurred in the left
eyes. No change had occurred in the control tissues.
Pieris brassica (cabbage white butterfly)- an endopterygote
The early stages of metamorphosis are controlled by
MH + JH -MH +JH MH positive feedback, that is to say increasing amounts of
thyroxine released by the growing thyroid gland stimulate
egg > | Wy >5 —Ppupa—> imago the further production of thyroxine. As the level of
(oh ath am eee pr eat Sie)
larval stages circulating thyroxine increases, different target organs
respond by showing cell death, cell division or cell
Rhodnius prolixus - an exopterygote
differentiation, as described in the examples quoted above.
MH+JH MH+JH MH+ JH + JH
MH MH
Once the adult form has been attained thyroxine secretion
egg
we 4
yiige et
e 5 & imago
is controlled by negative feedback (section 16.6.5). The
sequence of external morphological changes shown by the
nymphal stages tadpole during its metamorphosis is summarised in table
21,5:
Fig 21.33 The influence of growth hormones on The Mexican axolotl, Amblystoma mexicanum (fig
endopterygote and exopterygote insect life cycles (MH
21.34), does not normally undergo metamorphosis due to a
moulting hormone, JH juvenile hormone. The height of the
letters JH indicates relative concentration.) failure of the pituitary gland to produce thyroid stimulating
789
Fig 21.34 The axolotl to have played a significant role in the physiological
development of many animal groups, for example, the
Table 21.5. Changes in external features in frog during earliest vertebrates, which are thought to have descended
metamorphosis. Times and sizes can only be approximate as from neotenous larval forms of sea-squirts.
the rate of metamorphosis depends upon food supply, The axolotl can be induced to metamorphose by the
temperature and internal factors addition of the correct concentration of thyroxine to the
water in which it is living. The visible effects of this are loss
Time/ Size/ Stages in development of gills and most of the tail fin and the assumption of the
weeks mm shape and features of a close relative, the land salamander.
0 — Fertilisation
1 7 Hatches from jelly. External gills, tail, mouth
with horny jaws. Mucus glands beneath 21.8 Development in vertebrates
mouth
yy 9 External gills begin to atrophy, operculum The preceding section on metamorphosis in
grows over gills. Eyes prominent amphibia described the morphological changes which
4 1; External gills and mucus gland lost. occur during the development of the larval form into the
Spiracle develops. Tail widens to aid adult form. Whilst these changes are spectacular and have
swimming tremendous adaptive significance in preparing the organ-
a 28 Hindlimbs appear as buds ism to meet the demands of the adult habitat, they appear
35 Hindlimbs fully formed but not used for trivial when compared to the events following fertilisation
swimming. Head begins to broaden of the ovum. For example, of the 47 generations of cell
11-12 35 Left limb emerges through spiracle. Right division estimated to occur during the development of
limb enclosed by operculum. Hindlimbs used Man, 42 occur during the period of development which
for swimming
precedes birth. The study of embryonic development is
13 25 Eyes become prominent and mouth widens.
known as embryology and the events which occur during
14 20 Tail begins to be reabsorbed this time are fundamentally the same in all animal species.
16 dD All external larval features have disappeared. They increase our understanding of the origins and
Frog emerges on to land
functional relationships between tissues, organs and organ
systems. This development is conveniently divided into
hormone (TSH). Instead, the axolotl attains sexual three main phases, but it should be appreciated that the
maturity whilst retaining larval characteristics such as overall process of embryonic development is continuous
external gills, tail fins and a light coloration. This is an and one stage passes imperceptibly into the next. These
example of neoteny, the attainment of sexual maturity stages and their main features and implications are as
whilst retaining larval characteristics. Neoteny is believed follows.
790
Cleavage (or segmentation) This is the differentiate, and by a series of morphogenic movements
series of mitotic divisions of the zygote nucleus and
buckle inwards (invaginate) through the blastopore and
cytoplasm that follows fertilisation. In the frog, the point of
obliterate the blastocoel forming a new cavity, the
sperm entry into the egg prior to fertilisation imposes archenteron. The cells become rearranged as three distinct
symmetry on the future embryo and marks the position of
layers, called primary germ layers, within a double-layered
the future dorsal lip of the blastopore. Fertilisation structure called the gastrula. The germ layers, the
stimulates cleavage which produces daughter cells or ectoderm, mesoderm and endoderm, are now spatially
blastomeres forming a hollow ball of cells, the blastula,
organised so as to undergo organogeny.
enclosing a cavity called the blastocoel.
By injecting cells with different coloured dyes (a Organogeny (organogenesis). Further cell
technique known as intra vitam staining) the fate of the division, growth and differentiation occurs involving many
cells of the blastula can be followed throughout the next complex morphogenic activities including the formation of
two processes of gastrulation and organogeny. Vogt has the notochord, the formation of the central nervous system
used this technique and produced fate maps which show the and the development of the mesoderm. In vertebrates
prospective or presumptive fate of cells of different regions, these give rise to the tissues, organs and organ systems of
as shown in fig 21.35. the embryo as shown in figs 21.36 and 21.37. Organogeny is
terminated by the hatching or birth of the embryo.
ectoderm neural plate
neural crest
archenteron
neural tube
blastocoele e notochord dermatome
myotome
<« dorsal lip of
notochord
blastopore
sclerotome
nephrotome
yolk plug
ectoderm
mesoderm somatic mesoderm
Fig 21.35 VS of developing gastrula showing presumptive splanchnic mesoderm

areas. Later growth and cell migration of the ectoderm and intestine
mesoderm is indicated by arrows. Mesoderm spreads coelom
inwards and occupies a position beneath the ectoderm yolk cells of endoderm
Fig 21.36 Diagram of TS of newly hatched frog tadpole

showing derivatives of the germ layers shown in fig 21.37.
Gastrulation. The onset of gastrulation is
The tadpole is still dependent upon food provided by yolk
marked by the appearance of a circular opening, the since the ectoderm has not differentiated sufficiently to form
blastopore. In the frog, cells of the blastula divide rapidly, the complete alimentary canal
| epidermis, epidermal glands, hair, feathers, scales, enamel, conjunctiva, lens, ears,
oa epithelia of nose, mouth, anus and vagina, anterior pituitary
/\\
akVS central nervous system, sympathetic and parasympathetic ganglia, somatic motor
ectoderm ———— EE s = She . 5
ae neurones, retina, posterior pituitary
neural crest jaws, dorsal root ganglia, somatic sensory neurones, adrenal medulla
myotome skeletal muscle, diaphragm

sclerotome vertebrae, notochordal sheath
somatic
‘ dermatome dermis, dentine of teeth
body wall muscles
nephrotome kidney tubules, ureters, oviducts, uterus, parts of ovaries and testes, adrenal cortex
blastomeres mesoderm
smooth muscle, heart, blood vessels, blood, lymphatic system, lungs, connective
splanchnic (mesenchyme) tissue (including cartilage, bones, mesenteries), sclerotic, choroid and cornea
notochord
SU most of alimentary canal
A
epithelia ——_—_—_—_—— lining of gut, bladder, lungs, Eustachian tubes
endoderm
glands liver, pancreas, thyroid, parathyroid, thymus
gonads primary germ cells
Fig 21.37 The embryological fate of the three germ layers in vertebrates
791
21.8.1 Mammalian growth hormones and differentiation to restore the size of the organ. This
process is known as compensatory hypertrophy. The cells
There are four glands which secrete hormones of the central nervous system, on the other hand, are
influencing growth and development; they are the thyroid, incapable of regeneration if damaged or lost.
liver, adrenal cortex and gonads. In all cases these glands Regeneration is the replacement, by growth, of parts of
secrete hormones under the control of other hormones an organism lost for one of the reasons stated above. It is
released from the pituitary gland. The pituitary gland in common in plants, particularly in the angiosperms where
turn is under the influence of specific releasing and whole plants can be grown from cuttings which involve no
inhibiting factors produced by the hypothalamus (section more than asingle leaf and its petiole, such as Tradescantia.
16.6.2). Examples of these three levels of hormonal control The regenerative power of plants is used as a means of
are described with reference to the release of the growth vegetative propagation (section 20.1.1).
hormone, somatomedin by the liver. The degree to which animals are able to regenerate new
During the early part of this century numerous experi- structures appears to be related to their structural
ments involving ablation (removal) of the pituitary gland complexity. Sponges, coelenterates and flatworms have
and replacement therapy confirmed the role of the pituitary remarkable powers of regeneration. Sponges and some
in determining the rate and extent of growth in animals. A coelenterates, such as Hydra, are capable of being
pituitary extract was isolated which increased body mass macerated and regenerating complete organisms. Planaria
when injected into animals. This substance was called can be cut in half longitudinally or transversely and then
‘srowth hormone’. Subsequent research has revealed that regenerate new halves. Another flatworm, Dugesia can
this growth hormone is a trophic hormone, somatotrophin, completely regenerate from a small piece cut from an adult.
which is released from the pituitary gland under the Most species of annelids are able to regenerate missing
influence of two other hormones produced by the hypotha- regions of the body, for example the earthworm can
lamus. These are somatostatin, a peptide made up of 40 regenerate the first five segments if they are lost and a
amino acids and growth hormone-releasing factor (GH- substantial portion of the posterior end. Crustacea, insects
RF). These two hormones are thought to have an and echinoderms can regenerate new limbs and arms, just
antagonistic effect, with somatostatin being an inhibitor as amphibia and lizards are able to regenerate limbs and
and GH-RF a stimulator of somatotrophin release. The tails as a result of attack by others of the species or
former is thought to have a dominant influence over the predators. Lizards with their long tails are able to shed
latter. them if attacked. The wriggling movements of the detached
Human somatotrophin is a protein of 190 amino acids. It tail act as a diversion enabling the lizard to escape. This
acts via receptor sites in the hepatocytes of the liver, and ability to voluntarily shed limbs is shared by lobsters and
stimulates synthesis and release of somatomedin into the
crabs and is called autotomy.
hepatic vein. Somatomedin stimulates growth throughout
the body by increasing the synthesis of nucleic acids in
preparation for mitosis and increasing the amino acid
uptake into cartilage and muscle. It also produces a rise in
21.10 Dormancy
the intracellular levels of potassium, phosphorus, calcium
During their life cycle many plants and
and sodium, and changes in blood glucose and lipid levels
animals undergo periods when metabolic activity is
aimed at providing extra metabolites for cell synthesis and
reduced to a minimum and growth ceases. This is
growth.
dormancy. It is a physiologically determined behavioural
response and may involve the adult organism, a develop-
ment stage such as the pupa, or a specialised reproductive
21.9 Repair and regeneration body such as a spore. Dormancy is induced by environmen-
tal changes and has the biological significance of enabling
Cell growth does not cease with the attainment the species to survive periods when the supply of energy is
of adult size. The loss of cells and tissues as a result of inadequate for normal growth and metabolism.
ageing, damage sustained by disease, accident or attack by Most plant species of tropical and temperate zones, and
other organisms can act as a stimulus for cell division and all arctic plant species, exhibit dormancy in response to
differentiation, resulting in the healing, repair and replace- changing light, temperature and moisture conditions. The
ment of damaged or missing tissues. mechanisms of control of dormancy by plant growth
Many tissues in mammals, such as skin, gut epithelial regulator substances are described in section 15.5.
cells and blood cells, are continuously replaced throughout Animals show three forms of dormancy involving
life at a steady rate determined by the normal rate at which physiological adaptations which enable them to tolerate
they are lost. Other tissues, for example the liver, thyroid environmental extremes, and these are considered here.
and pancreas, normally show a very low rate of cell division Some animals though, because of well-developed mechan-
in the adult stage. If any part of these organs is lost in any isms of locomotion, are able to avoid unfavourable en-
way, the remaining tissues will undergo rapid cell division vironmental conditions by moving to more favourable
792
regions, such as by migration. These responses, although p~<q_adult
coordinated by physiological mechanisms, are strictly
high
behavioural and are described in section 16.8.
21.10.1 Diapause
Diapause is a form of arrested development level of pro-
thoracicotrophic
shown by insects, but this term is also used to describe the hormone
period of inactivity shown by protozoan cysts and the
winter eggs of many freshwater aquatic organisms. Insect Temperature/°C
life cycles are adapted to produce offspring at times when
food is plentiful for the larval and nymphal stages.
Diapause is a means of coordinating these two events.
Development can be arrested at any stage in the life cycle, low
I 2 3 4 5 6 i 8
egg, larva, pupa or adult, and for any species the exact Time/weeks
Stage or stages is determined genetically. Some insects
always undergo a period of diapause known as obligatory Fig 21.38 Graph showing the relationship between the
levels of prothoracicotrophic hormone in the brain and blood
diapause, for example the locust Nomadacris, whereas of silkworm at 3 °C and 25 °C
others enter diapause only if conditions become un-
favourable, such as the locust Schistocerca, and this is
termed facultative diapause. é They show a long-day response and only enter diapause if
The main factor inducing diapause in all species is exposed to periods of short daylength. Other species, such
daylength (photoperiod), but food availability, tempera- as the silkworm, Bombyx mori, which has an embryonic
ture and moisture may be influential. In order for diapause diapause (egg stage), grow and develop normally during
to be effective, the factor inducing diapause must act in periods of short daylength and only enter diapause
advance of the adverse conditions. Therefore the mechan-.
following exposure to periods of long daylength. This is a
ism of synchronisation often acts on a stage preceding that short-day response. Diapause eggs of the silkworm begin to
which undergoes diapause. For example, diapause in the grow and develop during the short daylengths of spring,
and the adult emerges during early summer. As develop-
egg stage of the silkworm, Bombyx mori, is the result of the
ment occurs during periods of short daylength the eggs laid
photoperiod experienced by the female parent during
by these adults hatch and develop during the summer
development of the egg. Adult diapause, on the other
without entering diapause. However, the eggs laid by these
hand, is induced by light conditions occurring during the
summer silkworms enter diapause in preparation for
early larval stages.
winter. Since long photoperiods induce diapause in this
Light is thought to have a direct effect upon the brain
type of insect this limits the number of generations per
which, in turn, influences the activity of the neurosecretory
year. In the case of the silkworm there are two, as shown in
system. In aphids, light is thought to act directly on
fig 21.39.
neurosecretory cells. In the case of the giant silkworm,
Hyalophora cecropia, the short photoperiod of autumn
short long short
inactivates neurosecretory cells in the brain which produce
prothoracicotrophic hormone. This prevents the release of photoperiodi, ar
ecdysone from the thoracic glands and, in the absence of
further development, the species overwinters in the pupal
lifecycle egg —»larva—» adult—» egg —» larva—»adult—» diapause egg»
stage. Normal secretion and further development is only stage @ @)
restored after an extended period of exposure to low
temperatures. Paradoxically, it is the adverse conditions season winter spring summer autumn winter
themselves, which diapause avoids, which act as the
Fig 21.39 Diagram showing the relationship between
stimulus to break the period of dormancy (fig 21.38). daylength, diapause and stages in the life cycle of Bombyx
mori. This is an example of a short-day response
--—-24.40 Using the experimental data
in fig 21.38, account for the mechanisms
presented Investigations have revealed the existence of a diapause
_ which terminate diapause. hormone in Bombyx which determines whether or not the
egg will undergo diapause. The long photoperiods of
Most species of insect exposed to long periods of daylight summer stimulate the brain to release a diapause hormone
- undergo continuous growth and development and produce from a pair of neurosecretory cells situated in the
several generations per year, for example the black bean suboesophageal ganglion. This has a direct effect on the
aphid, Aphis fabae. These species do not undergo diapause ovary which releases darkly pigmented eggs which undergo
if artificially maintained in conditions of long daylength. diapause. Silkworms which develop during the short
793
short photoperiod brain long photoperiod may wake up, if the environmental temperature rises, in
order to feed on supplies kept in the burrow or to urinate
and defaecate.
no stimulus stimulus The onset of hibernation is related to decreasing
circumoesophageal
commissure
environmental temperature and photoperiod and the
suboesophageal gland
accumulation of adequate supplies of stored food. There is
.
no diapause hormone KE
4
~ diapause hormone some experimental evidence that suggests a metabolite is
present in the blood which could trigger hibernation, since
non-pigmented egg __¢

C) oO
(oy
@) @——Pigmented egg
blood taken from a hibernating ground squirrel induced
hibernation in a non-hibernating squirrel within 48 h.
Likewise hibernation has been induced in ground squirrels
no diapause diapause at times of the year when they are normally active by
injecting blood from a hibernating squirrel. Some species
Fig 21.40 Diagram showing the effects of photoperiod on
such as the woodchuck, Arctomys, enter hibernation
the production of eggs in Bombyx mori
within a few hours whereas other species allow their core
photoperiods of later winter and spring do not release temperature to fall steadily over a period of several nights
diapause hormone and lay non-pigmented eggs which do before achieving their hibernation temperature.
not undergo diapause (fig 21.40). As the body temperature falls to about 1-2 °C above the
environmental temperature the heart rate may fall from
21.10.2 Aestivation 400 to 8 beats per minute, as in the ground squirrel, and the
basal metabolic rate may fall to less than 2% of the
Aestivation (aestas, summer) is a particular non-hibernating rate. Hibernating animals are unable to
form of dormancy found in certain fish and amphibia in withstand a considerable drop in temperature, but they
response to hot dry periods. In such cases the organisms avoid freezing conditions by hibernating in nests or
bury themselves in mud whilst it is still soft and undergo a burrows where the microclimate is several degrees above
period of metabolic inactivity which enables them to freezing. Curling up the body aids in reducing the body
survive conditions of extreme dehydration. During this surface area and conserving heat, as does burying the nose
time they breathe atmospheric air. Normal metabolic and mouth into the fur. If conditions become severe and
activities are restored only when the river, pool or lake body temperature falls to about 2 °C there is a danger
floods again following the period of drought. Two of the of tissue damage by frost. The animal prevents this either
three genera of lungfish, Lepidosiren in South America and by waking up and becoming active, thereby raising its
Protopterus in Africa, are able to aestivate for at least six temperature, or by increasing its metabolic rate without
months when the rivers they inhabit dry up. waking.
Many hibernating animals store energy as lipid droplets
21.10.3 Hibernation in a special type of adipose tissue known as brown fat.
Hibernation (hiber, winter) is a period of These cells contain many spherical mitochondria, and it is
relatively low metabolic activity associated with periods of the presence of these that gives the cells their characteristic
low temperature. This enables hibernating species of colour. Lipids in these cells are stored throughout the
amphibia, reptiles, birds and mammals to survive at times cytoplasm as droplets and not as a solid. Throughout
when food is in short supply by reducing their energy hibernation stored lipids are used up steadily, but as
demands to a low level. This is especially important in the animals come out of hibernation the metabolic activity of
case of homeotherms, which normally require a high these cells rises sharply. This release of energy enables the
metabolic rate to maintain their core temperature. True animal to shiver vigorously and generate heat, which is
hibernation involves a fall in body temperature to that of dispersed by the blood system as the heart rate increases.
the environment (the ambient temperature) and a reduc- The heart, brain and lungs receive this heat first as a result
tion in heart rate, ventilation rate, metabolic rate, growth of vasoconstriction of blood vessels to other parts of the
and development. In such conditions animals become body. This is initiated by the sympathetic nervous system.
almost poikilothermic. Examples of organisms showing Gradually the whole body warms up and normal metabolic
true hibernation include insectivores such as shrews and and physiological activities are resumed.
hedgehogs: rodents including marmots, ground squirrels, Some small birds and mammals with large surface area to
dormice and hamsters: insectivorous bats and a few birds. volume ratios have extremely high metabolic rates, and in
Most other species which show some form of hibernation order to survive they lower their body temperatures at
are in fact in a state of torpor, sleep or pseudohibernation. night when they are unable to feed. This is known as
These include snakes, lizards, tortoises, salamanders, diurnal hibernation and is seen in humming birds and small
toads, newts and frogs. During this period the organism insectivorous bats.
794
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Chapter Twenty-two
Continuity of life
The development of the cell theory by Schleiden and cell during division since they are responsible for the
Schwann is described at the beginning of chapter 7. transmission of hereditary information from generation to
Rudolph Virchow extended this theory in 1855 by declaring generation and have a regulatory role in cell metabolism
that ‘omnis cellula e cellula’ (‘every cell is from a cell’). (section 23.2). The chromosomes of eukaryotic cells are
Recognition of the continuity of life stimulated further composed of DNA, proteins and small amounts of RNA
workers throughout the later part of the nineteenth century (section 5.6). In non-dividing cells the chromosomes are
to investigate the structure of the cell and the mechanisms extremely long and thin and are dispersed throughout the
involved in cell division. Improved techniques of staining nucleus. Individual chromosomes cannot be seen, but the
and better microscopes with increased resolution revealed chromosomal material may stain with certain basic dyes
the importance of the nuclei, and in particular the (appendix A2.4.2.) and is known as chromatin (coloured
chromosomes within them, as being the structures provid- threads). During cell division the chromosomes shorten
ing continuity between one generation of cells and another. and the stain intensifies so that individual chromosomes
In 1879 Boveri and Flemming described the events can be seen. In their extended form the chromosomes are
occurring within the nucleus leading to the production of involved in controlling the synthesis of all materials in the
two identical cells, and in 1887 Weismann suggested that a cell, but during cell division this function ceases.
specialised form of division occurred in the production of In all forms of cell division the DNA of each chromo-
gametes. These two forms of division are called mitosis and some undergoes replication so that two identical polynuc-
meiosis respectively. The basic processes are almost leotide chains of DNA are produced (section 22.4.2).
identical but the outcome is entirely different. These become ‘surrounded’ by a protein ‘coat’ and at the
Mitosis is the process by which a cell nucleus divides to onset of cell division appear as two identical strands lying
produce two daughter nuclei containing identical sets of side by side. Each strand is called a chromatid and they are
chromosomes to the parent cell. It is usually followed attached to each other by a non-staining region called the
immediately by an equal division of the cytoplasm, the centromere (kinetochore).
re-formation of a cell (plasma) membrane and cell wall
(plants), or cell (plasma) membrane only (animals), and
separation of the two daughter cells. This process is known 22.1 The cell cycle
as cell division. Mitosis with cell division results in an
increase in cell numbers and is the method by which The sequence of events which occurs between
growth, replacement and repair of cells occurs in all higher the formation of a cell and its division into daughter cells is
animals and plants. In unicellular organisms, mitosis called the cell cycle. It has three main stages.
provides the mechanism of asexual reproduction leading to (1) Interphase. This is a period of intense synthesis and
an increase in their numbers. growth. The cell produces many materials required for
Meiosis is the process by which a cell nucleus divides to its own growth and carrying out all the functions
produce four daughter nuclei each containing half the peculiar to it. DNA replication occurs during inter-
number of chromosomes of the original nucleus. An phase.
alternative name for meiosis is reduction division since it (2) Mitosis. This is the process of nuclear division involving
reduces the number of chromosomes in the cell from the the separation of chromatids and their redistribution as
diploid number (2n) to the haploid number (n).. The chromosomes into daughter cells.
significance of the process lies in the fact that it enables the (3) Cell division. This is the process of division of the
chromosome number of a sexually reproducing species to cytoplasm into two daughter cells.
be kept constant from generation to generation. Meiosis The entire cycle is laid out in fig 22.1.
occurs only during gamete formation in animals and during The length of the cycle depends on the type of cell and
spore formation in plants showing alternation of genera- external factors such as temperature, food and oxygen
tions (section 20.2), and produces haploid nuclei that fuse supply. Bacteria may divide every 20 min, epithelial cells
together during fertilisation to restore the diploid number of the intestine wall every 8-10 h, onion root-tip cells may
of chromosomes. take 20 h, whilst many cells of the nervous system never
Chromosomes are the most significant structures in the divide.
796
PROPHASE
Method
METAPHASE (1) Place a pin through a clove of garlic and suspend ina
ANAPHASE test- tube full of water so that the base of the clove is
covered with water. Leave for 3—4 days without any
cell division disturbance as this is likely to inhibit cell division
temporarily.
(2) When several roots have grown 1-2 cm, remove the
clove and cut off the terminal 1 cm of the root.
cellular synthesis
(3) Transfer the roots to a small corked tube containing
acetic alcohol and leave overnight at room tempera-
ture to fix the material.
CELE (4) Remove the root tips with forceps by grasping the cut
DIVISION end of the root, transfer to a Petri dish containing
distilled water and wash for a few minutes to remove
Phase Events within cell the fixative. ;
(5) Transfer the root tips to a test-tube containing molar
Intensive cellular synthesis. Mitochondria, chloroplasts (in hydrochloric acid which is maintained at 60 °C for
plants), ER, lysosomes, Golgi apparatus, vacuoles and
vesicles produced. Nucleolus produces rRNA. mRNA and
3 min (6-10 min for onion, peas and beans). This
G, tRNA and ribosomes synthesised. Cell produces breaks down the middle lamellae holding the cells
structural and functional proteins. Cell metabolic rate high together and hydrolyses the DNA of the chromo-
and controlled by enzymes. Cell growth occurs. Substances
produced to inhibit or stimulate onset of next phase. somes to form deoxyribose aldehydes which will react
with the stain.
DNA replication occurs. Protein molecules called histones
(6) Pour the root tips and acid into a Petri dish. Remove
are synthesised and cover each DNA strand. Each the roots into another Petri dish containing distilled
chromosome has become two chromatids. water and wash to remove the acid. Leave for 5 min.
Intensive cellular synthesis. Mitochondria and chloroplasts
(7) Transfer the roots to a small tube containing Feulgen
divide. Energy stores increase. Centrioles replicate (in cells stain and cork. Leave in a cool dark place (preferably
possessing them) and mitotic spindle begins to form. a refrigerator) for a minimum of 2 h.
(8) Remove a root tip and place in a drop of acetic alcohol
Nuclear division occurs in four phases.
on a clean microscope slide.
Equal distribution of organelles and cytaplasm into each (9) Cut off the terminal 1-2 mm of the root tip and discard
daughter cell. the rest of the root.
(10) Tease out the root tip using a pair of fine needles and
Fig 22.1 The cell cycle cover with a cover-slip. Place the slide on a flat
surface, cover with several sheets of blotting paper
and press down firmly over the cover-slip with the ball
of the thumb. Do not allow the cover-slip to move
Experiment 22.1: To investigate the phases of sideways.
mitosis
(11) Examine the slide under the low and high powers of
Chromosomes can normally be observed only during the microscope and identify cells showing different
nuclear division. The apical meristem of roots (root tips) of phases of mitosis.
garlic (2n=16), onion (2n=16) and broad bean (2n=12) (12) Draw and label nuclei showing the various phases.
provide suitable material for the experiment. The material
is set up so that root development is initiated and the root
tips are removed, fixed, stained and macerated so that the 22.2 Mitosis
chromosomes may be observed under the microscope.
The events occurring within the nucleus
Materials during mitosis are usually observed in cells which have
pins pair of fine needles been fixed and stained (appendix A2.4.2). Whilst this
test-tube containing water several sheets of shows the phases through which chromosomes pass during
scalpel blotting paper cell division, it fails to reveal the continuity of the process.
small corked tube clove of garlic The use of the phase-contrast microscope and time-lapse
forceps distilled water photography has enabled the events of nuclear division to
2 Petri dishes acetic alcohol be seen in the living cell as they happen. By speeding up the
water bath and test-tube molar hydrochloric acid film, mitosis is seen as a continuous process which occurs in
microscope slide Feulgen stain four active stages. The changes occurring during these
cover-slip stages in an animal cell are described in fig 22.2.
797
\
nuclear membrane
tnterph
nterphase
Often mistakenly called resting stage. Variable duration depending upon chromatin threads
function of the cell. Period during which cell normally carries out synthesis
of organelles and increases in size. The nucleoli are prominent and actively _ nucleolus
synthesising ribosomal material. Just prior to cell division the DNA &
histone of each chromosome replicates. Each chromosome now exists as a centrioles
pair of chromatids joined together by a centromere. The chromosomal
material will stain and is called chromatin but structures are difficult to see. cytoplasm
cell membrane
Interphase
aster
Prophase
Usually the longest phase of division. Chromatids shorten (to 4% of their
original length) and thicken by spiralisation and condensation of the DNA centriole
protein coat. Staining shows up the chromatids clearly but the centromeres nuclear membrane
do not stain. The position of the centromere varies in different chromatid nucleolus
pairs. In animal cells and lower plants the centrioles move to opposite poles pair of chromatids
of the cell. Short microtubules may be seen radiating from the centrioles.
These are called asters (astra, a star). The nucleoli decrease in size as their centromere
nucleic acid passes to certain pairs of chromatids. At the end of prophase
the nuclear membrane disintegrates and a spindle is formed.
Prophase
Metaphase
The pairs of chromatids become attached to the spindle by spindle fibres
(microtubules) at their centromeres. The chromatids move upwards and
downwards along the spindle until their centromeres line up across the
‘equator’ of the spindle and at right-angles to the spindle axis. ame
centromere on
‘equator’ of spindle
Anaphase
This stage is very rapid. The centromeres split into two and the spindle
fibres pull the daughter centromeres to opposite poles. The separated
chromatids, now called chromosomes are pulled along behind the
centromeres.
daughter chromosomes
move apart led by their
centromeres
Telophase
pair of centrioles
The chromosomes reach the poles of the cell, uncoil, lengthen and lose the
ability to be seen clearly. The spindle fibres disintegrate and the centrioles nucleolus
replicate. A nuclear membrane forms around the chromosomes at each
pole and the nucleoli reappear. Telophase may lead straight into chromatin threads
cytokinesis (cell division). nuclear membrane
Fig 22.2 Summary notes and diagrams of the stages of

mitosis in an animal cell Telophase
798
Af
SF
Mitosis
Four stages of mitotic cell division from root-tip cells of the plant Crocus balansae (2n = 6). These are from squash
eee with only the stained chromosomes showing clearly. Details of nucleoli, spindles and cytoplasm are unstained and
not visible
At the start of division (1) the chromosomes are long, thin, but visibly double. They coil up and shorten. When fully
contracted the nuclear membrane breaks down and the chromosomes become orientated on the spindle (2: polar view). After a
while, spindle fibre activity on the centromeres (dotted in drawings) pulls the sister chromatids apart (3). Spindle fibres ;
pull centromeres and chromatids to the poles (4) with the formation of two identical polar groups. Following cell division these form
the nuclei of the two daughter cells
Fig 22.2 (continued) Photomicrographs and drawings based on them showing the stages in mitosis in plant cells.
(Courtesy of Dr S. A. Henderson, Department of Genetics, University of Cambridge)
799
22.2.1 Centrioles and spindle formation organelles become evenly distributed towards the two
poles of the telophase cells along with the chromosomes. In
Centrioles are organelles situated in the animal cells the cell membrane begins to invaginate during
cytoplasm close to the nuclear envelope in animal and telophase towards the region previously occupied by the
lower plant cells. They occur in pairs and are double spindle equator. Microfilaments in the region are thought
structures which lie at right-angles to each other. to be responsible for drawing in the cell membrane to form
Each centriole is approximately 500 nm long and 200 nm a continuous furrow around the equatorial circumference
in diameter and is composed of nine groups of microtubules of the cell. The cell membranes in the region of the furrow
arranged in triplets. Adjacent triplets are believed to be eventually join up and completely separate the two cells.
attached to each other by fibrils (fig 22.3). In plant cells the spindle fibres begin to disappear during
telophase everywhere except in the region of the equatorial
2S iniplet of
microtubules plane. Here they move outwards in diameter and increase
in number to form a barrel-shaped region known as the
phragmoplast. Microtubules, ribosomes, mitochondria,
endoplasmic reticulum and Golgi apparatus are attracted
to this region and the Golgi apparatus produces a number
of small fluid-filled vesicles. These appear first in the centre
of the cell and, guided by microtubules, coalesce to form a
‘cartwheel’
structure cell plate which grows across the equatorial plane (fig 7.23).
The contents of the vesicles contribute to the new middle
lamella and cell walls of the daughter cells, whilst their
Fig 22.3 (a) Electron micrograph of a TS of a centriole from membranes form the new cell membranes. The spreading
embryonic chick pancreas. (b) Diagram of TS through a
plate eventually fuses with the parent cell wall and
centriole as seen under the electron microscope
effectively separates the two daughter cells. The new cell
walls are called primary cell walls and may be thickened at
Centrioles also occur at the bases of cilia and flagella, a later stage by the deposition of further cellulose and other
where they are known as basal bodies (section 17.6.2). substances such as lignin and suberin to produce a
Spindle fibres are tubular and have a diameter of secondary cell wall (fig 22.4). In certain areas the vesicles of
approximately 25nm. They form during mitosis and the cell plate fail to fuse and cytoplasmic contact remains
meiosis, and are composed of microtubules made up of the between the daughter cells. These cytoplasmic channels
protein tubulin and other protein molecules. At one time it are lined by the cell (plasma) membrane and form
was thought that the function of the centrioles was to act as structures known as plasmodesmata.
organisers of the spindle fibres, but this is not now thought
to be the case. The majority of plant cells lack centrioles | secondary cell
wall of
but are capable of producing spindle fibres made up of daughter cell parental cell
microtubules identical to those found in animal cells. Some middle lamella primary cell
spindle fibres run from pole to pole, others are bound to the wall of
centromeres in bundles. It is thought that the relative parental cell
movement of these spindle fibres accounts for the separa- intercellular
tion of daughter chromosomes during anaphase of mitosis. S ~~ O
space
Electron microscopy provides evidence of the presence of
cross-bridges between the two types of fibre. These : plasmodesma r middle
primary cell walls \
cross-bridges suggest that the fibres may slide past each lamella
cell membrane___4
other during nuclear division using a ratchet-type mechan- |
ism similar to that associated with the movement of muscle daughter cell |
fibrils (section 17.4.6).
Fig 22.4 The structure of the plant cell wall formed as a
The addition of colchicine (section 23.9.1) to actively result of the cytokinesis of a parental cell
dividing cells inhibits spindle formation and the chromatid
pairs remain in their metaphase positions. This technique
enables the number and structure of chromosomes to be
22.2.3 Mitosis in animal and plant cells
examined under the microscope.
The most important event occurring during
22.2.2 Cell division
mitosis concerns the equal distribution of duplicate
chromosomes between the two daughter cells. This process
Cytokinesis is the division of the cytoplasm. is almost identical in animal and plant cells but there are a
This stage normally follows telophase and leads into the G, number of differences, and these are summarised in table
phase of interphase. In preparation for division, the cell 204s
800
Table 22.1. Differences between mitosis in plant and animal
cells
a
22.3 Meiosis
Plant Animal Meiosis (meio, to reduce) is a form of nuclear
oe
No centriole present division involving a reduction from the diploid number (2n)
Centrioles present
of chromosomes to the haploid number (n). In its simplest
No aster forms Asters form
terms it involves a single duplication of chromosomes
Cell plate forms No cell plate forms (DNA replication, as in mitosis) in the parent cell followed
No furrowing of cytoplasm at Furrowing of cytoplasm at by two cycles of nuclear divisions and cell divisions (the first
cytokinesis cytokinesis
meiotic division and the second meiotic division). Thus a
Occurs mainly at meristems Occurs in tissues throughout single diploid cell gives rise to four haploid cells as shown in
the body
ooo outline in fig 22.5.
Meiosis occurs during the formation of sperm and ova
(gametogenesis) in animals (sections 20.3.1 and 2) and
during spore formation in most plants (those showing
alternation of generations, section 20.2.2). Some lower
plants do not show alternation of generations and meiosis
The rate of mitosis varies in different organisms and occurs here at gamete formation. The stages of meiosis can
tissues, being greatest in bacteria and embryonic stages of be seen in spermatocyte nuclei from the testes of locust or
multicellular organisms and least in highly differentiated grasshopper or nuclei in immature pollen sacs of Crocus.
Like mitosis, meiosis is a continuous process but is
cells. Isolated cells taken from plants or animals and grown
conveniently divided into prophase, metaphase, anaphase
in nutrient culture solutions providing optimum conditions
and telophase. These stages occur once in the first meiotic
for cell division show the highest rate of mitosis. Such a
division and again in the second meiotic division. The
population of cells derived from a single parent cell is
behaviour of chromosomes during these stages is illus-
known as a clone (section 20.1.1). The cells found in a clone
need not be identical in structure or function. For example, trated in fig 22.6, which shows a nucleus containing four
chromosomes (2n=4), that is two homologous pairs of
single cells taken from an organism can give rise to a new
chromosomes.
organism or to a new tissue identical to that from which it
was taken, such as a single lung cell giving rise to lung tissue Diploid parental cell (with one pair of
(2n) homologous
having alveoli and ducts. chromosomes)
22.2.4 Significance of mitosis maternal in origin paternal in origin

The following points outline the main signi-
ficances of mitosis.
Each chromosome duplicates (DNA synthesis) forming two pairs of
Genetic stability. Mitosis produces two nuc- chromatids
lei which have the same number of chromosomes as the
parent cell. Moreover, since these chromosomes were
derived from parental chromosomes by the exact replica-
tion of their DNA, they will carry the same hereditary
information in their genes. Daughter cells are genetically
first meiotic ae
identical to their parent cell and no variation in genetic (Meiosis I)
information can therefore be introduced during mitosis.
This results in genetic stability within populations of cells
derived from parental cells, as in a clone.
Growth. The number of cells within an ee meiotic division
organism increases by mitosis (a process called hyperpla- (Meiosis IT)
sia) and this is a basic component of growth (chapter 21).
Asexual reproduction, regeneration and cell
replacement. Many animal and plant species are propa-
gated by asexual methods involving the mitotic division of
cells. These methods of vegetative reproduction are (n)
described more fully in section 20.1.1. In addition to oe sae
asexual reproduction, regeneration of missing parts, such Fig 22.5 The basic characteristics of meiosis showing one
as legs in crustacea, and cell replacement occurs, to varying chromosome duplication followed by two nuclear and cell
degrees, in multicellular organisms. divisions
801
Interphase
Variable length depending upon species. Replication of cell
organelles and increase in size of cell. Most of the DNA and
histone is replicated in premeiotic interphase but some is delayed
and prolonged into early meiotic prophase I. Each chromosome
now exists as a pair of chromatids joined together by a
centromere. Chromosomal material will stain but no structure
clear except prominent nucleoli, (see Fig. 22.2 as for mitosis).
nuclear membrane
Prophase I
The longest phase. It is often described in five stages called
leptotene, zygotene, pachytene, diplotene, and diakinesis but will
be considered here as a progressive sequence of chromosomal
changes.
(a) Chromosomes shorten and become visible as single
structures. In some organisms they have a beaded appearance due
to regions of densely stained material called chromomeres
alternating with non-staining regions. Chromomeres are regions
where the chromosomal material is tightly coiled.
(a) early prophase /
pair of homologous
(b) Chromosomes derived from maternal and paternal gamete chromosomes = a
nuclei come together and pair up. These are homologous bivalent
chromosomes. Each pair is the same length, their centromeres are
in the same positions and they usually have the same number of
genes arranged in the same linear order. The chromomeres of the
homologous chromosomes lie side by side. The pairing process is
called synapsis and it may begin at several points along the
chromosomes which then completely unite as if zipped up centromeres
together. The paired homologous chromosomes are often
described as bivalents. The bivalents shorten and thicken. This
involves both molecular packaging and some visible coiling (or
spiralisation). Each chromosome and its centromeres can now be
seen clearly.
bivalent
Fig 22.6 (a) —(k) Summary notes and diagrams of the stages (b) prophase |
of meiosis in an animal cell.
802
(c) The homologous chromosomes of the bivalents now fall
apart and appear to repel each other partially. Each chromosome
Is NOW seen to be composed of two chromatids. The two pair of sister
chromosomes are seen to be joined at several points along their chromatids
length. These points are called chiasmata (chiasma, a cross). It can
be seen that each chiasma is the site of an exchange between
chromatids. It is produced by breakage and reunion between two
of the four strands present at each site. Asa result, genes from one
chromosome (e.g. paternal, A, B, C) become attached to genes
from the other chromosome (maternal, a, b, c) leading to new
gene combinations in the resulting chromatids. This is called
genetic crossing over. The two chromosomes do not fall apart after
crossing over (chiasma formation) because sister chromatids (of
both chromosomes) remain firmly associated until anaphase.
pair of sister
chromatids
(c) crossing over during prophase 1
——
rotation
(d) (i) Bivalent with a single chiasma

(d) The chromatids of homologous chromosomes continue to Rotation of arms usually occurs during diplotene so that the
repel each other and bivalents assume particular shapes X-shaped chiasma at the start of chiasma formation ends up
depending upon the number of chiasmata. Bivalents having a looking like an open cross.
single chiasma appear as open crosses, two chiasmata produce a
ring shape and three or more chiasmata produce loops lying at
right angles to each other. By the end of prophase I, all
chromosomes are fully contracted and deeply stained. Other
changes have occurred within the cell including: the centrioles
(if present) migrate to the poles, the nucleoli and nuclear
membrane break down and then the spindle fibres form.
(d) (ii) Bivalent with double chiasmata

Rotation of the arms produces a ring shape.
803
spindle fibres
Metaphase I
The bivalents become arranged across the equatorial plate of the
spindle. Their centromeres (though often visibly double) behave
as though single and organise spindle fibres pointing towards only
one of the poles. Gentle pulling from these fibres places each
bivalent on the equator, with each centromere equidistant above
and below it.
cell membrane
(e) ate metaphase |
Anaphase I
The two centromeres of each bivalent do not divide, but sister
chromatid adhesion ends. Spindle fibres pull whole centromeres,
each attached to two chromatids, towards opposite poles of the
spindle. This separates the chromosomes into two haploid sets of
chromosomes in the daughter cells.
(f) anaphase |
Fig 22.6 (continued)
804
Telophase I
The arrival of homologous centromeres and their pairs of
chromatids at opposite poles marks the end of the first meiotic
division. Reduction of chromosome number has occurred but
each pole possesses chromosomes composed of two chromatids.
As a result of crossing over, or chiasma formation, these chromatids
arenot genetically identical and must be separated in the second
meiotic division. Spindles and spindle fibres usually disappear.
In animals and some plants the chromatids usually uncoil and a
nuclear membrane forms at each pole and the nucleus enters
interphase. Cleavage (animals) or cell wall formation (plants) then
occurs as in mitosis. In many plants there is no telophase, cell wall
formation or interphase and the cell passes straight from anaphase
I into prophase of the second meiotic division.
Interphase II
This stage is present usually only in animal cells and varies in
length. There is no S-phase and no further DNA replication
occurs. The processes involved in the second meiotic division are
mechanically similar to those of mitosis. They involve separation
of the chromatids of both daughter cells produced during the first (g) telophase /in an animal cell
meiotic division. The second meiotic division differs from mitosis
)
mainly in that (a) these sister chromatids are often widely separated
at metaphase II of meiosis and (b) the haploid number of
chromosomes is present.
‘
S ‘ 9
Prophase II
This stage is absent from cells omitting interphase II. The length of
the stage is inversely proportional to the length of telophase I. The
nucleoli and nuclear membranes break down and the chromatids
shorten and thicken. Centrioles, if present, move to opposite
poles of the cells and spindle fibres appear. The chromatids are
arranged with their axes at right angles to the spindle axis of the
first meiotic division.
(h) prophase II
(
805
Metaphase II
At this division the centromeres now behave as structurally
double. They organise spindle fibres on each side to both poles and
hence become aligned on the equator of the spindle.
(i) metaphase //
SS
(k) stages following telophase II and cell cleavage in animal
(j) anaphase I/ cell
Anaphase II Telophase II
The centromeres divide and the spindle fibres pull the centromeres This stage is very similar to that found at mitosis. The
to opposite poles. The separated chromatids, now called chromosomes uncoil, lengthen and become very indistinct. The
chromosomes, are pulled along behind the centromere. spindle fibres disappear and the centrioles replicate. Nuclear
membranes re-form around each nucleus which now possesses
half the number of single chromosomes of the original parent cell
(haploid). Subsequent cleavage (animals) or cell wall formation
(plants) will produce four daughter cells from the original single
Fig 22.6 (continued) parent cell.
806
Fig 22.7 (a) Eight stages of meiosis
from spermatocytes of the desert
locust Schistocerca gregaria (2n = 22
+ X 0’). As at mitosis, these are from
squash preparations with only the
stained chromosomes showing clearly.
Details of nucleoli, spindles and
cytoplasm are unstained and not
visible. At the start of the first meiotic
division (1) the chromosomes are
long, thin, fuzzy and not clearly
visible, while the X is deeply staining.
Pairing of homologous (maternal and
paternal) chromosomes takes place
with the formation of the haploid
number (11) of bivalents and the X
(2,3). Molecular exchange occurs
between maternal and paternal
chromosomes when intimately paired.
They then fall apart along their length
except at points where exchanges
took place (4). These are called
chiasmata. (part (b) overleaf)
807
YS
an ¢
A
Fig 22.7 (b) Further contraction

produces short, fat bivalents which,
following breakdown of the nuclear
membrane, become attached to the
spindle by their centromeres (5: polar
view). Spindle fibre activity pulls the
haploid number of chromosomes apart
(6: side view). The X, which is now
weakly staining, goes undivided to
one pole. At the second meiotic
division the haploid number of
chromosomes lie on the equatorial
plate (7: polar view) and single
chromatids are pulled to the poles by
spindle fibre activity (8: side view)
(Photomicrographs and
drawings by courtesy
of Dr S. A. Henderson,
Department of Genetics, University
of Cambridge)
808
=
l).
wae
x ctraide te
—
SS
a
7
4 N Pa
Vf mew enre te eG
a anc
Fig 22.8 Diplotene (prophase |) showing chiasmata. The chromosomes from a single cell at diplotene in the locust Locusta
migratoria, showing bivalents with one or two chiasmata. Maternal and paternal chromosomes are represented by solid and dotted
lines in the drawing. At each chiasma a genetic exchange has occurred. The shape of the bivalent will vary from rod-shaped, to
cross-shaped or ring-shaped, depending on the number and position of chiasmata. The unpaired X chromosome is deeply
staining at this stage (Dr S. A. Henderson)
Fig 22.9 Meiosis in living

cells. Pairing and cell division
at meiosis in living
spermatocytes of the locust
Locusta migratoria. These
preparations are photographed
with an optical technique known
as Nomarski interference
contrast, which uses polarised
light and produces images of
remarkably 3-D appearance in
living, unstained cells. Two cells
show chromosome pairing in
early prophase | nuclei, with
chromomere alignment
(arrowed). The other two cells
are at the end of the first meiotic
division. It can be seen that,
after the two polar groups have
formed, cleavage of the cell
takes place and two
approximately equal daughter
cells are produced. The fibrous
structures stretching between
the two polar groups, pinched in
two like a wheat sheaf at
cleavage, are the mitochondria
809
22.3.1 Significance of meiosis II are random. The subsequent separation of these
during anaphase I and II respectively produces new
The following points outline the main signi- allelic recombinations in gamete cells. This is called
ficances of meiosis. independent assortment and results in the random
Sexual reproduction Meiosis results in the assortment of maternal and paternal chromosomes
formation of four daughter cells, each with half the number between daughter nuclei. This is the basis of Mendel’s
of chromosomes of the parent cell in all organisms carrying second law (section 23.1.3).
out sexual reproduction. During fertilisation the nuclei of (3) As a result of chiasmata, crossing over of segments of
the two gamete cells fuse and produce a zygote which has a chromatids occurs between homologous chromosomes
fixed number of chromosomes for each species. In all during prophase I, leading to the formation of new
organisms this number of chromosomes represents the combinations of alleles on chromosomes of the gamete
diploid condition (2n). If meiosis did not occur fusion of cells. This breaks established linkage groups and
gametes would result in a doubling of the chromosomes for produces new ones.
each successive sexually reproduced generation. (An The significance of these three mechanisms in the
exception to this is shown in polyploidy which is described process of inheritance and the production of variation is
in section 23.9.) This situation is prevented in the life cycle described in detail in section 23.8.4.
of all sexually reproducing organisms by the occurrence, at
some stage, of cell division involving a reduction in the 22.3.2 Similarities and differences between
diploid number of chromosomes (2n) to the haploid mitosis and meiosis
number (7). The main similarities between mitosis and
meiosis involve the mechanisms by which the chromo-
22.1 The amount of DNA present per cell somes and other cell organelles replicate and are man-
during several nuclear divisions is represented oeuvred within the cell prior to and during cell division.
diagrammatically in fig 22.10. The mechanism of cell division, too, is similar in mitosis
(a) Which type of nuclear division is represented by and meiosis.
~ fig 22.10? — The differences between the two processes are shown in
_ (b) What phases are represented by the dashed table 22.2.
lines W, X and Y? -. Differences between males and females, animals and plants
_ (c) What type of cells are represented by theline Z? _ The above description of meiosis applies in a general way to
all animals and plants, and males and females, but there are
important differences also.
4n
(a) Males
In animals, cleavage occurs at the end of both first and
DNA content
per cell an second meiotic divisions and the daughter cells separate
immediately and behave independently thereafter. All
nA
four meiotic products survive, form spermatids and
develop into spermatozoa. In plants, cell wall formation
occurs and the daughter cells remain associated at the end
Fig 22.10 Diagram for use in question 22.1 of the first meiotic division to form a diad, and at the second
division, to form a tetrad. All products again survive to give
rise to pollen grains.
(b) Females
Genetic variation Meiosis also provides Female meiosis in both plants and animals ends up with
opportunities for new combinations of genes to occur in the three of the four meiotic products lost — only one survives
gamete cells. This leads to genetic variation in the genotype to form an egg nucleus.
and phenotype of offspring produced by the fusion of In animals, first division cleavage is unequal and results
gametes. The mechanisms of meiosis that contribute to this in the production of a primary oocyte and a polar body.
variation may be summarised as follows: Second division is also unequal and the oocyte gives rise to
(1) The reduction of chromosomes from the diploid to the a secondary oocyte and a second polar body (the first polar
haploid number segregates (separates) alleles so that body may also divide and all three polar bodies abort).
each gamete cell carries only one allele for a particular Only one functional oocyte is produced. In plants, a linear
gene locus (section 23.2). meiosis results in four nuclei lying in the embryo sac.
(2) The orientations on the equatorial spindle of bivalents Three abort and only one remains to give rise to the embryo
during metaphase I and of chromosomes in metaphase sac nuclei and the egg nucleus.
810
Table 22.2. Differences between stages of mitosis and meiosis
Ee eee eee
SaanannInRImcisaramimememsmeesssmneeee
Mitosis Meiosis
a eee ee Ree Nek a llee Bt Be he fe ee
Prophase Chromomeres not visible Chromomeres visible
Homologous chromosomes remain separate Homologous chromosomes pair up
No chiasmata formation Chiasmata occur
No crossing over Crossing over may occur
Metaphase Pairs of chromatids line up on the equator Pairs of chromatids line up on the equator only in
of the spindle second meiotic division
Centromeres line up in the same plane on the Centromeres lie equidistant above and below the
equator of the spindle equator in the first meiotic division
Anaphase Centromeres divide Centromeres divide only during second meiotic
division
Chromatids separate Chromatids separate only in the second meiotic division.
In the first division whole chromosomes separate.
Separating chromatids Separating chromosomes may not be identical due
identical to crossing over
Telophase Same number of chromosomes present in Half the number of chromosomes present in
daughter cells as parent cells daughter cells
Both homologous chromosomes present in Only one of each pair of homologous chromosomes
daughter cells if diploid present in daughter cells
Occurrence May occur in haploid, diploid or polyploid cells Only occurs in diploid or polyploid cells
Occurs during the formation of somatic cells and In formation of gamete cells or spores
some spores. Also occurs during the formation
of gametes in plants with alternation of generations
22.4 The structure of chromosomes
Histochemical and cytological analyses of Single DNA double

: helix chromomeres
chromosomes of eukaryotic cells have shown them to be
composed of deoxyribonucleic acid (DNA) and protein,
with small amounts of chromosomal RNA. The ‘chromo-
somes’ of prokaryotic cells (bacteria and blue-green algae)
are composed of DNA only and strictly should not be called
chromosomes. DNA has negative charges distributed
along its length, and positively charged (basic) protein
molecules called histones are bonded to it. This DNA
protein complex is called chromatin. The protein mole-
cules increase the diameter of the chromosome and may act
as an outer protective coat for the DNA. The exact
structural relationship between the DNA and histones in a
chromosome is still far from certain, but it would appear (o) single DNA
that a chromosome is not simply a single molecule of DNA double helix
surrounded by a layer of histones, but something much
more. Several models of the structure of chromosomes chromomere
have been proposed.
The diameter of a single DNA double helix is 2 nm, several
coiled DNA
whereas the thinnest chromosomes seen with the light strands
microscope are between 100 and 200 nm in diameter. The
explanation may be either that a chromosome contains
several DNA double helices or that one long DNA double
helix is tightly coiled. Electron micrographs taken of Fig 22.11 (a) Lampbrush chromosomes of amphibian oocyte
lampbrush chromosomes (so-called because of their re- showing centromere and three chiasmata. (b) and (c) The
effects of stretching lampbrush chromosomes to show the
semblance to brushes used to clean the glass of oil lamps) of
central filament of DNA and the loops of DNA where mRNA
amphibian oocytes during metaphase show that each synthesis takes place. The dense regions are thought tobe
chromatid appears to be composed ofa tightly coiled axis chromomeres. Each chromomere and its associated loop is
with several lateral loops made up of a single DNA double thought to be associated with a specific gene locus. (From
helix (fig 22.11). These loops may represent DNA which H. G. Callan (1963) Int. Rev. Cytology, 15, 7.)
811
has been exposed for the purpose of transcription (section Table 22.3. Results of Griffith’s experiments
22.6.6). Recent investigations into the structure of chromo- Injected form of Pneumococcus Effect
somes suggest that the DNA helix combines with groups of mice live
eight histone molecules to form structures known as Live non-capsulated
Live capsulated mice die
nucleosomes having the appearance of ‘beads onastring’. Heat-killed capsulated mice live
These nucleosomes, and the DNA strands linking them, Heat-killed capsulated +
are packed closely together to produce a 3 nm diameter live non-capsulated mice die
helix with about six nucleosomes per turn which has the
Sooo!
dimensions and other features of a chromosome. The

proposed structure of the chromosome is shown in fig It was suspected that protein might be the only molecule
Zale: with sufficient variety of structure to act as genetic
In the chromosomes of sperm of some species, such as material.
salmon and herring, protamines replace histones as the Frederick Griffith, an English bacteriologist, made an
proteins bound to the DNA. observation in 1928 which was later to prove significant in
resolving the problem. In the days before the development
<<. 36nm ————_> of antibiotics, pneumonia was often a fatal disease. Griffith
was interested in developing a vaccine against the bacte-
rium Pneumococcus which causes one form of pneumonia.
Two forms of Pneumococcus were known, one covered
with a gelatinous capsule and virulent (disease-producing)
and the other non-capsulated and non-virulent. The ability
of these bacteria to cause pneumonia appeared to be
surface view of helix related to the presence of the gelatinous capsule. Griffith
of nucleosomes
hoped that by injecting patients with the non-capsulated,
or heat-killed encapsulated forms, their bodies would
produce antibodies which would afford protection against
pneumonia. In a series of experiments Griffith injected
mice with both forms of Pneumococcus and obtained the
results shown in table 22.3. Post mortems carried out on the
single DNA dead mice revealed the presence within their bodies of live
lecul encapsulated forms. On the basis of these results Griffith
aie re of eight
histone molecules concluded that something must be passing from the
heat-killed encapsulated forms to the live non-capsulated
forms which caused them to develop capsules and become
virulent. However, the nature of this transforming prin-
ciple, as it was known, was not isolated and identified until
1944. For ten years Avery, McCarty and McCleod analysed
and purified the constituent molecules of heat-killed
Fig 22.12 Proposed structure of the nucleosome and its encapsulated pneumococcal cells and tested their ability to
relationship to the chromosome and the DNA molecule. (From
E. J. Du Praw School of Medicine, University of Maryland,
bring about transformation in live non-encapsulated cells.
(metaphase chromosome). ) Removal of the polysaccharide capsule and the protein
fraction from the cell extracts had no effect on transforma-
tion, but the addition of the enzyme deoxyribonuclease
22.4.1 Evidence for the role of DNA in
(DNAase), which breaks down (hydrolyses) DNA,
inheritance
prevented transformation. The ability of extremely
Following the proposal by Sutton and Boveri purified extracts of DNA from encapsulated cells to bring
at the beginning of this century that chromosomes were the about transformation finally demonstrated that Griffith’s
structures by which genetic information passed between principle was in fact DNA. Despite this evidence many
generations (section 23.2), it took many years to clarify scientists still refused to accept that DNA, not protein, was
whether the genetic material was the DNA or the protein of the genetic material. In the early 1950s a wealth of
the chromosomes. In a series of experiments, Alfred irrefutable evidence, based upon the study of viruses,
Mirsky demonstrated that all the somatic cells of a given eventually demonstrated the ubiquitous nature of DNA as
species contain the same amount of DNA, and this is the carrier of hereditary information.
double the amount found in gamete cells. However, the Viruses became one of the major experimental materials
same applies to the protein content of the chromosomes so in genetic research in the 1940s and since then experiments
this knowledge did little to clarify the nature of the genetic involving their use have become as classic as those
material. involving the garden pea, the fruit fly and, as will be
812
described later, the bread mould Neurospora. Virus radioactivity. The results are shown in fig 22.13.
particles have an extremely simple structure consisting of a On the basis of these results Hershey and Chase
mainly protein coat enclosing a molecule of nucleic acid, concluded that it was the phage DNA which entered the
either DNA or RNA (section 2.5.2). As such they provided bacterial cell and gave rise to large numbers of phage
ideal research material to investigate whether protein or progeny. These experiments had demonstrated that DNA
nucleic acid is the genetic material. In 1952 Hershey and is the hereditary material. Confirmatory evidence that only
Chase began aseries of experiments involving a particular the DNA contained within the phage is introduced into the
type of virus which specifically attacks bacterial cells and is bacterial cell has been provided by electron microscopy
called a bacteriophage. Bacteriophage T, attacks the colon and increased knowledge of the life cycle of viruses. (The
bacillus Escherichia coli (E. coli) and causes it to produce life cycle of virus and phage particles is described in
large numbers of T,-phage particles in a very short time. sections 2.5.3 and 2.5.4.)
The essence of Hershey and Chase’s experiment involved
growing T,-phage particles in E. coli which had been grown
22.4.2 DNA replication
on a medium containing radioactive isotopes of either
sulphur (*S) or phosphorus (’P). The phage particles The double helical structure of DNA, as
formed in E. coli labelled with radioactive sulphur had determined by Watson and Crick, is described in section
incorporated this into their protein coats, whereas those 5.6.3. One of its most attractive features is that it
formed in phosphorus-labelled E. coli contained radio- immediately suggests a method by which replication could
actively labelled 32P DNA. This selective distribution of occur. Watson and Crick proposed that the two strands
isotopes is due specifically to the fact that proteins do not were capable of unwinding and separating, and acting as
contain phosphorus and nucleic acids do not contain templates to which a complementary set of nucleotides
sulphur. The labelled T,-phage particles were allowed to would attach by base pairing. In this way each original
infect non-radioactively labelled E. coli and after a few DNA molecule would give rise to two copies with identical
minutes the cells were agitated in a blender or liquidiser structures.
which stripped off the phage particles from the bacterial In 1956 Kornberg succeeded in demonstrating the in
walls. The bacteria were then incubated and examined for vitro synthesis of a DNA molecule using a single strand of
T, phage
protein coat
DNA
infect E.coli grown in

( infect E.coli grown in
medium containing **P medium containing **S
radioactivity only in
Y radioactivity only in
32P of DNA 3° of sulphur coat
phage particles attach

to wall of E.coli
phage particles ‘inject’

DNA
phage particles are

stripped off E.coli by
mechanical action
no trace of
this cell emits
radioactivity
radioactivity in this cell
from DNA
both bacterial cells

produce new phage
particles
Fig 22.13 Summary diagrams of Hershey and Chase’s work on Tz phage and E. coli
813
DNA as a template. (This technique was used by later Kornberg later demonstrated, in 1967, that DNA
researchers to investigate the nature of the genetic code polymerase only adds nucleotides to the strand running in
(section 22.5).) Kornberg extracted and purified an the 5-3 direction. Since the two DNA strands lie in
enzyme from E. coli which was capable of linking free opposite directions, that is are anti-parallel as shown in fig
DNA nucleotides, in the presence of ATP as an energy 22.12, DNA polymerase can only continuously produce
source, to form a complementary strand of DNA. This one new DNA molecule at a time. Short sections of the
enzyme he named DNA polymerase. In subsequent other daughter DNA molecule are produced by the action
experiments Kornberg used nucleoside triphosphates of DNA polymerase moving in the opposite direction.
(ATP, GTP, TTP, CTP) instead of nucleotides and ATP, These short sections of newly synthesised polynucleotide
and they too produced a complementary strand of DNA. chains are joined together by the action of another enzyme
Later evidence confirmed that this is the form in which called DNA ligase (fig. 22.15). DNA molecules produced
nucleotides readily attach themselves to the DNA template by in vitro synthesis in the presence of DNA ligase are
and each other. As the nucleoside triphosphates link up, biologically active and can be used in protein synthesis.
the two terminal phosphate groups are broken off. The Those DNA molecules produced in 1956 by Kornberg,
remaining phosphate group of the nucleotide and the whilst having the correct DNA structure, were biologically
energy released are used to form the ester linkage between inactive due to the failure of sections of the synthesised
the 5—3 carbon atoms of the sugar molecules of adjacent 5—3 chain to connect up.
nucleotides (fig. 22.14). The method of DNA replication proposed by Watson
parental DNA double helix
and Crick and shown in fig 22.16 is known as semi-
conservative replication since each new double helix retains
one strand of the original DNA double helix. The evidence
for this mechanism was provided byaseries of classic
experiments carried out by Meselsohn and Stahl in 1958.
E. colihas asingle circular chromosome, and when cultures
direction of
of these cells were grown for many generations ina medium
movement of containing the heavy isotope '°N all the DNA became
DNA polymerase
labelled with '°'N. These cells containing DNA labelled
DNA polymerase with N were transferred to a culture medium containing
the normal isotope of nitrogen ('*N) and allowed to grow.
After periods of time corresponding to the generation time
eres QO
for E. coli (SO min at 36 °C) samples were removed and the
DNA extracted and centrifuged at 40 000 times gravity for
20h in a solution of caesium chloride (CsCl). During
free
nucleoside
centrifugation the heavy caesium chloride molecules began
free nucleoside ae triphosphates to sediment to the bottom of the centrifuge tubes producing
triphosphates
an increasing density gradient from the top of the tube to
the bottom. The DNA settles out where its density equals
CO icleascc
that of the caesium chloride solution. When examined
CG phosphate under ultra-violet light the DNA appeared in the centrifuge
A-[; tube as a narrow band. The positions of the bands of DNA
VL
template A-K
extracted from cells grown in °N and “‘N culture media and
§
trand G
iO new strand
the interpretation of their structure are shown in fig 22.16.
These experiments conclusively demonstrated that DNA
DST,
<M replication is semi-conservative.
Se eee 22.2 There were three “hypotheses

pair of identical DNA double helices
advanced to explain the process of DN, replication.
Fig 22.14 Replication of the DNA double helix. The parental One of these is known as semi-cor tive and is _
DNA helix unwinds and the complementary strands separate described above and in fig 22.14.
due to the breaking of weak hydrogen bonds between
The other hypo-
complementary bases. The DNA polynucleotide chains do not theses are known as conservative replication and
break due to the strength of their phosphodiester linkages. sensi replication and are summarised in fig
Base pairing occurs between the bases of the template 17
strands and free nucleoside triphosphates link up to form a Draw diagrams to show the distribution of the
polynucleotide chain. In this way two identical DNA molecules different types of DNA in a density gradient which
are produced. The enzyme DNA polymerase is involved in the
separation of the parental strands and the formation of the Meselsohn and Stahl would have found in the first
new polynucleotide chain but other enzymes are also thought two pee ifthese latter hypotheses had oe) .
to influence replication correct. | .
814
Fig 22.15 (above) The role of DNA ligase
in DNA replication. The arrows show the cells from A grown in '!N
direction in which DNA polymerase moves
in producing new DNA double helices. DNA cells grown Ist 2nd 3rd
in °N generation generation generation
ligase is active in joining short sections of
polynucleotide chains at the points labelled
with an asterisk caesium
chloride
density eee —-'4N-containing DNA
gradient
Fig 22.16 (right) The results and —-4N/N- containing DNA
interpretation of Meselsohn and Stahl’s — ->N- containing DNA
experiment into the process of DNA
replication. The widths of the DNA bands in A B Cc D
the centrifuge tubes reflect the proportions
of the various types of DNA molecules as ROOK OK OSS Sere
shown in the diagrams (right). In C the ratio allstrands _ alll strands all '*N
) ratio is 3:1
of the width is 1:1 and in D the contain *N contain '*N/?°N SOOO COCCST F strands
(heavy) DNA ‘hybrid’ DNA one half of the
strands contain
SN (light) DNA, --!4N strand
Fig 22.17 (below) Diagrams explaining the other half
~<)N strand
contain ‘hybrid’
two further theories of DNA replication DNA
were later termed genes and shown to be located on, and

transmitted from generation to generation by chromo-
parental DNA somes (section 23.2).
double helices Despite our current knowledge of chromosomes and the
structure and function of DNA, it is still extremely difficult
to give a precise definition of a gene. Investigations into the
nature of the gene have so far produced three possible
definitions of a gene and these are outlined below, since
Q OQ
they have implications in the fields of genetics and
evolution.
ees.
”, double helices
° A unit of recombination. From his studies
of chromosome mapping in Drosophila (section 23.3),
Bee ©
Morgan postulated that a gene was the shortest segment of a
chromosome which could be separated from adjacent
© Se
segments by crossing-over. This definition regards the gene
as a large unit, a specific region of the chromosome
conservative replication dispersive replication determining a distinct characteristic in the organism.
A unit of mutation. Investigations into the
nature of mutations revealed that changes in characteristics
occurred as a result of random and spontaneous alterations
22.5 The nature of genes in the structure of a chromosome, a sequence of bases or
even a single base (section 23.9). It seemed therefore that a
The particulate nature of heredity has always gene might be as small as a single pair of complementary
bases in the nucleotide sequence of DNA, that is the
been a feature of the study of inheritance. Mendel
shortest segment of a chromosome which could undergo
proposed in 1866 that the characteristics of organisms were
mutation.
determined by hereditary units called ‘elementes’. These
815
Aunitoffunction. Since genes were known to Neurospora is an ascomycete fungus and has the following
determine structural, physiological and biochemical char- advantages for genetic research:
acteristics of organisms it was suggested that a gene was the (1) it is easy to grow,
shortest segment of a chromosome responsible for the (2) it can be bred in large numbers,
production of a specific product. (3) it has a very short life cycle (10 days),
(4) it has a haploid vegetative stage.
The third definition is the most acceptable but it lacks The latter point is very significant. Since only one set of
precision concerning the nature of the specific product. In chromosomes is present for most of the life cycle there is no
some cases one ‘gene’ is known to affect several character- masking of recessive genes. If a mutation arises and
istics (section 23.1.1) whilst in other cases several ‘genes’ produces a recessive gene the effects of the mutation will be
(polygenes) may determine one specific characteristic seen immediately. Neurospora normally produces asexual
(section 23.7.6). spores (conidia) which germinate and give rise to a
It was, however, as a result of research carried out by mycelium. Segments of mycelia of opposite mating strains
Beadle and Tatum that the third definition of the gene may fuse and produce a diploid zygote. This immediately
gained recognition. This became known as the one gene, undergoes a meiotic division followed by a single mitotic
one enzyme hypothesis and was largely substantiated by the division to form an ascus or fruiting body containing eight
development of a new field of biology called molecular haploid ascospores, four of these ascospores being derived
genetics. (Subsequent research has narrowed this function- from each parental strain. Each ascospore may germinate
al concept to that of the one cistron, one polypeptide and produce a new mycelium. A mycelium can therefore
hypothesis (section 22.5.1).) arise from an asexually produced conidium or a sexually
produced ascospore (fig 22.18).
22.5.1 Genes and enzymes
The relationship between genes and enzymes
was first suggested, although not in those terms, by an
English physician, Sir Archibald Garrod in 1908. Garrod
postulated that certain ‘inborn errors of metabolism’ were
the result of the failure of the body to produce certain
important chemical substances which, in turn, were
determined by hereditary mechanisms. It was almost 40
years later that the full extent of this hypothesis was
demonstrated and found to be true as a result of the
pioneer work in molecular genetics.
Initial investigations in molecular genetics involved
establishing the various compounds and enzymes of
metabolic pathways. Certain metabolic disorders produce
defects in organisms which behave as though controlled by
single genes. They may appear spontaneously, as in the
case of mutations, or be inherited according to convention-
al genetic mechanisms. For example, the amino acids
phenylalanine and tyrosine are normally used in the
manufacture of cell proteins and other structural and
physiological compounds, and the excess is broken down
into carbon dioxide, water and nitrogenous wastes. In all
cases their fate is determined by simple metabolic pathways
involving a series of enzymes. Defective enzymes or lack of
enzymes at four points in these pathways produce the
metabolic disorders phenylketonuria, goitrous cretinism,
albinism and alkaptonuria. In all four cases these disorders
Fig 22.18 Ascospores in an ascus
are inherited in such a way as to suggest that they are each
controlled by a single recessive gene.
Further important evidence supporting the one gene, Neurospora can grow on a culture medium containing
one enzyme hypothesis was gained from work begun by only agar, sugars, salts and the vitamin biotin. This is
Beadle and Tatum in 1941, investigating the mechanisms of known as minimal medium. Growth under these conditions
inheritance of enzymes in the pink bread-mould Neuro- demonstrates that Neurospora is able to synthesise all the
spora crassa. In common with Mendel and Morgan they carbohydrates, fats, amino acids and vitamins essential for
were careful in their choice of experimental organism. its growth using enzymes produced by its cells.
816
Beadle and Tatum carried out the following procedure in strain, the other half were able to grow in minimal
their investigations: medium.
(1) Conidia were exposed to X-rays in order to increase the This suggested that the mutant gene behaved asasingle
mutation rate. recessive gene and was transmitted according to the rules of
(2) These conidia were transferred to complete medium normal Mendelian inheritance. Beadle and Tatum inter-
(containing all the necessary amino acids and vitamins preted these results as demonstrating that, in each case, the
for normal growth) and grown. X-rays had induced a mutation in one gene controlling the
(3) The mycelia which formed were then crossed with production of one enzyme necessary in the synthesis of an
mycelia from conidia not exposed to X-rays (‘wild amino acid. This formed the basis of their ‘one gene, one
type’). enzyme’ hypothesis (fig 22.19).
(4) The asci produced contained four ascospores from each This early work established the experimental procedure
parental mycelial strain (mutant and wild type). of using minimal media which has been used and modified
(5S) The ascospores were dissected out and separately in microbial genetics to produce a vast amount of
transferred to complete medium where they all grew. information concerning the role of genes.
(6) Samples of mycelia were placed on vitamin-enriched It is now clearly established that genes exercise control
minimal medium. In some cases no growth occurred. within the cell by the synthesis of enzymes and_ other
(7) The strains which did not grow were unable to proteins. These enzymes in turn direct the synthesis of all
synthesise certain amino acids. In order to determine other materials within the cell.
which amino acids could not be synthesised, these Over the years, however, the exact definition of the gene
strains were then transferred to a series of minimal has been modified. Research carried out by Benzer in 1955
media cultures each containing a single different amino into the structure of genes in the bacteriophage T, gave rise
acid. to the concept of the cistron as a unit of function. A cistron
(8) The specific medium in which growth occurred there- is aregion of DNA carrying information for the production
fore contained the amino acid which the strain of of a polypeptide chain. This may function on its own as a
Neurospora was unable to synthesise. In this way biologically active molecule or may form part of a larger
mutant strains of Neurospora were identified. Analysis macromolecule. Current views on the nature of the gene
of the results showed that in all cases where half of the have moved from the ‘one gene, one enzyme’ concept to
ascospores from a given ascus produced a mutant the ‘one cistron, one polypeptide’ concept.
X-rays
CONG 1a CGV ES 2a a,
(2) complete medium (3) complete medium

(1) (mutant strain) (wild-type)
(4) 00000008 ascus
ascospores
<a
(5) each ascospore transferred to complete
medium where they all grow
sample of each mycelium minimal medium

transferred to minimal medium: complete medium
minimal medium (all ascospores (growth, therefore
(no growth, therefore growth indicates wild type, no wild type)
growth indicates mutant strain produce mycelium)
mutant strain)
1 ase oe
yy
Se aT kg er ly Mee Ce
NRAYERY Ye Yocov any

(7)
mutant strains transferred to 20
minimal media each containing
a single amino acid (8)
this strain lacks the enzyme(s)
capable of producing valine
+ alanine + valine
(no growth) (growth)
! s involved
ing the synthesis of enzyme
identifyi ing mutant alleles controlling
; i
Flow diagra m ji
illustrating jt steps inin identify
j the main
on the diagram correspond with those in the text.)
poe ae ore icdon Full details are given in the text. (The numbers
817
22.5.2 The genetic code 7 228 Using different pairs of the bases/ co
When Watson and Crick proposed the double G, T and Clist the 16 possible combinations ofbases —
helical structure for DNA in 1953 they also suggested that that can be produced.
the genetic information which passed from generation to
generation, and which determined cell metabolism, might Only a code composed of three bases could incorporate all
reside in the sequence of bases of the DNA molecule. Once 20 amino acids into the structure of protein molecules.
it had been demonstrated that DNA only specified the Such a code would produce 64 combinations of bases.
production of protein molecules it became clear that the
DNA nucleotide base sequence must determine the amino 22.4 If four bases usedsingly would code
acid sequence of protein molecules. This relationship for four amino acids, pairs of bas 16 amino
between bases and amino acids is known as the genetic acids and triplets of bases code mino acids,
code. The problems remaining were to demonstrate that a deduce a mathematical expression to explain this
nucleotide code existed, to break the code and determine situation. .
how the code is translated into the amino acid sequence of a
protein molecule. Evidence for the existence of a triplet code was provided by
Francis Crick in 1961 by producing mutations involving the
addition or deletion of bases in T, phages. These additions
22.5.3 The code and deletions which produced frame-shifts in the code, as
There are four nucleotide bases in the DNA shown in fig 22.20, were expressed in T, phages as
molecule, adenine (A), guanine (G), thymine (T) and mutations. These frame-shifts produced base triplet se-
cytosine (C) (section 5.6). They are arranged as a quences which failed to result in the synthesis of protein
polynucleotide strand that can be indicated by the initial molecules with the original amino acid sequence (primary
letters of the bases. This ‘alphabet’ of four letters is structure). Only by adding a base and deleting a base at
responsible for carrying the code that results in the specific points could the original base code sequence be
synthesis of a potentially infinite number of different restored. Restoring the original base sequence prevented
protein molecules. If one base determined the position of a the appearance of mutants in the experimental T, phages.
single amino acid in the primary structure of a protein, the These experiments also demonstrated that the code is
protein could only contain four amino acids. If a combina- non-overlapping, that is to say, each triplet is discrete. No
tion of pairs of bases coded for each amino acid then 16 base of a given triplet contributes to part of the code of the
amino acids could be specified into the protein molecule. adjacent triplet (fig 22.21).
Normal triplet code
SSC AE ek
I
ee ee
|
ee ne|
| | I
Gee Aeeran: ADS G ALT eG
Beeman
(a tee triplet ‘frames’
Cron
pa age sate
rs Gigag | C ERS eT OO | ee | |
' |
ee
| |
(+C) :
Base removed (A)
| '
| I
GEA Tae iG aA Se i Gm Acme Gust AGMAETE Gy bol |
I | ea | | | [ | ! a? pote wae fe
|
| |
ey 7 | |
Base added (C) and removed (A)
tiesGtie GG)
teen ies
| ! | ' ' ‘
Cree TAM apis Ok TG SURG OMe G GRA

|
|
Te ole
|
, | |
(+C) Goa | a
triplet code in phase again
Fig 22.20 Diagrammatic explanation of the effect of adding and deleting bases to the triplet iti
produces a frame shift which makes the original message GAT, GAT, . Z read as TGA, Tn eaerie Secor
ern hee A e
produces a frame shift changing the original message from GAT, GAT, ... to ATG, ATG, ... The
addition of the base Cat th
point indicated and the deletion of the base A at the point indicated restores the original message GAT, GAT. (After
F. H.C. 4
Crick (1962) The genetic code |, Scientific American Offprint No. 123, Wm. Saunders & Co.) ee che rth
818
Fig 22.21 Base sequences indicating non-overlapping and
repeated many times. This was called polyuridylic acid
overlapping codes
(poly-U) and acted as a code. Aseries of 20 test-tubes was
| prepared, each containing cell-free extracts of E. coli
Ce See eaGe Tac! fe
aa l ee original triplet including ribosomes, tRNA, ATP, enzymes and a different
( )|\ ) sequence (non- radioactive labelled amino acid. Poly-U was added to each
| overlapping)
test-tube and left to allow in vitro synthesis of polypeptides
| to occur. Analysis of the contents of the test-tubes showed
oC [haa
=e G : that a polypeptide had been formed only in the test-tube
first overlapping
sequence containing the amino acid phenylalanine. Thus the genetic
code had been partly solved. Nirenberg had shown that the
base triplet of the mRNA, or codon, UUU, determines the
WD G
position of phenylalanine in a polypeptide chain. Niren-
‘€ 5
al second overlapping berg and his co-workers then set about preparing synthetic
\ ) sequence
polynucleotide molecules of all 64 possible codons and by
1964 had translated the codes for all 20 amino acids (table
22:4):
As can be seen from table 22.4 some amino acids have
several specified codons. This type of code where the
number of amino acids is less than the number of codons is
termed degenerate. Analysis of the code also shows that for
many amino acids only the first letters appear to be
significant. Three of the codons shown in table 22.4 act as
22.5.4 Breaking the code ‘full stops’ in determining the end of the code message.
These presumably mark the end-point of a functional unit
In order to understand the experimental of the DNA, acistron. In all of these experiments involved
procedures used to determine which triplet base sequence in breaking the genetic code, mRNA was used as the source
codes for which amino acid (that is, break the genetic code) of triplet bases. The ‘genetic code’, however, is transmitted
it is necessary to appreciate, in outline, the mechanism by from cell to cell and between generations by the triplet base
which the triplet code is translated into the structure of a sequence of DNA. Since mRNA is formed directly on
protein molecule. the DNA polynucleotide strand by the method of com-
Protein synthesis involves the interaction of two types of plementary base pairing, the inheritable DNA genetic code
nucleic acid, deoxyribonucleic acid, DNA and ribonucleic is the complement of the mRNA code. This code can be
acid, RNA. There are three kinds of RNA: messenger obtained by translating the RNA bases into their comp-
RNA (mRNA), ribosomal RNA (rRNA) and transfer RNA lementary DNA bases according to the rules shown in table
(tRNA). Apart from a number of organelles such as 2a:
mitochondria and chloroplasts, DNA is confined to the
nucleus where its base sequence is copied (transcribed) on
to strands of messenger RNA (mRNA) which leave the
nucleus. These become attached to ribosomes in the
cytoplasm where the base sequence of mRNA is translated
into an amino acid sequence. Specific amino acids become
attached to tRNA molecules which link up with comp-
lementary triplet bases on the mRNA. Adjacent amino One of the remarkable features of the genetic code is that it
acids brought together in this way react together to form a is thought to be universal. All living organisms contain the
polypeptide chain. The process of protein synthesis, same 20 common amino acids and the same five nitro-
therefore, depends upon the presence of DNA, mRNA, genous bases, A, G, T, C and U. Nirenberg demonstrated
ribosomes, tRNA, amino acids, ATP as an energy source that introducing mRNA from species A into a cell-free
and various enzymes and cofactors which catalyse each system from species B produced the same polypeptide as
stage in the process. would normally be produced in species A. For example,
Nirenberg used this information and various research mammalian haemoglobin molecules have been synthesised
techniques which had been developed during the late 1950s in cell-free extracts of E. coli supplied with mammalian
and designed aseries of experiments to break the code. The haemoglobin mRNA.
essence of his experiments involved using a known base Certain codons act as ‘start signals’ for the initiation of
sequence of mRNA as a coded message and analysing the polypeptide chains, such as AUG (methionine), whereas
amino acid sequence of the polypeptide chain produced others, such as UAA, are ‘nonsense’ codons and do not
from it. Nirenberg was able to synthesise a mRNA code for amino acids but act as ‘stop signals’ for the
molecule that consisted of the same triplet (UUU) termination of polypeptide chains.
819
acids for which they code
Table 22.4. The base sequences of the triplet code and the amino and T
DNA. The DNA genetic code would have complementary bases
NB These are codons, i.e. base sequences of mRNA and not
would replace U.
o
nA
3
Oo
»
na
eo base
Third
fy
*c.t., chain termination codon, equivalent to a full stop in the message.
Table 22.5. The RNA bases which are complementary to those (3) It is degenerate: a given amino acid may be coded for by
of DNA more than one codon.
(4) It is non-overlapping: for example, an mRNA sequence
DNA bases Complementary RNA bases
beginning AUGAGCGCA is not read AUG/UGA/
A (adenine) U (uracil) GAG ... (an overlap of two bases) or AUG/GAG/
G (guanine) C (cytosine) GCG . . . (an overlap of one base). (However, recent
T (thymine) A (adenine) studies have shown overlapping of certain genes in the
C (cytosine) G (guanine)
bacteriophage ®X174. This seems likely to be excep-
tional and may be an economy measure since it has very
Advances in molecular biology have reached the point few genes.)
now where it is becoming possible to determine the base
sequences for whole genes, and the genetic code for an
entire organism, the phage ®X174, has been determined. 22.6 Protein synthesis
This represented a major landmark and whole genes can
now be synthesised artificially, a practice which is of use in From the information given so far in this
genetic engineering (section 2.3.6). chapter it may be seen that the only molecules capable of
The main features of the genetic code are summarised being synthesised directly from the hereditary material of
below. the cell are proteins. These may have a structural role, such
(1) A triplet of bases in the polynucleotide chain of DNA is as keratin and collagen, or a functional role, as in insulin,
the code for the incorporation of one amino acid into a fibrinogen and most importantly enzymes, which are
polypeptide chain. responsible for controlling cell metabolism. It is the
(2) It is universal: the same triplets code for the same particular range of enzymes in the cell which determines
amino acids in all organisms. what type of cell it becomes. The ‘instructions’ for the
820
manufacture of these enzymes and all other proteins are
to higher plants and animals. It is found in the cytoplasm
located in the DNA, which is generally confined to the where it is associated with protein molecules which
nucleus; but, as was shown in the early 1950s, the actual
together form the cell organelles known as ribosomes (see
synthesis occurs in the cytoplasm and involves ribosomes. section 7.2.6).
This led to the realisation that a mechanism had to exist Ribosomes are the site of protein synthesis. Here the
for carrying the genetic information from nucleus to mRNA ‘code’ is translated into a sequence of amino acids
cytoplasm. In 1961 two French biochemists, Jacob and in a growing polypeptide chain. Ribosomes are often found
Monod, postulated, on theoretical grounds, the existence in clusters linked together by a strand of mRNA. This
of a specific form of RNA functioning as an intermediate complex is known as a polyribosome or polysome and
molecule in the synthesis of protein. This compound was enables several molecules of the same polypeptide to be
identified later as MRNA. produced simultaneously.
22.6.1 The role of RNA 22.6.4 Transfer RNA

RNA exists as a single-stranded molecule in all The existence of transfer RNA (tRNA) (or
living cells. It differs from DNA in possessing the pentose soluble RNA (sRNA) as it is sometimes referred to) was
Sugar ribose instead of deoxyribose and the pyrimidine postulated by Crick and demonstrated by Hoagland in
uracil instead of thymine. Analysis of the RNA content of 1955. Each amino acid has its own tRNA molecule which
cells has shown the existence of three types of RNA which transfers amino acids present in the cytoplasm to the
are all involved in the synthesis of protein molecules. These ribosome. Consequently it acts as an intermediate mole-
are messenger RNA (mRNA), transfer RNA (tRNA) and cule between the triplet code of mRNA and the amino acid
ribosomal RNA (rRNA). All three types are synthesised sequence of the polypeptide chain. It constitutes about
directly on DNA, which is said to act as a template for 15% of the total RNA of the cell and, having on average 80
RNA production, and the amount of RNA in each cell is nucleotides per molecule, it is the smallest of all the RNAs.
directly related to the amount of protein synthesis. There are more than 20 different tRNA molecules (60 have
so far been identified) carrying specific amino acids. All
22.6.2 Messenger RNA tRNA molecules have the same basic structure as shown in
fig 22.22.
Analyses of cells have shown that 3-5% of the The 5’-end of the tRNA always ends in the base guanine
total RNA of the cell is mRNA. This is a single-stranded whilst the 3’-end always ends in the base sequence of CCA.
molecule formed on a single strand of DNA by a process The nucleotide sequence of the rest of the molecule is
known as transcription. In the formation of mRNA only variable and may include some ‘unusual bases’ such as
one strand of the DNA molecule is copied. As yet the inosine (I) and pseudouracil (w). The triplet base sequence
mechanism determining which strand is to be copied has at the anticodon (fig 22.22) is directly related to the amino
not been demonstrated. It may involve the activity of a acid carried by that tRNA molecule. Each amino acid is
promoter gene and operator gene (section 22.7.1). RNA attached to its specific tRNA by its own form of the enzyme
nucleotides are attracted to the DNA strand according to amino-acyl-tRNA synthetase. This produces an amino
the rules of base pairing and link up to form an mRNA acid-tRNA complex with sufficient energy in the bond
polynucleotide strand under the influence of the enzyme between the terminal A nucleotide (of CCA) and the
RNA-polymerase. The base sequence of mRNA is a amino acid to later form a peptide bond with the carboxyl
complementary copy of the template DNA strand and group of the adjacent amino acid. In this way a polypeptide
varies in length according to the length of the polypeptide
chain for which it codes. The smallest mRNA molecule is A
approximately 300 nucleotide units long. Most mRNA C
GC
exists within the cell for a short time. In the case of bacteria
this may be a matter of minutes whereas mammalian
reticulocytes, which lose their nuclei as they become red
blood cells, may continue to produce haemoglobin for
several days.
hydrogen bonding
22.6.3 Ribosomal RNA between complementary
bases
Ribosomal RNA was the first RNA to be
identified, and it makes up approximately 80% of the total —_—’
RNA of the cell. It is synthesised by genes present on the

anticodon
DNA of several chromosomes found within a region of the Fig 22.22 A proposed model for the structure of transfer
nucleolus known as the nucleolar organiser. The base RNA (tRNA). The whole molecule is composed of 80
sequence of rRNA is similar in all organisms from bacteria nucleotides but only 20 show complementary base pairing
821
DNA double helix
chain is synthesised. Experiments using ribosomes from rat
liver cells in a cell-free extract of E. coli have shown that non-transcribing transcribing strand
E. coli proteins can be manufactured despite the presence strand
of ‘foreign’ ribosomes. This demonstrates the universal direction of

nature of coding mechanisms involving mRNA, tRNA and transcription
rRNA in the production of proteins.
22.6.5 The mechanism of protein synthesis

Information from various sources and invol-
RNA
ving a variety of experimental techniques carried out on a polymerase G ©
range of organisms from viruses to mammals has shown _L pp XQ
that protein synthesis is a two-stage process which may be = a
summarised by fig 22.23. @ a
ell membrane
single strand of jp} P-P
CYTOPLASM messenger RNA free nucleoside
transcribed on triphosphates
NUCLEUS tRNA/amino acids the DNA
molecule
DNA =e mRNA—®— ribosome
transcription transcribing and
translation non-transcribing
DNA strands rejoin
nuclear envelope polypeptide
Fig 22.23 Summary diagram of the main steps involved in

protein synthesis
Fig 22.24 Diagram showing the mechanism of transcription.
In the presence of RNA polymerase the DNA double helix
22.6.6 Transcription unwinds by breakage of the hydrogen bonds between
complementary bases, and a polynucleotide strand of mRNA
Transcription is the mechanism by which the is formed from free RNA nucleoside triphosphates which link
up opposite complementary DNA bases on the transcribing
base sequence of a cistron of a DNA strand is converted
strand of template DNA. (After E. J. Ambrose & D. M. Easty
into the complementary base sequence of mRNA. The (1977) Cell biology, 2nd ed., Nelson.)
histone coat protecting the DNA double helix in the region
of the cistron is thought to be stripped away, exposing the
polynucleotide sequences of the DNA molecule. The 22.6.7 Translation
double helix unwinds by breakage of the relatively weak
hydrogen bonds between the bases of complementary Translation is the mechanism by which the
strands exposing single strands of DNA. By some mechan- triplet base sequences of mRNA molecules are converted
ism, as yet not understood, one of these strands is selected into a specific sequence of amino acids in a polypeptide
as a template for the formation of a complementary single chain. This occurs on ribosomes. Several ribosomes may
strand of mRNA. This molecule is formed by the linking become attached to a molecule of mRNA like beads on a
of free ribonucleotides under the influence of RNA string and the whole structure is known as a polysome.
polymerase and according to the rules of base pairing These structures, which can be seen under the electron
between DNA and RNA (table 22.5 and fig 22.24). microscope (fig 22.25) have a common strand with a
The exact nature of the copying of DNA bases into RNA diameter of 1.5 nm. This is the diameter of a single strand
bases has been demonstrated using synthetic DNA com- of mRNA. The advantage of such a complex is that it
posed solely of thymine nucleotides (TTT). When intro- allows several polypeptides to be synthesised simul-
duced into acell-free system containing RNA polymerase taneously (section 22.6.3). Each ribosome is composed of a
and all four nucleotides (A, U, C and G) the messenger small and a large subunit, resembling a ‘cottage loaf’ (fig
RNA formed was composed entirely of adenine nu- 7.18). Messenger RNA is thought to form a reversible
cleotides. attachment to the surface of the small subunit in the
When the mRNA molecules have been synthesised they presence of magnesium ions (Mg’*). Having become
leave the nucleus via the nuclear pores and carry the attached to the ribosome it is thought that two mRNA
genetic code to the ribosomes. When sufficient numbers of codons are exposed to the larger ribosome subunit. The
mRNA molecules have been formed from the cistron the first codon binds the tRNA molecule having the comp-
RNA polymerase molecule leaves the DNA, the two lementary anticodon and which is carrying the first amino
strands ‘zip’ up re-forming the double helix and the acid (usually methionine) of the polypeptide being synthe-
protective protein coat is added again. sised. The second codon then attracts a tRNA-—amino acid
822
tRNA! tRNA?
anticodon large subunit of

codon (UAC ribosome
3’ end
mRNA
small subunit of ribosome
(b)
. od
peptide bond (—C-—N-) now
inactive chromosome tRNA! forms between the two amino
segment acids
tRNA?
, RNA polymerase second tRNA/amino acid
active chromosome
molecule now binds to mRNA
segment polyribosome
ribosome
q direction of protein synthesis (6)

messenger RNA
Fig 22.25 Transcription and formation of a polysome in

bacteria. (a) Electron micrograph of a chromosome segment tRNA! released
aon
tRNA?
to collect another
showing stages in the development of mRNA and the methionin
attachment of ribosomes. (b) Diagrammatic representation of molecule
the structure shown in the electron micrograph in (a) \ a third tRNA/amino acid
AUG approaches the mRNA
complex showing the complementary anticodon (figs
ee et jel al —!
22.26a and b). The function of the ribosome is to hold in complex has moved forward one
frame relative to the ribosome
position the mRNA, tRNA and the associated enzymes (c)
controlling the process until a peptide bond forms between
the adjacent amino acids. Fig 22.26 (a) and (b) Consecutive stages in the attachment
of tRNA/amino acid complexes by their anticodons to the
Once the new amino acid has been added to the growing
codons on MRNA and the formation of a peptide bond
polypeptide chain the ribosome moves along the mRNA to between adjacent amino acids. (c) The relative movements of
enclose a new codon. The tRNA molecule which was mRNA and ribosome exposing a new triplet (frame) for the
previously attached to the polypeptide chain now leaves attachment of the tRNA/amino acid complex. The initial tRNA
the ribosome and passes back to the cytoplasm to be molecule is now released from the ribosome and cycles back
into the cytoplasm to be reactivated by enzymes to form a
reconverted into a new tRNA-amino acid complex (fig new tRNA/amino acid complex
22.26c).
This sequence of the ribosome steadily ‘reading’ and
‘translating’ the mRNA code continues until it comes to a Evidence that it is the complementary base pairing
codon signalling ‘stop’. These terminating codons are between the mRNA codon and the tRNA anticodon which
UAA, UAG and UGA. At this point the polypeptide determines the incorporation of an amino acid into the
chain, now with its primary structure as determined by the polypeptide chain, and not the amino acid, was demon-
DNA cistron, leaves the ribosome and translation is strated by the following experiment. The tRNA-cysteine
complete. The main steps involved in translation may be complex normally pairs up, via its anticodon ACA, with
summarised under the following headings: the mRNA codon UGU. Exposure of this tRNA-cysteine
(1) binding of mRNA to ribosome, complex to a catalyst, Raney nickel, converted the cysteine
(2) amino acid activation and attachment to tRNA, to the amino acid alanine. When the new tRNA-alanine
(3) polypeptide chain initiation, complex (carrying the tRNA-cysteine anticodon) was
(4) chain elongation, placed in a cell-free system containing poly- UGU-mRNA
(5) chain termination, the polypeptide chain formed contained only alanine. This
(6) fate of MRNA, experiment demonstrated the importance of the role of the
and the process is summarised in fig 22.27. mRNA-codon-tRNA-anticodon mechanism in translating
As the polypeptide chains leave the ribosome they may the genetic code.
immediately assume either secondary, tertiary or quatern- The whole sequence of protein synthesis occurs as a
ary structures (section 5.5.3). continuous process and is summarised in fig 22.28.
823
EL
large subunit of ribosome
growing polypeptide chain amino acid

8
as
> ye
oN
7
tRNA molecule passes back into cytoplasm
tRNA molecule
6 ate
\
Cae Game
\ ——
anticodon
l
A U Aj _________» ribosome reading in this direction
/
GA. YA CeCe ey
\ oi
U A AVAree UU GGG aU A UG ee a
5’ end 3’ end
Sa ae Ree eee) a, SS)
codon 4 codon 5 codon 6 codon 7 codon 8
single stranded mRNA i ;
\ |
\ J
s SS Position of ribosome
Sy a
> we
SS ~ oe area + at
small subunit of ribosome

=
SS _—— — —_— eae
its
Fig 22.27 Diagrammatic representation of translation. The anticodon of each specific tRNA/amino acid complex pairs with
complementary bases of the mRNA codon in the ribosome. In the example above a peptide bond would form between leucine
and glycine and in this way an additional amino acid would be added to the growing polypeptide chain
-_—
__~
amino-acyl-tRNA synthetase a
—— << DNA double helix
(.
| ——_——
a Se ie Red
RNA polymerase
nuclear envelope pee ay acid

P tRNA
ribosomes ae
()—~_ nuclear pore
(rRNA + protein) — anticodon
mRNA transcription || DNA template

2A KGC UGAGC Y mRNA (together with
PQOAMDFSFACIO
HANAFANFOAOD ribosomes forms a
wh fo a i ee SS
mRNA a) polysome)
codon translation
polypeptide chain released

from ribosome
Fig 22.28 Simplified summary diagram of the major structures and processes involved in protein synthesis in the cell
824
22.7 Genetic control instead of glucose it will not grow immediately but after a
short delay begins to show the same growth rate as seen on
In this chapter the mechanisms whereby cells a glucose medium. Investigations revealed that growth on
transfer genetic material from generation to generation lactose medium required the presence of two substances
have been described. The structure and methods of not normally synthesised, called $-galactosidase, which
functioning of the genetic material are now known in hydrolyses lactose to glucose and galactose, and lactose
considerable detail, but there are still many areas of permease, which enables the cell to take up lactose. This is
genetics where there are many questions and few answers. an example of where a change in environmental conditions,
Genetic research has come a long way in the last 30 years lactose instead of glucose, has induced the synthesis of a
and produced many answers to fundamental questions. particular enzyme. Other experiments involving E. coli
The major breakthroughs were undoubtedly the discovery showed that high concentrations of the amino acid
of the structure of DNA and the breaking of the genetic tryptophan in the culture medium suppressed the produc-
code. Both of these provided inspiration and incentive to tion of the enzyme tryptophan synthetase used to synthe-
other scientists to delve deeper into understanding the sise tryptophan. B-galactosidase synthesis is an example of
apparent mysteries of molecular genetics. Most of these enzyme induction, whereas the suppression of tryptophan
unsolved problems which face molecular geneticists are synthetase is an example of enzyme repression. On the
concerned with the mechanisms by which gene activity is basis of these observations and experiments, Jacob and
controlled in the processes of metabolism, development Monod proposed a mechanism to account for induction
and differentiation. and repression, the mechanism by which genes are
Classical genetics has demonstrated that all somatic cells ‘switched on and off’.
of an organism carry the same genetic complement, that is
they contain the same number of chromosomes carrying 22.7.1 The Jacob—Monod hypothesis of
the same alleles. Despite this, cells in a multicellular genetic control
organism show wide variation in structure and function. The genetic blueprint determining the amino
Even within a single cell the rate at which certain protein acid sequence of the proteins described above is located on
molecules are synthesised varies according to circum- structural genes, those for B-galactosidase and lactose
stances and demand. Evidence for the mechanism by which permease being closely linked on the same chromosome.
genes are regulated within the cell was first obtained from The activity of these genes is controlled by another gene
studies into the control of enzyme synthesis in E. coli. called a regulator gene which is thought to prevent the
In 1961 Jacob and Monod carried out a series of structural genes from becoming active. This may be
experiments to investigate the nature of induction of situated some distance from the structural genes. Evidence
enzyme synthesis in E. coli. Of the 800 enzymes thought to for the existence of a regulator gene comes from studies of
be synthesised by E. colisome are synthesised continuously mutant E. coli which lack this gene and as a consequence
and are called constitutive enzymes; others are synthesised produce ($-galactosidase continuously. The regulator gene
only in the presence of an inducer compound, which may carries the genetic code which results in the production of a
not be the substrate, and are called inducible enzymes. One repressor molecule that prevents the structural genes from
of the latter enzymes is called B-galactosidase. being active. The repressor molecule does not directly
E. coli will grow rapidly on a culture medium containing affect the structural genes but is thought to influence a gene
glucose. When transferred to a medium containing lactose immediately adjacent to the structural genes known as the
OPERON
structural genes
DNA regulator gene operator gene a b

|
| c
double helix E = ]
|
leseee
Qa) |transcription (4) transcription
VSI
I FF SS mRNA OOS IS AAO DES ORIN
@ translation G) Gs) translation
may or may not bind
with the operator gene
repressor molecule (6) polypeptide synthesis
Fig 22.29 The basic structures and processes involved in the control of protein synthesis according to the hypothesis
produced by Jacob and Monod. The numbers indicate the sequence of events
operator gene. The operator and structural genes are The operator gene becomes active and ‘switches on’ the
collectively known as the operon (fig 22.29). structural genes.
The repressor molecule is thought to be a particular type In the case of E. coli grown on glucose medium, the
of protein known as an allosteric protein which can either regulator gene produces a repressor substance which
bind with the operator gene and suppress its activity combines with the operator gene and switches it ‘off’. The
(‘switch it off’) or not bind and permit the operator gene to structural genes are not activated and no 6-galactosidase
become active (‘switch it on’). When the operator gene is and lactose permease are produced. When transferred to a
‘switched on’ the structural genes carry out transcription lactose medium the lactose is thought to act as an inducer of
and mRNA is formed which the ribosomes and tRNA protein synthesis by combining with the repressor molecule
translate into polypeptides. When the operator gene is and preventing it combining with the operator gene. The
‘switched off’ no mRNA is formed and no polypeptides structural genes become active, mRNA is produced and
are formed (fig 22.29). proteins are synthesised. Lactose is thus an inducer of its
The mechanism controlling whether or not the allosteric own breakdown (fig 22.30).
protein binds to the operator gene is simple, yet sensitive to
varying intracellular conditions. It is thought that the 22.7.3 Enzyme repression
repressor molecule has at least two active sites to which If a co-repressor molecule binds with its active
either an inducer molecule or a co-repressor molecule may site on the repressor molecule it reinforces the normal
become attached, depending upon their relative concentra- binding response of the repressor molecule with the
tions at any given time, as described in section 22.7.4. operator gene. This inactivates the operator gene which, in
effect, prevents the structural genes from being ‘switched
22.7.2 Enzyme induction on’.
The binding of an inducer molecule to its E. coli synthesises the amino acid tryptophan in the
active site on the repressor molecule alters the tertiary presence of the enzyme tryptophan synthetase. When the
structure of the repressor (allosteric effect) (section 6.6) so cell contains an excess of tryptophan some of it acts as a
that it cannot bind with the operator gene and repress it. co-repressor of enzyme synthesis by combining with the
regulator gene operator gene structural gene
©.| (4) transcription
repressor
molecule inducer-repressor
lactose acts as unable to bind to translation
e O-O Cs neha
an inducer operator (5)
f-galactosidase glucose +
lactose (enzyme) galactose
Fig 22.30 (above) The method of induction of B-galactosidase synthesis according to the Jacob—Monod hypothesis. The
numbers indicate the sequence of events
Fig 22.31 (below) The mechanism of repression of tryptophan synthetase synthesis according to the Jacob—Monod
hypothesis. The numbers indicate the sequence of events. Solid lines indicate actual processes. The dotted lines represent
repressed stages
regulator gene operator gene structural gene
eS Creal
binds to operator gene
transcription
functional <<
—---
rear) P ISS
repressor
inactive
repressor molecule
NS
NSIDQIIO
molecule |
(3) translation
@) tryptophan acts as
a cO-repressor
Y
tryptophan synthetase
©
precursor (6) p tryptophan
826
structural genes
regulator gene Operator gene
repressor molecule
P P
AHS,
x
ee |
BAA <
|
1 :
{ar \
enzyme enzyme enzyme enzyme
——$——_> —_.—_> ———_> —_—___
a b © d
substrate —— end-product
negative feedback
Fig 22.32 Mechanisms of control of the metabolic pathways A-E. The solid lines represent the mechanism operating during
negative feedback. The dotted lines represent the mechanisms operating during repression. X represents blocks in the enzyme
synthesis
repressor molecule. Co-repressor and repressor molecules ‘operational’ or not. Secondly, the base sequence of the
combine with the operator gene and inhibit its activity. The promoter gene determines which strand of the DNA
structural genes are ‘switched off’, no mRNA is produced double helix attracts the RNA polymerase. In this way the
and no further tryptophan synthetase is synthesised. This is promoter gene determines which strand of the DNA
an example of feedback inhibition acting at the gene level double helix acts as the template for mRNA transcription.
(fig 22.31).
22.7.4 Control of metabolic pathways

22.8 The genetic control of development
This dual mechanism enables the cytoplasm The life cycle of the majority of multicellular
and nucleus to interact in a delicate control of cell animals and plants begins witha single cell, the zygote. This
metabolism. In the case of a simple metabolic pathway as undergoes repeated mitotic and cytoplasmic divisions to
shown in fig 22.32 the initial substrate and final product can give rise to a complex and highly differentiated organism.
act as inducer and co-repressor respectively. This mechan- This process is known as growth and development and
ism enables the cell to produce the amount of enzyme includes the process of differentiation. Differentiation, that
required at any given time to maintain the correct level of is the process whereby cells assume particular structures
product. This method of metabolic control is highly which enable them to carry out a restricted number of
economical. Negative feedback involving the inactivation particular functions more efficiently, is one of the remark-
of the initial enzyme (a) by combination with the able events of development. Why should the cells of an
end-product (£) would rapidly halt the pathway but would organism produced by repeated cell divisions and contain-
not prevent the continued synthesis of the other enzymes ing identical genetic material show the range of diversity
(b, cand d). In the system proposed by Jacob and Monod, which typifies higher multicellular organisms? The answer
the end-product (£), by combining with the repressor is far from clear but must involve the induction and
molecule to increase its repressive effect on the operator repression of genes, perhaps by mechanisms similar to
gene, would prevent the synthesis of all enzymes (a, b, c those described in the previous section. Evidence has
and d) and halt the pathway. suggested that there are three factors which act together in
various ways to bring about differentiation. They are the
22.7.5 Modification to the ‘operon’ nucleus, the cytoplasm and the environment.
hypothesis
22.8.1 The role of the nucleus
Since 1961 when Jacob and Monod suggested
a mechanism by which genes are ‘switched on and off’, The importance of the nucleus as the storage
further evidence has accumulated which has helped to site of the genetic material and its primary role in
clarify aspects of the mechanism. Genetic evidence has determining phenotypic characteristics has been appreci-
suggested the existence of a promoter gene situated ated for a long time. The German biologist Hammerling
adjacent to the operator gene which acts between it and the was one of the earliest workers to demonstrate the primary
regulator gene. It is thought to have two functions. First the role of the nucleus. He chose as his research organism the
promoter gene is the site to which RNA polymerase binds unusually large acellular marine alga Acetabularia. There
before moving along the DNA to begin the transcription of are two closely related species, A. mediterranea and A.
mRNA on the structural genes. This movement will, of crenulata, which differ only in the shape of their ‘head’
course, depend upon whether the operator gene is region (fig 22.33).
827
(a) In a series of experiments, including some that involved
\IZ separating the ‘head’ region from the ‘basal’ region (which
SF
‘head’
contained the nucleus), he was able to demonstrate the
<> necessity of the nucleus for normal development. Further
experiments involving the reciprocal grafting of the
nucleus-containing ‘basal’ region of one species to the
enucleate stalk region of the other species always produced
hybrids that developed the ‘head’ which was characteristic
‘stem’ of the nucleate portion of the graft.
However, in considering this model of nuclear control,
due regard must be taken of the primitive nature of the
organism used. Later experiments performed by two
American scientists, Briggs and King, in 1952, made use of
the technique of grafting but in this case they used cells
basal region
from the frog Rana pipiens. The nuclei were removed from
unfertilised egg cells and replaced with nuclei from late
blastula cells which showed signs of differentiation. In most
nucleus
cases the recipient cells developed and grew into normal
adult frogs.
A. mediterranea A. crenulata
“What did the results of the above
experimen Jemonstrate?
Fig 22.33 The marine alga Acetabularia used by
Hammerling to demonstrate the role of the nucleus. (a) Two
species of Acetabularia. (b) Transplant and excision
experiments carried out by Hammerling
22.8.2 The role of the cytoplasm
(b) A. mediterranea A. crenulata
Further evidence for the role of the cytoplasm
is provided by embryological studies. From an early stage
in the embryological development of many organisms from
algae to mammals the egg cytoplasm is not homogeneous.
The cytoplasm appears to be stratified, with less dense
material at the upper pole and dense granular material at
the lower pole. In those species, where the early cleavage
planes are vertical, such as amphibia, all resulting cells, if
separated, give rise to normal offspring. In other species
where early cleavage planes are horizontal, such as
molluscs, the cells, if separated, do not undergo normal
‘head’
removed
‘head’ development. In the former case all cells contain an equal
removed
along along X-X distribution of the different layers of cytoplasm, whereas in
W-W the latter there is an unequal distribution of cytoplasm.
stem removed
between W-W Eggs where the cytoplasm is differentiated in this way in
and Y-Y
order to give rise to certain regions of the embryo are
known as mosaic eggs. In all cases the nuclei of the cells
basal region
removed below
contain the same genetic complement. Differential de-
Z-Z velopments would therefore appear to result from some
form of cytoplasmic influence on the genes.
In a series of now classic embryological experiments,
Spemann and Mangold demonstrated in 1924 that dif-
ferentiation is largely controlled by the cytoplasmic
influence which one cell type has over another. In one of
their experiments, they removed tissue from the dorsal lip
of the blastopore of an amphibian gastrula (fig 22.34) and
new ‘head’ develops implanted it into a ventral region of another gastrula (the
as for A. crenulata host). The cells of the dorsal lip normally develop into the
notochord, mesodermal somites (myotomes) and neural
tube. The host gastrula in this experiment developed a
secondary notochord, extra myotomes and a neural tube in
828
blastopore neural plate other tissues to act as secondary and tertiary organisers and
notochord
so on until all the organs and organ systems of the embryo
section of have differentiated and developed in their normal posi-
Qs. b } blastopore
tions.
22.8.3 The nature of the organiser

Se - endoderm
(a) tissue from (b)
Experiments have been carried out involving
(b) ectoderm the removal of a tissue known to act as an organiser and
transplanted here
placing it on a piece of agar for several hours. Implantation
neural tube of the agar into regions known to be competent has induced
notochord them to differentiate as directed by the organiser tissue.
This technique demonstrates that the organiser is a
chemical substance which has diffused into the agar.
Attempts to isolate the substance have so far produced no
mesodermal somites clear-cut result. Steroids, proteins and nucleic acids have
notochord
all been suggested as likely organiser substances. Paul and
Gilmour have observed that DNA molecules in different
neural tube tissues are differentially ‘coated’ with histone and non-
(c) histone proteins. They suggest that histones ‘cover’ those
regions of the DNA which are non-functional, that is their
genes are irreversibly repressed, whilst non-histone pro-
teins ‘cover’ those genes which are to be transcribed. This
(d) embryo derived from host gastrula idea fits in well with the situation in mosaic eggs where a
©
distribution of histones throughout the cytoplasm would
lead to the production of cells with different regions of
DNA ‘blocked off’. In this way various genes would be
©
effectively ‘switched off’ in different cells and this would
embryo derived from transplanted
tissues from dorsal lip of blastopore provide a possible mechanism of differentiation.
Fig 22.34 Technique and results of Spoemann and Mangold’s 22.8.4 The role of the environment
experiments on embryonic induction. (a) Surface view of the
developing gastrula showing the blastopore. (b) LS through It is largely as a result of experiments such as
gastrula of an amphibian showing the region of the dorsal lip those of Jacob and Monod that the extent of environmental
of the blastopore which is excised and transplanted into
gastrula (a). (c) TS through the developing embryo showing influence can be seen on development. For example,
the development of two identical groups of embryological lactose is an environmental factor that has a direct
tissues. (d) ‘Siamese’ embryos produced by transplanting influence on the functioning of genes in E. coli. Light,
tissue from the dorsal lip of the blastopore temperature, water, nutrients and gas supplies can all
influence the extent of development and growth in plants
and animals as described in chapters 9, 10, 15, 16 and 21.
the region of the transplant which gave rise to a second The effect of the environment on differentiation is
tadpole as shown in fig 22.34. probably normally through an intermediate influence on
On the basis of these observations Spemann and the cytoplasm which in turn has a direct effect on the genes.
Mangold advanced a hypothesis of differentiation known
as embryonic induction. According to this hypothesis 22.8.5 The role of genes in development
certain cells act as organisers of other cells described as
competent (or determined). Organisers are capable of The relationship between specific regions of
inducing competent cells to develop into cell types having a the DNA molecule and morphological development has
structure and function different from those which would been studied extensively in organisms possessing giant
appear in the absence of organisers. These misplaced chromosomes. These are found, for example in the salivary
structures are described as atopic (a, without; topos, glands of many dipteran larvae, including Drosophila. The
place). reasons for the size of these polytene chromosomes are
In the embryological development of organisms acertain given in section 23.5.1. They are relatively easy to see
region known as the primary organiser determines the under the light microscope and show distinct banding
entire further course of development. In the case of patterns when stained with Feulgen stain. During meta-
amphibia this is the dorsal lip of the blastopore, whereas in morphosis the Drosophila larvae pass through several
birds it is a region known as the primitive streak. ‘These stages, or instars, each separated by a period of intense
primary organisers establish the embryonic axis and induce cellular activity called ecdysis followed by moulting of the
829
Fig 22.35 The relationship between puffing
and mRNA synthesis is clearly
demonstrated by these autoradiograms of
chromosome IV of the midge Chironomus
tentans. The black dots indicate the position
of radioactive uridine taken up during mRNA
synthesis. (a) shows that RNA synthesis is
closely related to the region of puffing. (b)
Very little puffing and uridine uptake has
4 occurred following the addition of smail
att: *% amounts of actinomycin D which inhibits
é mRNA synthesis. (From W. Beerman & U.
Clever (1964) Chromosome puffs, Scientific
American Offprint No. 180. Wm. Saunders &
* Co)
old cuticle. The final two ecdyses are the most dramatic and characteristic, puffing at this locus is only seen in the
produce the pupal stage and the imago (adult) stage. These chromosome carrying the dominant allele.
are stages of intense metabolic activity and differentiation. Further evidence for the link between chromosome puffs
Metamorphosis is controlled by hormones as described in and mRNA synthesis is provided by the effect of injecting
section 21.4. During metamorphosis ‘bands’ along the the drug actinomycin D into organisms having giant
chromosomes enlarge and form structures known as chromosomes. Actinomycin D inhibits transcription by
chromosome puffs or rings of Balbiani (after the scientist preventing the synthesis of mRNA, and no puffing is seen
who first observed them in 1890). Specific stains which in organisms treated in this way (fig 22.35).
show up RNA, and autoradiographic studies involving Another factor influencing growth and development in
labelled RNA nucleotides have shown these ‘puffs’ to be plants and animals is hormones. In many cases this occurs
regions of RNA synthesis. The size of the ‘puffs’ has also at the level of transcription of mRNA. The exact way in
been shown to be directly related to the rate of RNA which a given hormone affects transcription and protein
synthesis. The puffing effect is thought to be produced by synthesis is extremely variable, but some are thought to
the unwinding of DNA molecules, the separation of exert their influence on receptor sites on the cell mem-
complementary strands and the formation of mRNA brane. Following binding of the hormone on to the
during transcription. receptor site, thought to be the enzyme adenylate cyclase,
There is a definite sequence of puffing during meta- cyclic AMP is released into the cytoplasm and this acts as a
morphosis and this is induced by the moulting hormone second messenger which induces transcription. Further
ecdysone. During the various larval stages and the pupal details of this mechanism are described in section 16.6.1.
stage different regions of the chomosomes show ‘puffing’, This chapter has attempted to describe some of the
suggesting that the puffs correspond to the structural genes processes associated with the continuity of life. Living
postulated in the Jacob—Monod hypothesis. Evidence that systems appear to require both short-term genetic stability
regions of puffing correspond to regions of genetic activity and long-term genetic flexibility. Genetic stability is seen to
was produced by Beerman using two species of midge be achieved by the mechanisms of mitosis, whereas the
belonging to the genus Chironomus. Certain cells in the mechanisms of meiosis introduce an enormous amount of
salivary gland of one species are granulated whereas those genetic variation, enabling organisms to adapt to changing
of the other species are non-granulated. Genetic mapping environment.
based upon crosses between these two species has shown Having established the cytological mechanisms of inheri-
that the alleles determining these characteristics are tance, research has shifted to considerations of the nature
situated near the centromere of one of the chromosomes of the genetic material and the mechanisms involved in its
and the allele for granulated cells is dominant. Examina- control of cellular activities such as development, growth
tion of the chromosomes of these species showed that and differentiation. The next chapter describes the ways in
puffing is only seen in the region where this gene is located which characteristics are inherited, the rules governing
in the species producing granulated cells. Furthermore, in these processes and the way variations are introduced into
the case of midges which are heterozygous for this populations.
830
Chapter Twenty-three
Variation and genetics
Genetics may rightly be claimed to be one of the most the process of hybridisation in plants and, in particular, the
important branches of biology. For thousands of years, different forms in which hybrid progeny appear and the
Man has used the techniques of genetics in the improve- statistical relationships between them. This formed the
ment of domestic animals and crops without having any basis of Mendel’s scientific investigations on inheritance
real knowledge of the mechanisms which underlie these which he began in the summer of 1856.
practices. Various pieces of archaeological evidence dating Mendel’s success was due, in part, to his careful choice of
back 6 000 years suggest that Man understood that certain experimental organism, the garden pea, Pisum sativum.
physical characteristics could be transmitted from one He ascertained that it had the following advantages over
generation to another. By selecting particular organisms other species:
from wild stocks and interbreeding these, Man has been (1) There were several varieties available which had quite
able to produce improved varieties of plants and animals to distinct characteristics.
suit his needs. (2) The plants were easy to cultivate.
It is only since the beginning of this century, though, that (3) The reproductive structures were completely enclosed
scientists have begun to appreciate fully the principles and by the petals so that the plant was normally self-
mechanisms of heredity. Whilst advances in microscopy pollinating. This led to the varieties producing the same
have revealed that the sperm and the ova transmitted the characteristics generation after generation, a phe-
hereditary characteristics from generation to generation, nomenon known as pure breeding.
the problem nevertheless remained of how minute particles (4) Artificial cross-breeding between varieties was possible
of protoplasm could carry the vast number of characteris- and resulting hybrids were completely fertile. From the
tics that make up an individual organism. 34 varieties of garden pea, Mendel selected 22 varieties
The first really scientific advance in the study of which showed clear-cut differences in characteristics
inheritance was made by the Austrian monk Gregor and used these in his breeding experiments. The seven
Mendel who published a paper in 1866 which laid the basic characteristics, or traits, that Mendel was in-
foundations for the present-day science of genetics. He terested in were length of stem, shape of seed, colour of
demonstrated that characteristics do not blend but pass seed, shape and colour of pod, position and colour of
from parents to offspring as discrete units. These units, flower.
which appear in the offspring in pairs, remain discrete and Many scientists before Mendel had performed similar
are passed on to subsequent generations by the male and experiments on plants but none had produced results which
female gametes which each contain a single unit. The had the accuracy and detail of Mendel’s, nor were they able
Danish botanist Johannsen called these units genes in 1909, to explain their results in terms of a mechanism of
and the American geneticist Morgan, in 1912, demon- inheritance. The reasons for Mendel’s success may be
strated that they are carried on the chromosomes. Since the taken as a model of how to carry out a scientific
early 1900s the study of genetics has made great advances in investigation. They may be summarised as follows:
explaining the nature of inheritance at both the level of the (1) Preliminary investigations were carried out to obtain
organism and at the level of the gene. familiarity with the experimental organism.
(2) All experiments were carefully planned so that atten-
tion was focussed on only one variable at any time, thus
23.1 Mendel’s work simplifying the observations to be made.
(3) Meticulous care was taken in carrying out all tech-
Gregor Mendel was born in Moravia in 1822.
niques, thus preventing the introduction of contami-
In 1843 he joined an Augustinian monastery at Brinn in nating variables (see below for details).
Austria (now Brno, in Czechoslovakia) where he took (4) Accurate records were kept of all the experiments and
Holy Orders. From there he went to the University of the results obtained.
Vienna where he spent two years studying natural history (5) Sufficient data were obtained to have statistical
and mathematics before returning to the monastery in significance.
1853. This choice of subjects undoubtedly had a significant As Mendel stated,
influence on his subsequent work on inheritance in pea
‘The value and utility of any experiment are
plants. Whilst in Vienna, Mendel had become interested in
831
determined by the fitness of the material to the The F, plants then had their flowers enclosed in bags (to
purpose for which it is used.’ prevent cross-pollination occurring) and were left to
However, it is worth stating that there was an element of self-pollinate. The seeds collected from these F, plants
luck in Mendel’s choice of experimental organism. The were counted and planted the following spring to produce
characters chosen by Mendel lacked many of the more the second filial generation or F, generation. (An F,
complex genetic features which were later discovered, such generation is always the result of allowing the F, generation
as incomplete dominance (section 23.7.1), characteristics to inbreed or, as in this case, to self-pollinate.) When these
controlled by more than one pair of genes (section 23.7.6) plants flowered, some bore axial flowers and others
and linkage (section 23.3). terminal flowers. In other words, the characteristic ‘ter-
minal flower’, which was absent in the F, generation, had
reappeared in the F, generation. Mendel reasoned that the
23.1.1 Monohypbrid inheritance and the terminal characteristic must have been present in the F,
principle of segregation generation but as it failed to be expressed in this generation
Mendel’s earliest experiments involved select- he termed it recessive. Of the 858 F, plants that Mendel
ing plants of two varieties which had clearly differentiated obtained, 651 had axial flowers and 207 had terminal
characteristics, such as flowers distributed along the main flowers. Mendel carried out a series of similar experiments
stem (axial) or flowers at the tip of the stem (terminal). involving in each case the inheritance of a single pair of
These plants, showing a single pair of contrasted character- contrasting characteristics. Seven pairs of contrasting
istics, were grown for a number of generations. Seeds characteristics were studied and the results of the ex-
collected from axial plants always produced plants with perimental crosses are shown in table 23.1. In all cases the
axial flowers, whilst those from terminal plants always analyses of the results revealed that the ratios of dominant
produced terminal flowers. This demonstrated to Mendel to recessive characteristics in the F, generation were
that he was using pure-breeding plants. With this informa- approximately 3:1.
tion he was in a position to carry out hybridisation The example quoted above is typical of all Mendel’s
experiments (experimental crosses) using these plants. experiments involving the inheritance of a single character-
His experimental technique involved removing the anthers istic (monohybrid inheritance) and may be summarised as
from a number of plants of one variety before self- follows.
fertilisation could have occurred. These he called ‘female’
Observations
plants. Pollen was then transferred, by means of a brush,
Parents axial flowers X< terminal flowers
from the anthers of another plant of the same variety to the
F; all axial flowers
stigmas of the ‘female’ plant. The experimental flowers
F, 651 axial flowers 207 terminal flowers
were then enclosed in a small bag to prevent pollen from
F, ratio 3 : 1
other plants reaching their stigmas. Reciprocal crosses
were carried out by transferring pollen grains from axial On the basis of these, and similar results, Mendel drew the
plants to terminal plants and pollen grains from terminal following conclusions.
plants to axial plants. In all cases the seeds subsequently (1) Since the original parental stocks were pure breeding,
collected from both sets of plants gave rise to plants with the axial variety must have possessed two axial factors
axial flowers. This characteristic, ‘axial flower’, shown by and the terminal variety two terminal factors.
these first generation hybrid plants (subsequently called (2) The F, generation possessed one factor from each
the first filial generation or F, generation by Bateson and parent which were carried by the gametes.
Saunders in 1902) was termed dominant by Mendel. (3) These factors do not blend in the F, generation but
None of the F, plants produced terminal flowers. retain their individuality.
Table 23.1 The results of Mendel’s experiments on the inheritance of seven pairs of contrasted characteristics. (The
observed ratio of dominant to recessive characteristics approximates to the theoretical value of 3:1.)
Characteristic Parental appearance F, appearance Ratio

(dominant) (recessive) (dominant) (recessive)
length of stem tall dwarf 787 Zine 2.84:1

shape of seed round wrinkled 5474 1850 2.96:1
colour of seed yellow green 6022 2001 3.01:1
shape of pod inflated constricted 882 oe 2.95:1
colour of pod green yellow 428 eZ 2.82:1
position of flower axial terminal 651 207 3.14:1
colour of flower red white 705 224 ee
total 14.949 5.010 2.98:1
oso SS——__C—W——aaa00 009050 OOo
832
(4) The axial factor is dominant to the terminal factor Let:
which is recessive. A represent axial flower (dominant)
The separation of the pair of parental factors, so that one a represent terminal flower (recessive)
factor is present in each gamete, became known as

Parental phenotypes pure breeding axial X_ pure breeding terminal
Mendel’s first law, or the principle of segregation. This flower flower
states that Parental genotypes (2n) AA x aa
the characteristics of an organism are determined by Meiosis

internal factors which occur in pairs. Only one of a pair Gametes (n) (A) (A) x (a) (a)
of such factors can be represented in a single gamete.
We now know that these factors determining characteris-
tics, such as flower position, are regions of the chromosome
‘Random fertilisation
F genotypes (2n)
jes cae ala
Aa Aa Aa Aa
known as genes.
The foregoing experimental procedure carried out by
F, phenotypes all heterozygous axial flower (the alleles A and
Mende! in the investigation of the inheritance of a single a remain distinct in spite of the dominance of A)
pair of contrasted characteristics is an example of a
monohybrid cross. This may be represented in terms of
symbols and placed in a modern context of gamete The F, generation were self-pollinated
formation and fertilisation. By convention, the initial letter
of the dominant characteristic is used as the genotypic
symbol and its capital form (e.g. A) represents the
F, phenotypes axial flower x axial flower
dominant allele and the lower case (e.g. a) represents the
recessive allele. All of the terms and symbols described F, genotypes (2n) x Aa
above are used in genetics and are summarised in table Meiosis
2D
Fig 23.1 shows the correct way to describe a monohybrid
Gametes (n)
a 2) aateei ae?
cross or arrive at the solution to a genetics problem
involving the inheritance of a single pair of contrasted
Random fertilisation
F, genotypes (2n) AA Aa Aa
aa aa
characteristics. ee eee,
The ratio of dominant phenotypes to recessive pheno- homozygous heterozygous homozygous
types of 3:1 is called the monohybrid ratio. Mendel’s
conclusions regarding the transfer of a single characteristic F, phenotypes 3 axial flower 1 terminal flower
by each gamete and the genotypic appearance can be Fig 23.1 Full genetic explanation of one of Mendel’s
demonstrated by mathematical probability. The probabil- monohybrid crosses. (2n represents the diploid condition,
ity of a gamete cell from a heterozygous F, parent n represents the haploid condition; see section 22.3.)
containing either the dominant allele A or the recessive
allele a is 50% or 4. If each gamete is represented by 2, the
Table 23.2 Glossary of common genetic terms with examples based on fig 23.1.
Genetic term Explanation Example
ene The basic unit of inheritance for a given characteristic flower position
ae Alternative forms of the same gene responsible for Aora
determining contrasting characteristics
locus Position of an allele on a chromosome
homozygous The diploid condition where both alleles are identical AA or aa
heterozygous The diploid condition where different alleles are present Aa .
phenotype The physical or chemical expression of a characteristic axial, terminal
genotype The genetic expression of a characteristic in terms of alleles AA, Aa, AA
dominant The allele which influences the appearance of the phenotype A
when present in the homozygous or heterozygous condition
The allele which only influences the appearance of the a
recessive
phenotype when present in the homozygous condition
F, generation The generation produced by crossing two parental stocks
F, generation The generation produced by crossing two F, organisms
Se
833
number of possible combinations of F, genotypes is 23.1.2 Backcross or test cross
represented by 4 x 4 = 4. Hence there are four possible F, The genotype of an F, organism, produced by
genotypes. The statistical probability of the A and a the breeding of homozygous dominant and homozygous
containing gametes combining by random fertilisation is recessive parents, is heterozygous but shows the dominant
shown in fig 23.2 As a result of dominance the phenotpyic phenotype. An organism displaying the recessive pheno-
appearance will be 3 dominant phenotypes : 1 recessive type must have a genotype which is homozygous for the
phenotype. The results of Mendel’s breeding experiments recessive allele. In the case of F, organisms showing the
bear out this theoretical ratio as shown in table 23.1.
dominant phenotype the genotype may be either homozy-
gous or heterozygous. It may be of interest to a breeder to
know the genotype and the only way in which it can be
Let the probability ofthe alleles A and a appearing in the
heterozygote (Aa) = 1,
therefore A = //, determined is to carry out a breeding experiment. This
a=, involves the use of a technique known as testcross or
Using these values the probability of each genotype and phenotype
backcross. By crossing an organism having an unknown
appearing in the F, generation can be demonstrated as shown below:
F, genotypes (2n) Aa x Aa genotype with a homozygous recessive organism it is
possible to determine an unknown genotype within one
®QH ~ ©
Meiosis
breeding generation. For example in the fruit fly, Dro-
Gametes (n) sophila, long wing is dominant to vestigial wing. The
genotype of a long wing Drosophila may be homozygous
(In terms of probability)
Os) W ; Cs)
SS
(LL) or heterozygous (LI). In order to establish which is the
Random fertilisation correct genotype, the fly is testcrossed with a double
F, genotypes (2n) Y, AA Y Aat+!/, Aa Yaa recessive (Il) vestigial wing fly. If the testcross offspring are
all long wing the unknown genotype is homozygous
F, phenotypes ¥, dominant: '/, recessive
i.e. 3 dominant: | recessive dominant. A ratio of 1 long wing : 1 vestigial wing indicates
that the unknown is heterozygous (fig 23.3).
Fig 23.2 Explanation of the 3:1 Mendelian monohybrid ratio
in terms of probability
23.2 Why is it not possible to use a
_ homozygous dominant organism (such as TT) in a
backcross experiment to determine the | .
an organism showing the domir
_---23.1_‘If a pure strain of mice wit yn- Illustrate your answer fully using appre
_ coloured fur are allowed to breed with a pure strain of | symbols. —
_ mice with grey-coloured fur they produce offspring
_ having brown-coloured fur. If the F, mice are allowed
_ to interbreed they produce an F, generation with fur 23.1.3 Dihybrid inheritance and the
colour in the proportion of three brown-coloured to principle of independent assortment
one grey. -
(a) Explain fully these results. Having established that it was possible to
(b) What would be the result of mating a brown- predict the outcome of breeding crosses involving a single
coloured heterozygote from the F, generation pair of contrasted characteristics, Mendel turned his
with the original grey-coloured parent? attention to the inheritance of two pairs of contrasted
characteristics. Since two pairs of alleles are found in
Let: L represent long wing (dominant)

l represent vestigial wing (recessive)
homozygous long wing parent heterozygous long wing parent
Testcross phenotypes long wing (homozygous) X vestigial wing long wing (heterozygous) vestigial wing
Testcross genotypes (2n) LL ss Il Ll x ll
Meiosis
Gametes (n) (L) (L) x C) C1) (L) CG) x ay CG)

Offspring genotypes (2n) LI LI LI LI LI LI ll ll

Offspring phenotypes all long wing (heterozygous) long wing (heterozygous) vestigial wing (homozygous)
1 : 1
Fig 23.3 A/ full genetic explanation of how to determine the genotype of an organism showin a dominant
I ch isti i
technique is known as a testcross, and produces offspring phenotypes as Sta m oS anes
834
(a) Let: R represent round seed (dominant) Y represent yellow seed (dominant)
r represent wrinkled seed (recessive) y represent green seed (recessive)
Parental phenotypes round seed and yellow seed (homozygous) x wrinkled seed and green seed (homozygous)
Parental genotypes (2n) RRYY x rryy
Meiosis
Gametes (n) all(RY) x all
(ry)
F, genotypes (2n) all RrYy
F, phenotypes all heterozygous round and yellow seeds
(b) F, phenotypes round and yellow seed x round and yellow seed
F, genotypes (2n) Rr Yy Xx RrYy
Meiosis
Gametes (n) (as shown byfand @). Seed RY Ry Ey ou
RY RY Ry rY ry
RY O RY © RY © RY O
R RY Ry rY ry
Sore as yi
Random fertilisation (as shown by Punnett square) Ry © Ry @| Ry © Ry @
RY Ry rY ry”
rY rY O rY O ry © rY i
Fy RY Ry ry ry
F, genotypes (2n) (listed in each square) a O a @ no O a ®
F, phenotypes 9 round yellow; 3 round green: 3 wrinkled yellow: 1 wrinkled green seeds
O
Fig 23.4 (a) Stages in the formation of F, phenotypes from homozygous parents. This is an example of a dihybrid cross since
two characteristics are being considered (b) Use of Punnett square to show all possible combinations of gametes to form F,
genotypes
the heterozygotes, this condition is known as dihybrid whilst combining in the F, generation, separate and behave
inheritance. independently from one another in subsequent genera-
In one of his experiments Mendel used pea shape and tions. This forms the basis of Mendel’s second law or the
pea cotyledon colour as the characteristics. Using the same principle of independent assortment which states that,
techniques as described in section 23.1.1, he crossed any one of a pair of characteristics may combine with
pure-breeding (homozygous) plants having round and either one of another pair.
yellow peas with pure-breeding plants having wrinkled and The above experiment can be written out in terms of our
green peas. The F, generation seeds were round and present knowledge of genetics as shown in fig 23.4a. As a
yellow. Mendel knew that these characteristics were result of separation (segregation) of alleles (R, r, Y and y)
dominant from earlier monohybrid breeding experiments and their independent assortment (rearrangement or
but it was the nature and number of organisms of the F, recombination) four possible arrangements of alleles can
generation produced from the self-pollination of the F, be found in each of the male and female gametes. In order
plants that now interested him. He collected a total of 556 to demonstrate all the possible combinations of gametes
F, seeds from the F, generation which showed the following that occur during random fertilisation a Punnett square is
characteristics: used. This is a grid named after the Cambridge geneticist
315 round and yellow, R. C. Punnett and its value lies in minimising the errors
101 wrinkled and yellow, which can occur when listing all possible combinations of
108 round and green and gametes. It is advisable when filling in the Punnett square
32 wrinkled and green. to enter all the ‘male’ gametes first in the vertical squares
and then enter all the ‘female’ gametes in the horizontal
The proportions of each phenotype approximated toa ratio squares. Likewise, when determining the F, phenotypes, it
of 9:3:3:1. This is known as the dihybrid ratio. Mendel is advisable to mark off identical phenotypes in some easily
made two deductions from these observations. identifiable way, as shown in fig 23.4b. From figs 23.4a and
(1) Two new combinations of characteristics had appeared b, which are based on Mendel’s first and second laws, it can
in the F, generation: wrinkled and yellow, and round be seen that each F, male and female genotype can give rise
and green. to gametes with the following combination of alleles;
(2) The ratios of each pair of allelomorphic characteristics
R can only be present with Y or y (not r), that is RY or
(phenotypes determined by different alleles) appeared
in the monohybrid ratio of 3:1, that is 423 round to 133 Ry,
r can only be present with Y or y (not R), thatisrY orry.
wrinkléd, and 416 yellow to 140 green.
Thus there is a 1 in 4 chance of any gamete containing
On the basis of these results Mendel was able to state that
any of the four allele combinations shown above.
the two pairs of characteristics (seed shape and colour),
835
From a consideration of monohybrid inheritance, where 7 (5) Each allele is transmitted from generation to genera-
of the F, phenotypes show the dominant allele and 4 the tion as a discrete unchanging unit.
recessive allele, the probability of the four alleles appear- (6) Each organism inherits one allele (for each characteris-
ing in any F, phenotype is as follows: tic) from each parent.
NB The mechanism of dihybrid inheritance, the examples
round (dominant) quoted in this section and the typical dihybrid ratio of
yellow (dominant) 9:3:3:1 only apply to characteristics controlled by genes on
wrinkled (recessive) different chromosomes. Genes situated on the same
green (recessive) mico
ico
mie chromosome may not show this pattern of independent
assortment as described in section 23.3.
Hence the probability of the following combinations of
alleles appearing in the F, phenotypes is as follows:
round and yellow lI 23.2 The chromosomal basis of

round and green = inheritance
wrinkled and yellow =
wrinkled and green lI HalCo
ICO
ale ex
xe ico
bie
Bio
mie
Sle
Sho
bale
par a)
Mendel published his research data and
hypotheses in 1866 in a journal, The Proceedings of the
The results of Mendel’s breeding experiments with two Briinn Natural History Society, which was sent to most of
pairs of contrasted characteristics approximated to the the learned scientific societies throughout the world. In all
theoretical values shown above. cases they failed to appreciate the importance of his
findings, possibly because scientists at the time were unable
23.3 In the guinea pig, (Cavia), there are to relate them to any physical structures in the gametes by
two alleles for hair colour, black and white, and two which the hereditary factors might be transmitted from
alleles for hair length, short and long. In a breeding parent to offspring.
experiment all the F, phenotypes produced from a By 1900, as a result of improvements in the optical
cross between pure-breeding, short black-haired properties of microscopes and advances in cytological
and pure-breeding, long white-haired parents had techniques, the behaviour of chromosomes in gametes and
_ short black hair. Explain (a) which alleles are zygotes had been observed. In 1875 Hertwig noted that
_ dominant, and (6) the expected Bioportoh - during the fertilisation of sea urchin eggs two nuclei, one
phenotypes. from. the sperm and one from the egg, fused together.
Boveri, in 1902, demonstrated the importance of the
oo 23.4 Flower colour in sweet peaplantsis Is
nucleus in controlling the development of characteristics in
determined by two allelomorphic pairs of genes (R,r,
and S,s). If at least one dominant gene from each organisms, and in 1882 Flemming clarified the chromo-
_ allelomorphic pair is present the flowers are purple. somal events involved in mitosis.
All other genotypes are white. In 1900 the significance of Mendel’s work was realised
If two purple plants, each having the genotype almost simultaneously by three scientists, de Vries,
RrSs, are crossed, whatwil be the phenotypic ratio Correns and Tschermak. In fact, it was Correns who
of the offspring? summarised Mendel’s conclusions in the familiar form of
two principles and coined the term ‘factor’, Mendel having
used the term ‘elemente’ to describe the hereditary unit. It
was an American, William Sutton, however, who noticed
23.1.4 Summary of Mendel’s hypotheses
the striking similarities between the behaviour of chromo-
The following summary includes terms taken somes during gamete formation and fertilisation, and the
from our present knowledge of the nature of genetics. transmission of Mendel’s hereditary factors. These have
(1) Each characteristic of an organism is controlled by a been summarised in table 23.3.
pair of alleles. On the basis of the evidence suggested above, Sutton and
(2) If an organism has two unlike alleles for a given Boveri proposed that chromosomes were the carriers of
characteristic, one may be expressed (the dominant Mendel’s factors, the so-called chromosome theory of
allele) to the total exclusion of the other (the recessive heredity. According to this theory, each pair of factors is
allele). carried by a pair of homologous chromosomes, with each
(3) During meiosis each pair of alleles separates (segre- chromosome carrying one of the factors. Since the number
gates) and each gamete receives one of each pair of of characteristics of any organism vastly outnumbers the
alleles (the principle of segregation). chromosomes, as revealed by microscopy, each chromo-
(4) During gamete formation in each sex, either one of a some must carry many factors.
pair of alleles may enter the same gamete cell (combine The term factor as the basic unit of heredity was replaced
randomly) with either one of another pair (the principle by Johannsen, in 1909, with the term gene. Whilst gene is
of independent assortment). used to describe the unit of heredity, it is the alternative
836
Table 23.3 A summary of the similarities Mendel’s principle of segregation of factors could now be
between events occurring during meiosis and explained in terms of the separation (segregation) of
fertilisation and Mendel’s hypotheses. homologous chromosomes which occurs during anaphase I
aeaN{Cq“_60@a=0eE2F—0—8—O"aa=ow"aoaoooooooooooonaououqQqqoeee
oo
of meiosis and the random distribution of alleles into
Meiosis and fertilisation Mendel’s hypotheses gamete cells. These events are summarised in fig 23.6.
Diploid cells contain pairs of Characteristics are controlled
chromosomes (homologous by pairs of factors 23.2.1 Chromosomal explanation of
chromosomes) independent assortment
Homologous chromosomes Pairs of factors separate Mendel’s principle of independent assortment
separate during meiosis during gamete formation may also be explained in terms of the movement of
One homologous chromosome Each gamete receives one chromosomes during meiosis. During gamete formation
passes into each gamete cell factor the distribution of each allele from a pair of homologous
Only the nucleus of the male Factors are transmitted from chromosomes is entirely independent of the distribution of
gamete fuses with the egg cell generation to generation as
nucleus
alleles of other pairs. This situation is described in fig 23.7.
discrete units
It is the random alignment or assortment of homologous
Homologous pairs of chromo- Each organism inherits one
somes are restored at fertilisa- factor from each parent
chromosomes on the equatorial spindle during metaphase I
tion, each gamete (C" and 9) of meiosis, and their subsequent separation during meta-
contributing one homologous phase I and anaphase I, that leads to the variety of allele
chromosome recombinations in the gamete cells. It is possible to predict
the number of allele combinations in either the male or
female gamete using the general formula 2”, where n =
forms of the gene or alleles which determine the expression haploid number of chromosomes. In the case of Man,
of the gene. Alleles are the alternative forms in which a where n = 23, the possible number of different combina-
gene may exist and they occupy corresponding positions or tions is 2” = 8 388 609.
loci (singular locus) on homologous chromosomes, as shown
in fig 23.5.
Fig 23.5 A cell showing two pairs of homologous

chromosomes. The positions of two different gene loci are
indicated by circles. In this example two gene loci are shown
situated on different pairs of homologous chromosomes and
each gene is present as two alleles
haploid cells
prophase II
anaphase I separate
haploid cells
homologous each carrying
chromosomes one allele of the
original pair
homologous chromosomes
(each chromosome appears
as a pair of chromatids
carrying the alleles A and a)
d in terms of the separation of homologous
Fig 23.6 Mendel’s principle of segregation of factors (alleles) A anda describe
chromosomes which occurs during meiosis
837
subsequent
nuclear divisions
during metaphase I homologous
have given rise
chromosomes may line up on the
to four types
equatorial spindle either as shown
of gametes as
above or as below; this is
shown above
independent assortment
and below
two pairs of homologous

chromosomes bearing the
alleles A, a, B, bas seen
during prophase I
Fig 23.7 Mendel’s principle of independent assortment of factors (alleles) A, a, B, b, described in terms of the separation of
homologous chromosomes which occurs during meiosis
different alleles): grey and black body, and long and

23.3 Linkage vestigial (short) wings. Grey body and long wing are
dominant. If pure-breeding grey-bodied long-winged Dro-
All the situations and examples discussed so
far in this chapter have dealt with the inheritance of genes sophila are crossed with black-bodied vestigial-winged
situated on different chromosomes. Cytological studies Drosophila, the expected F, phenotypic ratio would be
have revealed that Man possesses 46 chromosomes in all 9:3:3:1. This would indicate a normal case of Mendelian
the somatic (body) cells. Since Man possesses thousands of dihybrid inheritance with random assortment resulting
characteristics such as blood group, eye colour and the from the genes for body colour and wing length being
ability to secrete insulin, it follows that each chromosome situated on non-homologous chromosomes. However this
must carry a large number of genes. result is not obtained. Instead the F, show an approxi-
Genes situated on the same chromosome are said to be mately 3:1 ratio of parental phenotypes. This may be
linked. All genes on a single chromosome form a linkage explained by assuming that the genes for body colour and
group and usually pass into the same gamete and are wing length are found on the same chromosome, that is
inherited together. As a result of this, genes belonging to they are linked, as shown in fig 23.8.
In practice, though, this 3:1 ratio is never achieved and
the same linkage group usually do not show independent
assortment. Since these genes do not conform to Mendel’s four phenotypes are invariably produced. This is because
principle of independent assortment they fail to produce total linkage is rare. Most breeding experiments involving
the expected 9:3:3:1 ratio in a breeding situation involving linkage produce approximately equal numbers of the
parental phenotypes andasignificantly smaller number of
the inheritance of two pairs of contrasted characteristics
phenotypes showing new combinations of characteristics,
(dihybrid inheritance). In these situations a variety of
also in equal numbers. These latter phenotypes are
ratios are produced which may be explained quite simply
described as recombinants. From this it is possible to
now that we possess a basic understanding of the
mechanisms of inheritance as revealed by Mendel. (At this produce the following definition of linkage.
point it is worth re-emphasising Mendel’s good fortune in Two or more genes are said to be linked when pheno-
choosing to study the inheritance of pairs of characteristics types with new gene combinations (recombinants) occur
located on different chromosomes.) In Drosophila the less frequently than the parental phenotypes.
genes for body colour and wing length have the following The events leading to the discovery of linkage by the
allelomorphs (phenotypic characteristics determined by American Thomas H. Morgan may be summarised in one
838
Let:
G represent grey body (dominant) Fig 23.8 Genetic explanation of the 3:1 ratio produced in Fo
g represent black body (recessive)
phenotypes asa result of linkage
L represent long wing (dominant)
l represent vestigial wing (recessive)
Parental phenotypes grey body,long wing x __ black body, vestigial

wing
ei A of his experiments in which he predicted the results of a
8 8 backcross between heterozygous grey-bodied long-winged
Parental genotypes (2n) x Drosophila (the F, generation of the experimental cross
. x shown in fig 23.8) and homozygous recessive black-bodied
l l vestigial-winged Drosophila. The two possible outcomes
were predicted as follows:
Meiosis (1) If the four alleles for grey and black body, and long and
vestigal wings, were on different pairs of chromosomes
G g (that is not linked) they should show independent
assortment and produce the following phenotypic
Gametes (n) x ratios: P & P YP
L I 1 grey body, long wing: 1 grey body, vestigial wing: 1

black body, long wing: 1 black body, vestigial wing.
(2) If the alleles for body colour and wing length were
situated on the same pair of chromosomes (that is
linked) the following phenotypic ratio would be
G g produced:
F, genotypes (2n) 1 grey body, long wing:1 black body, vestigial wing.
An explanation of these predictions is given in fig 23.9.
L l Morgan carried out this backcross several times and
never obtained either of the predicted outcomes. Each
F, phenotypes all heterozygous grey body, long-winged offspring time he obtained the following results:
The F, generation was allowed to interbreed 41.5% grey body long wing
F, phenotypes grey body, long wing X_ grey body, long wing 41.5% black body vestigial wing
8.5% grey body vestigial wing
8.5% black body long wing
On the basis of these results he postulated that:
F, genotypes (2n) x (1) the genes were located on chromosomes,
(2) both the genes were situated on the same chromosome,
that is linked,
(3) the alleles for each gene were on homologous chromo-
somes,
Meiosis (4) alleles were exchanged between homologous chromo-
somes during meiosis.
/G The reappearance of recombinant alleles in 17% of the
offspring was explained in terms of point (4). This is known
Gametes (n) x as crossing-over.
f . |. - 23.7 nh ‘homozygous. -purple-flowered

/ yrt-stemmed plant was crossed with a homo- —
__zygous red-flowered long-stemmed plant and the F, —
Random fertilisation - phenotypes had purple flowers and short stems.
n theF, generation was testcrossed (back-
crossed) with a double homozygous recessiveee _
the follo ng progeny were EN
ae senor pes Or) 52purple flower, short stem—
7 purple flower, long stem-
Z a a ' a 9 red flower, short stem
46 red flower, long stem
——————OO
n fullythese results,
Foe 839
(a) If the four alleles are situated on different pairs of chromosomes Fig 23.9 (left) (a) and (b) Genetic explanation of Morgan’s
predictions
Testcross phenotypes grey body,longwing ~_ black body, vestigial wing
(heterozygous) (homozygous)
Testcross genotypes (2n) GgLl Xx gell
Meiosis 23.3.1 Crossing-over and crossover values

Gametes (n) In 1909 the Belgian cytologist Janssens
(as shown by observed chiasmata formation during prophase I of meiosis
cand 2) of (section 22.3). The genetic significance of this process was
GL | Gl | gL gl
clarified by Morgan who proposed that crossing-over of
(as shown in Punnett : GL Gl eL gl alleles occurred as a result of the breakage and recombina-
square) g ol al ‘l al tion of homologous chromosomes during chiasmata.
Subsequent research based on the microscopic examina-
tion of cells and recombinant phenotypic ratios has
confirmed that crossover of genetic material occurs
Offspring genotypes (2n) between virtually all homologous chromosomes during
(listed in each square) meiosis. The alleles of parental linkage groups separate
and new associations of alleles are formed in the
Offspring phenotypes 1 grey body, long wing: | grey body, vestigial wing:
1 black body, long wing: | black body, vestigial wing
gamete cells, a process known as genetic recombination.
Offspring formed from these gametes showing ‘new’
(b) If the four alleles are situated on the same pair of chromosomes combinations of characteristics are known as recombi-
Testcross phenotypes grey body, long wing black body, vestigial
nants. Thus crossing-over is a major source of observable
(heterozygous) wing (homozygous) genetic variation within populations.
The behaviour of a pair of homologous chromosomes in
G g s g Drosophila, carrying the alleles grey body and long wing
(both dominant) and black body and vestigial wing (both
Testcross genotypes (2n) x
recessive), during formation of chiasmata may be used to
L l l l illustrate the principle of crossing-over. A cross between a
male homozygous grey-bodied long-winged Drosophila
Meiosis and a female homozygous black-bodied vestigial-winged
Drosophila produced heterozygous F, offspring with
grey bodies and long wings as shown in fig 23.10.
Gametes (n) grey body, black body,

Testcross phenotypes long wing x vestigial wing
(heterozygous) (homozygous)
G g g g
Random fertilisation Testcross genotypes (2n) x
tut
Qa l ]
7a re ra
Offspring genotypes (2n)
Meiosis
L ] l I (showing crossing-over)
wi 7
Offspring phenotypes 1 grey body, 1 black body,
long wing vestigial wing
Gametes (n)
Fig 23.10 (right) Genetic explanation of crossing-over and the

appearance of recombinant genotypes. The recombination
Offspring genotypes (2n) aie,
frequency can be calculated by counting the number of
individuals showing recombination and the total number of
individuals and applying the following formula:
reootAOREN Geico
genotypes
DERE
(x)
recombination frequency (%) = 7 x 100
total genotypes (y)
840
Backcrossing the F, generation flies with homozygous
double recessive flies produced the following results.
Parental phenotypes grey body, long wing 965

black body, vestigial wing 944
Recombinant black body, long wing 206
phenotypes grey body , vestigial wing 185
These results indicate that the genes for body colour and
wing length are linked. (Remember that a dihybrid cross
between an F, heterozygote and a double homozygous
recessive would have produced a 1:1:1:1 phenotypic ratio if
the genes had been situated on different chromosomes and
therefore had undergone random assortment.) Using the
figures obtained from the above cross it is possible to
calculate the recombination frequency of the genes for
body colour and wing length.
The recombination frequency is calculated using the
formula
number of individuals showing recombination na

Sh — XX
number of offspring
From the example above the recombination frequency (%)

is
(206 + 185) .
100
(965 + 944) + (206 + 185)
= 391 x 100
2300
=17%. 23.4 Gene mapping
This value indicates the number of crossovers which have The major significance of calculating cross-
occurred during gamete formation. A. H. Sturtevant, a over frequencies is that it enables geneticists to produce
student of Morgan, postulated that the recombinant maps showing the relative positions of genes on chromo-
frequency or crossover frequency (crossover value (COV)) somes. Chromosome maps are constructed by directly
demonstrated that genes are arranged linearly along the converting the crossover frequency or value between genes
chromosome. More importantly, he suggested that the into hypothetical distances along the chromosome. A
crossover frequency reflects the relative positions of genes crossover frequency or value (COV) of 4% between genes
on a chromosome because the further apart linked genes A and B means that those genes are situated 4 units apart on
are on the chromosomes, the greater the possibility of the same chromosome. A COV of 9% for a pair of genes A
crossing-over occurring between them, that is the greater and C would indicate that they were 9 units apart, but it
the crossover frequency (fig 23.11). would not indicate the linear sequence of the genes, as
shown in fig 23.12.
—_——__—_@—_—_e——___M—_o——_
A B C C? A B Ge
Fig 23.11 Three gene loci represented by A, B and C are

9 >
shown on the chromosome. Crossing-over and separation of
genes is more likely to occur between A and C than between
Fig 23.12 Possible gene loci of A, B and C on the basis of
B and C or A andB since the frequency of crossing-over is
the data presented
related to the distance between the genes
841
In practice it is usual to determine crossover values for at
least three genes at once, as this triangulation process
enables the sequence of the genes to be determined as well
as the distances between them. Consider the following
crossover values as determined by a series of breeding
experiments involving four genes P, Q, R and S.
P-Q =24%
R-P =14%
R-S = 8% (b)
a B c
S-P = 6%
Fig 23.14 (a) A pair of homologous chromatids, one carrying
To calculate the sequence and distances apart of the genes, the dominant alleles A, B and C and the other carrying the
a line is drawn representing the chromosome and the recessive alleles a, b and c. Crossing-over occurs at two
following procedure carried out. points *; and *2. (b) The result of separation of the chromatids
in which the sequences of alleles are different, although the
(1) Insert the positions of the genes with the least COV in sequences of gene loci and the distances between them
the middle of the chromosome, that is S — P = 6% remain the same
(fig 23.132).
(2) Examine the next largest COV, that is R — S = 8%,
- 23.9 In maize the genes for coloured ©
and insert both possible positions of R on the chromo-
some, relative to S (fig 23.135).
seed and full seed are dominant to the genes for —
colourless seed and shrunken seed. Pure-breeding _
(3) Repeat the procedure for the next largest COV, that is
R — P = 14%. This indicates that the right-hand pos-
strains of the double dominant variety were crossed _
with the double recessive variety and a backcross of
ition of R is incorrect (fig 23.13c).
the F, generation produced the following results.
(4) Repeat the procedure for the COV for P — Q = 24%
(fig 23.13d). The position of Q cannot be ascertained coloured, fullseed = $380
without additional information. If, for example, the colourless, shrunken seed 396
COV for Q — R = 10% this would confirm the left- coloured, shrunken seed 14
hand position for gene Q. colourless, full seed = 10°
Ss P
Calculate the distance in units between the genes for
—________e_0——
Tp _ coloured seed and seed shape on the chromosomes.
(a) — 6 —>
R? S) P R?
——___—_@—__—_e——_6-.
$$$
(b) <— 8 —>—_6
—>
R
¢—
S
8 —>
E
23.5 Linkage groups and
——_oe—_——_—_o——_9 chromosomes
() ¢— 8 —pe—
6 —>
<—_———
14 —_»
Q? R S P Q? Much of the evidence presented in this chapter
—_@——_____@—___e—__e_________"__e—_-
(d) so far has shown how our knowledge of the mechanics of
i 24 bie 24 >
inheritance has gradually increased. Most of the research
Fig 23.13 Use of the triangulation process to establish the into genetics in the early part of this century involved
positions of genes P, Q, R and S on a chromosome establishing the role of genes in inheritance. Morgan’s
research with the fruit fly (Drosophila melanogaster)
established that the majority of phenotypic characteristics
were transmitted together in four groups and these were
called linkage groups. It was observed that the number of
linkage groups corresponded to the number of pairs of
chromosomes.
Studies on other organisms produced similar results.
Breeding experiments using a variety of organisms re-
A problem which arises in preparing chromosome maps vealed that some linkage groups were larger then others
is that of double crossover, particularly when considering (that is they carried more genes). Examination of chromo-
genes which are widely separated, since the number of somes in these organisms showed that they varied in length.
apparent crossovers will be less than the actual number. Morgan demonstrated that there was a distinct relationship
For example, if crossovers occur between alleles A and B between these observations. This provided further con-
and B and C in fig 23.14, A and C will still appear linked, firmatory evidence that genes were located on chromo-
but the chromosome will now carry the recessive allele b. somes.
842
23.5.1 Giant chromosomes and genes
23.6 Sex determination
In 1913 Sturtevant began his work on mapping
the positions of genes on the chromosomes of Drosophila The technique of relating phenotypic charac-
but it was 21 years before there was a possibility of linking teristics of organisms to the structure of their chromo-
visible structures on chromosomes with genes. In 1934, it somes, as described in earlier sections, is seen most clearly
was observed that the chromosomes in the salivary gland in the determination of sex. In Drosophila the observed
cells of Drosophila were about 100 times larger than phenotypic differences between the two sexes appear to be
chromosomes from other body cells. For some reason these related to the differences in the size of their chromosomes,
-o-
chromosomes duplicate without separating until there are as shown in fig 23.16. Examination of the chromosome
several thousand lying side by side. When stained they can structure of a range of animals revealed that males and
be seen with the light microscope and appear to be made up females showed certain chromosomal differences. Pairs of
of alternating light and dark bands. Each chromosome has
its own distinctive pattern of bands (fig 23.15). It was I \G We
originally thought, or rather hoped, that these bands were
genes, but this is not the case. Phenotypic abnormalities
may be artificially induced in Drosophila and these
correlate with changes in chromosomal banding patterns,
as observed with the microscope. These phenotypic and
chromosomal abnormalities in turn correlate with gene loci female male
shown on chromosome maps which have been constructed
on the basis of crossover values obtained from breeding Fig 23.16 Structure of chromosomes in male and female
Drosophila melanogaster. Four pairs of chromosomes are
experiments. Therefore it is possible to say that the bands shown. The sex chromosomes are numbered |
on the chromosomes indicate the positions of genes but are
not themselves genes.
chromosomes (homologous chromosomes) are found in all
cells, but one pair of chromosomes always shows dif-
ferences between the sexes. These are the sex chromosomes
or heterosomes. All other chromosomes are known as
autosomal chromosomes or autosomes. As can be seen in fig
23.16, Drosophila has four pairs of chromosomes. Three
pairs appear identical in both sexes (numbers II, III and
IV), but the other pair, whilst appearing identical in the
female, differ in the male. The chromosomes are known as
X and Y chromosomes, and the genotype of the female is
XX and that of the male is XY (fig 23.17). These char-
acteristic sex genotypes are found in most animals,
including Man; but in the case of birds (including poultry),
moths and butterflies the sex genotypes are reversed: the
females are XY and the males are XX. In some insects,
such as the grasshopper, the Y chromosome may be
absent entirely and so the male has the genotype XO.
Fig 23.15 Giant chromosomes from the salivary glands of

Drosophila melanogaster. Four pairs of chromosomes are Fig 23.17 Human sex chromosomes as they appear during
shown joined at their centromeres metaphase of mitosis
843
Fig 23.18 Genetic explanation of the sex ratio in humans as opposed to sex-linked inheritance and in humans is
thought to cause suppression of the genes for growth of
Parental phenotypes female x male
() beard in females.
(2)
Parental genotypes (2n) XX x XY Morgan and his co-workers noticed that inheritance of
eye colour in Drosophila was related to the sex of the
Meiosis
parent flies. Red eye is dominant over white eye. A
Gametes(n) (x) (x) x (x) (¥) red-eyed male crossed with a white-eyed female produced
equal numbers of F, red-eyed females and white-eyed
males (fig 23.19a). A white-eyed male, however, crossed
Offspring genotypes (2n) XX XY XX XY with a red-eyed female produced equal numbers of F,
Offspring phenotypes 2 o.. 2 a red-eyed males and females (fig 23.19b). Inbreeding these
sex ratio 1 female :1 male
F, flies produced red-eyed females, red-eyed males and
white-eyed males but no white-eyed females, (fig 23.19c).
In the production of gametes the sex chromosomes The fact that male flies showed the recessive characteristic
segregate in typical Mendelian fashion. For example, in more frequently than female flies suggested that the white
mammals each ovum contains an X chromosome; in males eye recessive allele was present on the X chromosome and
one half of the sperms contains an X chromosome and the that the Y chromosome lacked the eye colour gene. To test
other half contains a Y chromosome as shown in fig 23.18. this hypothesis Morgan crossed the original white-eyed
The sex of the offspring depends upon which type of sperm male with an F, red-eyed female (fig 23.19d). The offspring
fertilises the ovum. The sex having the XX genotype is included red-eyed and white-eyed males and females.
described as homogametic as it produces gamete cells From this Morgan rightly concluded that only the X
containing only X chromosomes. Organisms with the XY chromosome carries the gene for eye colour. There is no
genotype are described as heterogametic since half their gene locus for eye colour on the Y chromosome. This
gametes contain the X chromosome and half the Y phenomenon is known as sex linkage.
chromosome. In humans, the genotypic sex of an indi-
vidual is determined by examining non-dividing cells. One
X chromosome always appears in the active state, which
has the normal appearance. If another is present, it is seen 23.10 In Drosophila t for wing
in a resting state as a tightly coiled dark-staining body
called the Barr body. The number of Barr bodies is always
_ length and for eye colour aresé al wing
and red eye are dominant t to ngand
one less than the number of X chromosomes present, that is white eye.
male (XY) = 0, female (XX) = 1. The function of the Y (a) In a cross between a, miniature wing, red-eyed
chromosome appears to vary according to species. In Man
male and a homozygous normal wing, white-eyed ©
the presence of a Y chromosome controls the differentia- female, explain fully the appearance of (i) the F,
tion of the testis which subsequently influences the and (ii) the F, generations.
development of the genital organs and male characteristics
(6)Crossing a female from the F, generation above
(section 20.3.1). In most organisms, however, the Y
with a miniature wing; white-eyed male gave the
chromosome does not carry genes concerned with sex. In
fact it is described as genetically inert or genetically empty
ing,
s, white-eyed males and females 35 _
since it carries so few genes. In Drosophila it is thought that ing, red-eyed males and females 7
the genes determining male characteristics are carried on miniature wing, white-eyed males and females 18 —
the autosomes and their phenotypic effects are masked by miniature wing, red-eyed males and females 36
the presence of a pair of X chromosomes. Male characteris- Account for the appearance and numbers of ihe
tics, on the other hand, appear in the presence of a single X phenotypes shown above.
chromosome. This is an example of sex-limited inheritance,
844
Let: R represent red eye (dominant)
r represent white eye (recessive)
XX represent female fly (Q)
XY represent male fly (o’)
Parental phenotypes red-eyed S) x white-eyed 9 white-eyed S x red-eyed ©

Parental genotypes (2n) xRy x Xrxr xTy x xRxR
Meiosis
Gametes (n) (y) x Gr) (xr) (xr) (y) x (xR
Random fertilisation é x y xR)
F, genotypes (2n) xRxr xRxr xry Xry XxrxR XxrxR xRy xRy
F\ phenotypes red-eyedQ white-eyed red-eyed 9 red-eyed
(a) (b)
F, phenotypes red-eyed o” x red-eyed © white

-eyed 07 x red-eyed ©
F genotypes (2n) xRy Xx xrxR XTy x xRxr
co O-® © © ©-®@® ®@
Meiosis
F, genotypes (2n) xRxr xRxR_ xry xRy xrxR Xrxr xRy XTy
F, phenotypes red-eyed 2 white-eyed red-eyed red-eyed white-eyed, _red-eyed white-eyed
Ree aa Ace ene)
Q On
() (d)
Fig 23.19 (a) and (b) Morgan’s reciprocal experimental rate of blood clotting. The gene for substance VIII is
crosses between red-eyed and white-eyed Drosophila. Note carried on the non-homologous portion of the X chromo-
the low frequency of appearance of white eyes. (c) Morgan’s some and can appear in two allelomorphic forms: normal
confirmatory inbreeding experimental cross between an F,
(dominant) and mutant (recessive). The following possible
red-eyed male and an F, (heterozygous) red-eyed female.
(d) The experimental cross between a white-eyed male and genotypes and phenotypes can occur.
an F, (heterozygous) red-eyed female. Note the appearance of
the white-eyed characteristic only in homozygous white-eyed genotype phenotype
female flies xX X normal female
ue. normal female (carrier)
xXhy normal male
23.6.1 Sex linkage UY, haemophiliac male
Genes carried on the sex chromosomes are In all sex-linked traits, females who are heterozygous are
said to be sex-linked. In the case of the heterogametic sex described as carriers of the trait. They are phenotypically
there is a portion of the X chromosome for which there is normal but half their gametes carry the recessive gene.
no homologous region of the Y chromosome (fig 23.20). Despite the father having a normal gene there is a 50%
probability (probability 2) that sons of carrier females will
show the trait. In the situation where a carrier haemophi-
non-homologous portion liac female marries a normal male they may have children
of the X chromosome
with phenotypes as shown in fig 23.21.
homologous portion Let: H represent normal allele for blood clotting (dominant)
of the sex chromosomes h represent allele for haemophilia (recessive)
XX represent female chromosomes
x Y XY represent male chromosomes
Fig 23.20 Homologous and non-homologous regions of the Parental phenotypes normal female (carrier) X normal male
sex chromosomes Parental genotypes (2n) XH xh x xHy
Meiosis
Characteristics determined by genes carried on the non- Gametes (n) (xH) (xh) x ) (y)
homologous portion of the X chromosome therefore Random fertilisation
appear in males even if they are recessive. This special form Offspring genotypes (2n) XH XH xHy xh xH xhy
of linkage explains the inheritance of sex-linked traits such
Offspring phenotypes normal normal normal haemophiliac
as red-green colour blindness, premature balding and female male female male
haemophilia. Haemophilia or ‘bleeder’s disease’ is a (carrier)
sex-linked recessive condition which prevents the forma- Fig 23.21 Mechanism of inheritance of the
tion of factor VIII, an important factor in increasing the sex-linked allele for haemophilia
845
Queen Victoria (e) {| Prince Albert
ta ; [ ©)
Alice} Ludwig IV Leopold | Helen of Beatrice| Henry of
Frederick X]Victoria, _Edward VII} Queen Battenburg
German Empress of King of Alexandra of Hesse of Albany] Waldeck
Emperor |Germany Britain
(e) T] ©
Sophi George V ()
Irene of Prince Victoria}Prince Czarina |Czar Alice] Alexander Leopold Maurice Ones Alphonso
WaldemarSigigiismung ar Hesse Frederick Louis of | Alexandra| Nicholas of Teck ueen | XIII King
Battenburg of Russia of Spainjof Spain
YA GeorgeVI{_] Henry Waldemar () Alice G

lived to 56 Czarivitch Maurice Rupert Alfonso Gonzalo
Alexander died at 4 Pio Manvel
©) Alexis died in (Lord Trematon)
Elizabeth II Prince Philip murdered infancy BL eee
he Nota eatcic after accidents
Andrew Edward (@) Carrier female
Oars ba Kom male 7
Diana|Charles Anne] Mark Phillips
a e) Possible male haemophiliac
CI L Peter Zara i Male haemophiliac

William Henry
One of the best-documented examples of the inheritance Fig 23.22 Transmission of haemophilia in the descendants
of haemophilia is shown by the descendants of Queen of Queen Victoria. In the diagram only those descendants
Victoria. It is thought that the gene for haemophilia arose involved in the transmission and appearance of haemophilia
have been shown. The ancestry of the British Royal Family
as a mutation in Queen Victoria or one of her parents. has been given to show why haemophilia is absent from
Fig 23.22 shows how the haemophilia gene was inherited by seven generations of Queen Victoria's descendants
her descendants.
(2311. Body colour in cats and magpie moths iscontrolled bya sex-linked gene on the X chromosome. -
The following data were obtained in two breeding experiments where the homogametic sex was homozygous for
body colour in the parental generation. =
th a
Ca
colour dominant (black colour dominant _
to yellow colour) _
:Parental phenotypes pale male x normal female black male x yellow female
: Offspring phenotypes 1 normal male : 1 pale female - 1 yellow male : 1 black female
Which is the heterogametic sex in each of these organisms? —
23.7 Gene interactions situation described by Mendel in his monohybrid breeding

experiments. It is fortunate that he did not select organisms
The topics in this chapter so far have re- which show this condition as it may have unnecessarily
presented the simpler aspects of genetics: dominance, complicated his early work.
monohybrid and dihybrid inheritance, linkage, sex deter- Incomplete dominance is found in both plants and
mination and sex linkage. There are many situations in animals. In most cases the heterozygote has a phenotype
genetics where genes interact in ways other than those which is intermediate between the homozygous dominant
already described and it i: probable that the majority of and recessive conditions. An example is the production of
phenotypic characteristics in organisms result from these. blue Andalusian fowls by crossing pure-breeding black and
Several types of gene interaction will now be considered. splashed white parental stocks. The presence of black
plumage is the result of the possession of an allele for the
production of the black pigment melanin. The splashed
23.7.1 Incomplete dominance
white stock lack this allele. The heterozygotes show a
There are several conditions where two or partial development of melanin which produces a blue
more alleles do not show complete dominance or recessive- sheen in the plumage.
ness due to the failure of any allele to be dominant in the As there are no accepted genotypic symbols for alleles
heterozygous condition. This state of incomplete domi- showing incomplete dominance, the importance of specify-
nance (co-dominance or blending) is an exception to the ing symbols in genetic explanations is apparent. For
846
example, in the case of the Andalusian fowl, the following Table 23.4 Examples of incomplete dominance.
genotypic symbols may be used to illustrate the alleles:
black- B; splashed white— b, W, B” or B®”. The results Characteristic Alleles Heterozygous
of a cross between black and splashed white homozygous phenotype
fowl are shown in fig 23.23.
Antirrhinum flower red X white pink
(snapdragon)
Bet:
B represent the black allele Mirabilis flower red X white pink
BW represent the splashed white allele (four-o’clock flower)
Parental phenotypes black x splashed white Short-horn cattle red X white roan
(homozygous) (homozygous) Angora and rex long hair and intermediate silky fur
Parental genotypes (2n) BB x BW BW rabbits short hair
Meiosis
Gametes (n)
23.7.2 Multiple alleles
-
Random fertilisation In all the cases studied so far, each character-
istic has been controlled by a gene which may have
F, genotypes (2n) BBW BBW BBW BBW appeared in one of two forms or alleles. There are several
F,, phenotypes all ‘blue’ heterozygotes conditions where a single characteristic may appear in
Fig 23.23
several different forms controlled by three or more alleles,
The production of F; hybrids of Andalusian fowl
of which any two may occupy the same gene loci on
homologous chromosomes. This is known as the multiple
allele (or multiple allelomorph) condition and it controls
If the F, generation are allowed to interbreed, the F, such characteristics as coat colour in mice, eye colour in
generation shows a modification of the normal Mendelian mice and blood group in Man.
phenotypic monohybrid ratio of 3:1. In this case a
phenotypic ratio of 1:2:1 is produced where half the F, Inheritance of blood groups
generation have the F, genotype (fig 23.24). This ratio of Blood group is controlled by an autosomal gene. The gene
1:2:1 is characteristic of examples of incomplete domi- locus is represented by the symbol I (which stands for
nance. Other examples are shown in table 23.4. isohaemagglutinogen) and there are three alleles re-
presented by the symbols A, B and O. The alleles A and B
F, phenotypes blue x blue are equally dominant and O is recessive to both. The
F, genotypes (2n) BBW x BBW genotypes shown in table 23.5 determine the phenotypic
appearance of blood groups. The presence of a single
Meiosis
dominant allele results in the blood producing a substance
Gametes (n) x called agglutinin ywhich acts as an antibody. For example,
the genotype I“ I° would give rise to the agglutinogen A on
the red blood cell membrane, and the plasma would
F, genotypes (2n) BB BBW BBW BW BW contain the agglutinin anti-B (the blood group would be
F, phenotypes black blue splashed
A). Blood-grouping is described in section 14.14.6.
1 : 2 white 1
Table 23.5 Human blood group genotypes.
Fig 23.24 The production of Fz hybrids of Andalusian fowl
Blood group (phenotype)
847
2g
Fig 23.26 Variation in comb shape in domestic fowl (top left) sin gle comb, (top right) pea comb, (bottom left) rose comb,
(bottom right) strawberry comb
848
23.7.3 Lethal genes be controlled by the interaction of two or more genes
There are several examples of conditions situated at different loci. In the case of the inheritance of
where a single gene may affect several characteristics, the shape of the comb in domestic fowl there are genes at
including mortality. In the case of Man and other mam- two loci situated on different chromosomes which interact
mals a certain recessive gene may lead to internal adhesions and give rise to four distinct phenotypes, known as pea,
of the lungs resulting in death at birth. Another example rose, walnut and single combs (fig 23.26). The appearance
involving a single gene affects the formation of cartilage of pea comb and rose comb are each determined by the
and produces congenital deformities leading to foetal and presence of their respective dominant allele (P or R) and
neonatal death. the absence of the other dominant allele. Walnut comb
In chickens which are homozygous for an allele controll- results from a modified form of incomplete dominance in
ing feather structure called ‘frizzled’, several phenotypic which at least one dominant allele for pea comb and rose
effects result from the incomplete development of the comb is present (that is PR). Single comb appears only in
feathers. These chickens lack adequate feather insulation the homozygous double recessive condition (that is pprr).
and suffer from heat loss. To compensate for this they These phenotypes and genotypes are shown in table 23.6.
exhibit a range of structural and physiological adaptations, The F, genotypes and F, phenotypic ratios resulting from
but these are largely unsuccessful and there is a high crossing a pure-breeding pea-comb hen with a pure-
mortality rate. breeding rose-comb cock are shown in fig 23.27.
The effects of a lethal gene are clearly illustrated by the Table 23.6 Phenotypes and possible genotypes
inheritance of fur colour in mice. Wild mice have grey- associated with comb shape in poultry.
coloured fur, a condition known as agouti. Some mice
have yellow fur. Cross-breeding yellow mice produces Phenotype Possible genotypes
offspring in the ratio 2 yellow fur : 1 agouti fur. These
results can only be explained on the basis that yellow is pea PPrr, Pprr
rose RRpp, Rrpp
dominant to agouti and that all the yellow coat mice are
walnut PPRR, PpRr, PPRr, PpRr
heterozygous. The atypical Mendelian ratio is explained by single pprr
the foetal death of homozygous yellow coat mice (fig
23.25). Examination of the uteri of pregnant yellow mice
ee Y represent yellow fur (dominant)

y represent agouti fur (recessive)
Parental phenotypes yellow fur x yellow fur
Parental genotypes (2n) Yy x Yy
Meiosis
Gametes (n) (¥) (y) Xx (Y) Yy)
Offspring genotypes (2n) YY Yy Yy yy
Offspring phenotypes 1 yellow fur: 2 yellow fur : 1 agouti fur

die before birth
Fig 23.25 Genetic explanation of fur colour inheritance in

mice showing the lethal genotype YY
from the above cross revealed dead yellow foetuses.

Similar examination of the uteri of crosses between yellow
fur and agouti fur mice revealed no dead yellow foetuses.
The explanation is that this cross would not produce
homozygous yellow (YY) mice.
23.7.5 Epistasis
23.7.4 Gene-complex A gene is said to be epistatic (epi, over) when
The presence of a pair of alleles occupying a its presence suppresses the effect of a gene at another locus.
given gene locus and controlling the production of a single Epistatic genes are sometimes called ‘inhibiting genes’
phenotypic characteristic is true in some cases only and because of their effect on the other genes which are
exceptional in most organisms. Most characteristics are described as hypostatic (hypo, under).
determined by the interaction of several genes which form Fur colour in mice is controlled by a pair of genes
a ‘gene-complex’. For example, a single characteristic may occupying different loci. The epistatic gene determines the
849
Let: - :
P represent presence of pea comb (dominant)
p represent absence of pea comb (recessive)
R represent presence of rose comb (dominant)
r represent absence of rose comb (recessive)
pea comb x rose comb
Parental phenotypes
PPrr x RRpp
Parental genotypes (2n)
Meiosis
Gametes (n)
‘ (Pe)
F, genotypes (2n) all PpRr
F, phenotypes all walnut comb
F phenotypes walnut comb x walnut comb
F, genotypes (2n) PpRr x PpRr
Meiosis
os PR Pr pR pr
Gametes (n) (as shown byofandQ ) PR Pr pR PE
fe PR 6 PR PR 6 PR: ae
ame
Random fertilisation P PR Pr pR pr
Pr Oo} Pr Pr O Pr gO
F, genotypes (2n) (shown in Punnett square) R PR Pr pR pr
‘ PR pR pRittran pRig 7
PR Pr pR pr
ea Pro pr pore Pre
F, phenotypes 9 walnut comb:3 pea comb: 3 rose comb: 1 single comb
Offspring symbols O O A oO
Fig 23.27 Genetic explanation of comb inheritance in fowl
Let: A represent agouti fur (dominant)

a represent black fur (recessive)
C represent coloured fur (dominant)
c represent albino fur (recessive)
Parental phenotypes agouti x albino
Parental genotypes (2n) AaCc Aacc
Meiosis
Gametes (n)(as shown by of and Q) 9 ow AC Ac aC | ac
Ke AC Ac aC ac
Ac O Ac oO Ac O Ac i
at AC Ac aC ac
Offspring genotypes (2n) (as shown in Punnett square)
ac O ac Oo ac A ac oO
Offspring phenotypes 3 agouti : 4 albino: | black

Offspring symbols O O A
Fig 23.28 A genetic explanation of how unusual phenotypic
ratios can be produced in the case of epistatic genes Table 23.7 Some examples of the range of phenotypic
ratios which can be produced as a result of epistatic
presence of colour and has two alleles, coloured (domi- gene interactions (see fig 23.28 for explanation of
nant) and albino (white) (recessive). The hypostatic gene alleles).
determines the nature of the colour and its alleles are
agouti (grey) (dominant) and black (recessive). The mice Parental
may have agouti or black fur depending upon their phenotypes Genotypes Phenotypic ratios
genotypes, but this will only appear if accompanied by the agouti X agouti AaCe x AaCe ‘9 agouti:3 black:4 albino
allele for coloured fur. The albino condition appears in agouti X black AaCc X aaCc 3 agouti:3 black :2 albino
mice that are homozygous recessive for colour even if the agouti X albino AaCc X Aacc 3 agouti:1 black:4 albino
alleles for agouti and black fur are present. Three possible agouti X albino AaCc X aacc 1 agouti: 1 black:2 albino
phenotypes can occur and they are agouti, black and agouti X albino AACc X aacc 1 agouti:1 albino
agouti X black AaCe X aaCC 1 agouti:1 black
albino. A variety of phenotypic ratios can be obtained albino X black AAce X aaCC all agouti
depending on the genotypes of the mating pair (fig 23.28 albino x black AdAce X aaCc 1 agouti:1 albino
and table 23.7). Ee eee
850
23.15 In White Leghorn fowl, plumage
_ Colour is co trolled by two sets of genes, including
A (a) A (b)
__ the following:
__W (white) dominant over w (colour)
____ B (black) dominant over b (brown). .
__ The heterozygous F, genotype WwBb is white.
_ Account for this type of gene interaction and show
_ thephenotypic ratio of the F,generation.
Number
of
organisms of
Number
organisms
23.7.6 Polygenic inheritance

Many of the most obvious characteristics of
organisms are produced by the combined effect ofman
y Characteristic Characteristic
different genes. These genes form a special gene comp
lex Fig 23.29 Histograms representing frequency distribution in
known as a polygenic system. Whilst the effect of each gene
the case of (a) discontinuous variation and ( b) continuous
alone is too small to make any significant impression on the
variation
phenotype, the almost infinite variety produced by the
combined effect of these genes (polygenes) has been shown
to form the genetic basis of continuous variation, which is
described further in section 23.8.2.
23.8.1 Discontinuous variation

23.8 Variation
There are certain characteristics within a
The term variation describes the difference in population which exhibit a limited form of variation.
characteristics shown by organisms belonging to the same Variation in this case produces individuals showing
natural population or species. It was the amazing diversity clear-cut differences with no intermediates between them,
of structure within any species that caught the attention of such as blood groups in Man, wing length in Drosophila,
Darwin and Wallace during their travels. The regularity melanic and light forms in Biston betularia, style length in
and predictability with which these differences in charac- Primula and sex in animals and plants. Characteristics
teristics were inherited formed the basis of Mendel’s showing discontinuous variation are usually controlled by
research. Whilst Darwin recognised that particular charac- one or two major genes which may have two or more allelic
teristics could be developed by selective breeding, as forms and their phenotypic expression is relatively un-
described in section 24.4.2, it was Mendel who explained affected by environmental conditions.
the mechanism by which selected characteristics were Since the phenotypic variation is restricted to certain
passed on from generation to generation. clear-cut characteristics, this form of variation is alterna-
Mendel described how hereditary factors determine the tively known as qualitative inheritance, as opposed to
genotype of an organism which in the course of develop- quantitative inheritance which is characteristic of conti-
ment becomes expressed in the structural, physiological nuous variation.
and biochemical characteristics of the phenotype. Whilst
23.8.2 Continuous variation
the phenotypic appearance of any characteristic is ulti-
mately determined by the genes controlling that character- Many characteristics in a population show a
istic, the extent to which certain characteristics develop complete gradation from one extreme to the other without
may be influenced by the environment. any break. This is illustrated most clearly by characteristics
A study of phenotypic differences in any large popula- such as mass, linear dimension, shape and colour of organs
tion shows that two forms of variation occur, discontinuous and organisms. The frequency distribution for a character-
and continuous. Studies of variation in a character involve istic exhibiting continuous variation is a normal distribu-
measuring the expression of that characteristic in a large tion curve (section A2.7.3). Most of the organisms in the
number of organisms within the population, such as height population fall in the middle of the range with approx-
in Man. The results are plotted as a histogram or a graph imately equal numbers showing the two extreme forms of
which reveals the frequency distribution of the variations of the characteristic. Characteristics exhibiting continuous
that characteristic within the population. Typical results variation are produced by the combined effects of many
obtained from such studies are shown in fig 23.29 and they genes (polygenes) and environmental factors. Individually
highlight the difference between the two forms of varia- each of these genes has little effect on the phenotype but
tion. their combined effect is significant.
851
‘
a-
23.8.3 Environmental influences introducing unlimited genetic variation into the popul
tion. These may be summarised as follows:
The ultimate factor determining a phenotypic (1) Reciprocal crossing-over of genes between chromatids
characteristic is the genotype. At the moment of fertilisa- of homologous chromosomes may occur during proph-
tion the genotype of the organism is determined, but the ase I of meiosis. This produces new linkage groups and
subsequent degree of expression allowed to this genetic so provides a major source of genetic recombination of
potential is influenced greatly by the action of environmen- alleles (section 23.3 and 22.3).
tal factors during the development of the organism. For (2) The orientation of the chromatids of homologous
example, Mendel’s tall variety of garden pea normally chromosomes (bivalents) on the equatorial spindle
attained a height of six feet. However, it would only do so if during metaphase I of meiosis determines the direction
provided with adequate light, water and soil conditions. A in which the pairs of chromatids move during anaphase
reduction in the supply of any of these factors (limiting I. This orientation of the chromatids is random. During
factors) would prevent the gene for height exerting its metaphase II the orientation of pairs of chromatids
full effect. It was the Danish geneticist Johanssen who once more is random and determines which chromo-
demonstrated the effect of the interaction of genotypic and somes migrate to opposite poles of the cell during
environmental factors on phenotype. In a series of anaphase II. These random orientations and the
experiments on the mass of dwarf bean seeds he selected subsequent independent assortment (segregation) of
the heaviest and lightest seeds from each generation of the chromosomes gives rise to a large calculable
self-pollinating dwarf bean plants and used these to number of different chromosome combinations in the
produce the next generation. After repeating these gametes (section 23.2.1)
experiments for several years he found only small dif- A third source of variation in sexual reproduction results
ferences in the mean mass of seeds from the same selected from the fact that the fusion of male and female gametes
line, that is heavy or light, but large differences in mean containing complementary sets of haploid chromosomes to
mass of seeds from different selected lines, that is heavy produce a diploid zygotic nucleus is completely random (at
and light. This suggested that both heredity and environ- least in theory). Thus, any male gamete is potentially
ment were influencing the phenotypic appearance of the capable of fusing with any female gamete.
characteristic. From these results it is possible to describe These sources of genetic variation account for the
continuous phenotypic variation as being ‘the cumulative routine ‘gene reshuffling’ which is the basis of continuous
effect of varying environmental factors acting on a variable variation. The environment acts on the range of pheno-
genotype’. The results also indicated that the extent to types produced and those best suited to it thrive. This leads
which a characteristic is inherited is determined primarily to changes in allele and genotypic frequencies as described
by the genotype. In the development of human characteris- in chapter 25. However, these sources of variation do not
tics such as personality, temperament and intelligence, generate the major changes in genotype which are
there is evidence to suggest that both nature (hereditary necessary in order to give rise to new species as described
factors) and nurture (environmental factors) interact to by evolutionary theory. These changes are produced by
varying degrees in different individuals to influence the mutations.
final appearance of the characteristic. It is these genetic and
environmental differences which act to produce pheno-
typic differences between individuals. There is no firm 23.9 Mutation
evidence, as yet, to suggest that one factor is universally A mutation is a change in the amount or the
more influential than the other, but the environment can
structure of the DNA of an organism. This produces a
never increase the extent of the phenotype beyond that
change in the genotype which may be inherited by cells
determined by the genotype. derived by mitosis or meiosis from the mutant cell. A
mutation may result in the change in appearance of a
23.8.4 Sources of variation characteristic in a population. Mutations occurring in
It will be appreciated that, as a result of the gamete cells are inherited, whereas those occurring in
interaction between discontinuous and continuous varia- somatic cells can only be inherited by daughter cells
tions and the environment, no two organisms will possess produced by mitosis. The latter are known as somatic
identical phenotypes. Replication of DNA is so nearly mutations.
perfect that there is little possibility of variation occurring A mutation resulting from a change in the amount or
in the genotypes of asexually reproducing organisms. arrangement of DNA is known as a chromosomal mutation
Any apparent variation between these organisms is or chromosomal aberration. Some forms of these affect the
therefore almost certainly the result of environmental chromosomes to such an extent that they may be seen
influences. In the case of sexually reproducing organ- under the microscope. Increasingly the term mutation is
isms there is ample opportunity for genetic variation to being used to describe a change in the structure of the DNA
arise. Two processes occurring during meiosis and the at a single locus and this is known as a gene mutation or
fusion of gametes during fertilisation provide the means of point mutation.
852
yp
The concept of mutation as the cause of the sudden
appearance of a new characteristic was first proposed by
parental cell (2n) pair of homologous
the Dutch botanist Hugo de Vries in 1901, following his
(wheren=1) chromosomes
work on inheritance in the evening primrose Oenothera
lamarckiana. Nine years later T. H. Morgan began a series
yore hh
of investigations into mutations in Drosophila and, with the
assistance of geneticists throughout the world, identified
over 500 mutations. gametes @ es
23.9.1 Mutation frequency and causes of

mutation fusion of
gametes a es
Mutations occur randomly and spontane-
Ously; that is to say that any gene can undergo mutation at normal normal
haploid haploid
any time. The rates at which mutations occur vary between gamete gamete
organisms.
As a result of the work of H. J. Miiller in the 1920s it was
observed that the frequency of mutation could be increased
above the spontaneous level by the effects of X-rays. Since trisomy monosomy
then it has been shown that the mutation rates can be Fig 23.30 Non-disjunction in gamete cell formation and the
significantly increased by the effects of high energy results of fusion of these abnormal gametes with normal
electromagnetic radiation such as ultra-violet light, X-rays haploid cells. The resulting cells may show a form of
and gamma rays. High-energy particles, such as a and polysomy where the chromosome number may be (2n + 1)
particles, trisomy, (2n + 2) tetrasomy, (2n + 3) pentasomy etc., or
neutrons and cosmic radiation, are also monosomy (2n — 1) depending upon the number of
mutagenic, that is cause mutations. A variety of chemical homologous chromosomes which fail to separate normally
substances, including mustard gas, caffeine, formalde-
hyde, colchicine, certain constituents of tobacco and an
increasing number of drugs, food preservatives and
pesticides, have been shown to be mutagenic.
Fusion of either of these gametes with a normal haploid
gamete produces a zygote with an odd number of
23.9.2 Chromosome mutations chromosomes.
Chromosomal mutations may be the result of Zygotes containing less than the diploid number of
changes in the number or structure of chromosomes. chromosomes usually fail to develop, but those with
Certain forms of chromosomal mutation may affect several polysomic chromosomes may develop. In most cases where
genes and have a more profound effect on the phenotype this occurs in animals it produces severe abnormalities.
than gene mutations. Changes in the number of chromo- One of the commonest forms of chromosomal mutation in
somes are usually the result of errors occurring during Man resulting from non-disjunction is a form of trisomy
meiosis but they can also occur during mitosis. These is called Down’s syndrome (2n = 47). The condition, which
changes may involve the loss or gain of single chromo- is named after the doctor who first described it in 1866, is
somes, a condition called aneuploidy, or the increase in due to the non-disjunction of the G21 chromosomes
entire haploid sets of chromosomes, a condition called (fig 23.31). The symptoms of Down’s syndrome include
euploidy (polyploidy). mental retardation, reduced resistance to disease, congeni-
tal heart abnormalities, a short stocky body and thick neck,
Aneuploidy and the characteristic folds of skin over the inner corner of
In this condition half the daughter cells produced have an the eye which produce a superficial facial similarity to
extra chromosome (n +1), (2n + 1) and so on, whilst the Mongolians. The syndrome is commonly, but rather
other half have a chromosome missing (n — 1), (2n — 1) cruelly, termed mongolism. Down’s syndrome and other
and so on. Aneuploidy can arise from the failure of a pair, related chromosomal abnormalities occur more frequently
or pairs, of homologous chromosomes to separate during in children born to older women. The exact reason for this
anaphase I of meiosis. If this occurs, both sets of is unknown but appears to be related to the age of the
chromosomes pass to the same pole of the cell and mother’s egg cells.
separation of the homologous chromosomes during ana- Non-disjunction of the male and female sex chromo-
phase II may lead to the formation of gamete cells contain- somes may also occur and produce aneuploidy affecting
ing either one or more chromosomes too many or too few secondary sexual characteristics, fertility and, in some
as shown in fig 23.30. This is known as non-disjunction. cases, intelligence (table 23.8).
853
pentaploid and so on. Polyploidy is much more common in
plants than in animals. For example, approximately half
the 300 000 known species of angiosperms are polyploid.
The relatively low occurrence in animals is explained by the
fact that the increased number of chromosomes in
1 s 3 ) 4 Oe polyploids makes normal gamete formation during meiosis
much more prone to error. Since most plants are capable of
WhRHBONG BB KSNK
propagating themselves vegetatively they are able to
reproduce despite being polyploid. Polyploidy is often
associated with advantageous features such as increased
size, hardiness and resistance to disease. This is called
hybrid vigour (section 25.4.2). Most of our domestic plants
ST
OA AR OA MK KES
are polyploids producing large fruits, storage organs,
flowers or leaves.
There are two forms of polyploidy, autopolyploidy and
1 16 17 18 allopolyploidy.
{ n ——_—_—_,—_—_——’
, it
Nee vce
Autopolyploidy. This condition may arise
naturally or artificially as a result of an increase in number
4% 48 AbA ha of chromosomes within the same species. For example, if
chromosomes undergo replication (during interphase) and
1 9 20 ~~ 2|—> ae xX
en ence ‘caesarean gsiemens?
the chromatids separate normally (during anaphase) but
F G the cytoplasm fails to cleave (during cytokinesis), a
tetraploid (47) cell with a large nucleus is produced. This
Fig 23.31 The chromosomes of a female suffering from
Down's syndrome. The non-disjunction of chromosomes G21 cell will undergo division and produce tetraploid cells. The
in one of the gametes has led to these chromosomes being amount of cytoplasm in these cells increases to preserve the
trisomic in this female. A photograph, such as the one above, nucleo-cytoplasmic volume ratio and leads to an increase in
shows a complete set of chromosomes for an individual the size of the whole plant or some part of it. Autopoly-
known as a karyotype ploidy can be induced by the use of a drug called colchicine
which is extracted from the corm of the autumn crocus
(Colchicum). Concentrations of the order of 0.01% inhibit
Table 23.8 Phenotypic abnormalities resulting from spindle formation by disrupting microtubules so that the
non-disjunction of the sex chromosomes. chromatids fail to separate during anaphase. Colchicine
and related drugs have been used in the breeding of certain
Frequency in varieties of economically important crops such as tobacco,
Condition/ Western tomatoes and sugarbeet. Autopolyploids can be as fertile
Genotype Symptoms populations
as diploids if they have an even number of chromosome
Klinefelter’s CO’, possessing some 2 0.02% sets.
syndrome (XXY) secondary sexual characteris- A modified form of polyploidy can occur in animals and
tics, sterile, testes very small, give rise to cells and tissues which are polyploid. This
little facial hair, breasts may
develop, usually low intelli- process is called endomitosis and involves chromosome
gence replication without cell division. The giant chromosomes in
Turner’s Q, lacking normal secondary 0.03% the salivary glands of Drosophila and tetraploid cells in the
syndrome (XO) sexual characteristics and very human liver are produced by endomitosis.
short, nipples close together
Allopolyploidy. This condition arises when
XXX Q, normal appearance, 0.12%
fertile, but mentally retarded the chromosome number inasterile hybrid becomes
XYY O’, tall, variable intelligence, 0.1%
doubled and produces fertile hybrids. F; hybrids produced
may possess psychopathic from different species are usually sterile since their
traits or tendency for petty chromosomes cannot form homologous pairs during
criminal acts meiosis. This is called hybrid sterility. However, if
multiples of the original haploid number of chromosomes,
for example 2(n, + n,),3(n, + n2) and so on (where n, and
Euploidy (polyploidy) n, are the haploid numbers of the parent species) occur, a
Gamete and somatic cells containing multiples of the new species is produced which is fertile with polyploids like
haploid number of chromosomes are called polyploids, and itself but infertile with both parental species.
the prefixes tri-, tetra-, and so on, indicate the extent of Most allopolyploid species have a diploid chromosome
polyploidy, for example 3n is triploid, 4n is tetraploid, 5n is number which is the sum of the diploid numbers of their
854
parental species; for example rice grass (Spartina angli
ca syndrome, where the diploid number is normal, the effects
(2n = 122) isa fertile allopolyploid hybrid produced from a are produced by the translocation of an extra G21
cross between Spartina maritima (stricta) (2n = 60)
and chromosome onto a larger chromosome, usually D15.
Spartina alterniflora (2n = 62). (The F, hybrid forme
d The simplest form of chromosomal mutation is deletion,
from the latter two species is sterile and is called Spart which involves the loss of a region of a chromosome, either
ina
townsendii (2n = 62)). Most allopolyploid plants have from the ends or internally. This results in a chromosome
different characteristics from either parental species, becoming deficient in certain genes (fig 23.33). Deletion
and
include many of Man’s most economically impor can affect one of a homologous pair of chromosomes, in
tant
plants. For example, the species of wheat used to make which case the alleles present on the non-deficient
bread, Triticum aestivum (2n = 42), has been selectively chromosome will be expressed even if recessive. If deletion
bred over a period of 5 000 years. By crossing a wild variety affects the same gene loci on both homologous chromo-
of wheat, einkorn wheat (2n = 14), with ‘wild grass’ somes the effect is usually lethal.
(2n = 14) a different species of wheat, emmer wheat In some cases a region of a chromosome becomes
(2n = 28), was produced. Emmer wheat has crossed with duplicated so that an additional set of genes exists for the
another species of wild grass (2n = 14) to produce Triticum region of duplication. The additional region of genes may
aestivum (2n = 42) which actually represents the hexap- be incorporated within the chromosome or at one end of
loid condition (6n) of the original einkorn wheat. Another
example of interspecific hybridisation involving crossing
A B C D EF
the radish and cabbage is described in section 25.9. —ee__________+—___2—_
@G normal chromosome
Allopolyploidy does not occur in animals because there

are fewer instances of cross-breeding between species. inversion translocation
Polyploidy does not add new genes to a gene pool (section
25.1.1) but gives rise to a new combination of genes. Da Coe EEG AN ise J Gp E Ipy Ve
—_e—_e—__2—_2—_+#——__o___o—__
Structural changes in chromosomes

Crossing-over during prophase I of meiosis involves the
| i
reciprocal transfer of genetic material between homolo- D Ae Beem anes
gous chromosomes. This changes the allele sequence of
normal
parental linkage groups and produces recombinants, but chromosome
no gene loci are lost. Similar effects to these are produced KL eV CDi
inverted + a TeA ae ae
by the structural changes in chromosomes known as ee FOG chromosome
inversions and translocations. In other forms of change, (a) showing loop (b)
such as deletions and duplications, the number of gene loci
Fig 23.32 Diagrammatic representation of inversion and
on chromosomes is changed, and this can have profound translocation and their effects on the positions of genes A-G.
effects on the phenotypes. Structural changes in chromo- (a) Looping in prophase due to inversion. (b) Part of the
somes resulting from inversion, deletion and duplication, chromosome carrying genes C, D and E has broken off and
and in some cases from translocation, may be observed become attached to the chromosome carrying genes K, L and
under the microscope when homologous chromosomes M
attempt to pair during prophase I of meiosis. Homologous
genes undergo synapsis (pairing) (section 22.3) and a loop A B C D E F G _ normal chromosome
or twist is formed in one of the homologous chromosomes
as a result of the structural change. Which chromosome
forms the loop and the arrangement of its genes depends deletion duplication
upon the type of structural change.
Inversion occurs when a region of a chromosome breaks PX \3) 1B, IR AG
go ix 3} (1D) ie (© ID) IE OF G
off and rotates through 180° before rejoining the chromo-
—e—__e—__e—__8—_8— —e—_e__e___e__—___@___e—_0—__@—__3—
some. No change in genotype occurs as a result of inversion

but phenotypic changes may be seen (fig 23.32). This
suggests that the order of gene loci on the chromosome is
important; a phenomenon known as the position effect. D
Translocation involves a region of a chromosome C D normal e E _ duplicated
breaking off and rejoining either the other end of the same AB EFG chromosome A Bee DiE EG chromosome
chromosome or another non-homologous chromosome 0 deleted —o_e__0_e__o_©_
“oe normal
(fig 23.32). The position effect may again be seen in the ees aa! chromosome A B C D E F G chromosome
phenotype. Reciprocal translocation between non- Fig 23.33 Diagrammatic representations of deletion and
homologous chromosomes can produce two new ORE OS duplication and their effects on the positions of genes A-G. In
gous pairs of chromosomes. In some cases of Down’s both cases looping can be seen
855
the chromosome, or become attached to another chromo- cause haemoglobin-S-containing red blood cells to distort
some (fig 23.33). and appear sickle-shaped. The physiological effect is to
lower the amount of oxygen which can be carried by these
cells, leading to acute anaemia. This not only causes
23.9.3 Gene mutations physical weakness, but may lead to heart and kidney failure
Sudden and spontaneous changes in pheno- and an early death in individuals homozygous for the
type, for which there are no conventional genetic explana- mutant allele. In the heterozygous condition individuals
tions or any microscopic evidence of chromosomal muta- show the sickle cell trait. The red blood cells appear normal
tion, can only be explained in terms of changes in gene and only about 40% of the haemoglobin is abnormal. This
structure. A gene mutation or point mutation (since it produces only mild anaemia and in parts of the world where
malaria is endemic, Africa and Asia in particular, it
applies to a particular gene locus) is the result of a change in
the nucleotide sequence of the DNA molecule in a particu- prevents carriers of the trait from contracting the disease.
lar region of the chromosome. Such a change in the base This is because the protozoan Plasmodium, which causes
sequence of the gene is transmitted to mRNA during trans- malaria, cannot live in red blood cells containing the
cription and may result in a change in the amino acid abnormal haemoglobin (section 25.1.5).
sequence of the polypeptide chain produced from it during
23.9.4 Implications of mutation
translation at the ribosomes (section 22.6).
There are a variety of forms of gene mutation involving The effects of chromosome and gene muta-
the addition, loss or rearrangement of bases in the gene. tions are very variable. In many cases the mutations are
These mutations take the form of the duplication, lethal and prevent development of the organism, for ex-
insertion, deletion, inversion or substitution of bases. In all ample in humans about 20% of pregnancies end in natural
cases they change the nucleotide sequence and result in the abortion before 12 weeks and of these about 50% exhibit a
formation of a modified polypeptide. For example, chromosome abnormality. Some forms of chromosome
deletion causes a frame shift and the implications of this are mutation may bring certain gene sequences together, and
described in section 22.5. that combined effect may produce a ‘beneficial’ character-
Gene mutations occurring during gamete formation are istic. Another significance of bringing certain genes closer
transmitted to all the cells of the offspring and may be together is that they are less likely to be separated by cross-
significant for the future of the species. Somatic gene ing-over and this is an advantage with beneficial genes.
mutations which arise in the organism are inherited only by Gene mutation may lead to several alleles occupying a
those cells derived from the mutant cells by mitosis. Whilst specific locus. This increases both the heterozygosity and
they may affect that organism, they are lost on the death of size of the gene pool of the population and leads to an
the organism. Somatic mutations are probably very increase in variation within the population. Gene re-
common and go unnoticed, but in some cases they may shuffling as a result of crossing-over, independent assort-
produce cells with an increased rate of growth and division. ment, random fertilisation and mutations, may increase the
These cells may give rise to a tumour which may be benign amount of continuous variation but the evolutionary
and not affect other tissues, or malignant, which lives implications of this are often short-lived since the changes
parasitically on healthy cells, a condition known as cancer. produced may be rapidly diluted. Certain gene mutations,
The effects of gene mutation are extremely variable. on the other hand, increase discontinuous variation and
Most minor gene mutations pass unnoticed in the pheno- this has the more profound effect on changes in the
type since they are recessive, but there are several cases population. Most gene mutations are recessive to the
where a change inasingle base in the genetic code can have ‘normal’ allele which has come to form genetic equilibrium
a profound effect on the phenotype. Sickle cell anaemia in with the rest of the genotype and the environment as a
Man is an example of base substitution mutation affecting a result of successfully withstanding selection over many
base in one of the genes involved in the production of generations. Being recessive the mutant alleles may remain
haemoglobin. The respiratory pigment haemoglobin of in the population for many generations until they come
adults is made up of four polypeptide chains (two a chains together in the homozygous condition and are expressed
and two £ chains) attached to the prosthetic group haem. phenotypically. Occasionally a dominant mutant allele
The polypeptide chains influence the oxygen-carrying may arise in which case it will appear immediately in the
capacity of the haemoglobin molecule. A change in the phenotype (section 25.5, Biston betularia).
base sequence of the triplet coding for one particular amino
acid out of the 146 in the B chains gives rise to the production The information provided in this chapter accounts for
of sickle cell haemoglobin (HbS). The amino acid se- the origins of variation within populations and the
quences for the normal and abnormal f chains differ in the mechanisms by which characteristics are inherited, but it
substitution of valine for glutamic acid at one point in the does not explain how the amazing diversity of living
abnormal polypeptide chains of haemoglobin S. Such a organisms described in chapters 2-4 may have arisen.
minor change causes haemoglobin S to crystallise at low Possible answers to this problem form the basis of the next
oxygen concentrations. The histological effect of this is to two chapters.
856
Chapter Twenty-four
aee ee OE ee
Evolution — history of life

The nature of life, its origin, the diversity of living design of a Creator. Science, contrary to popular belief,
organisms and the unifying structural and functional cannot contradict the idea of a divine origin for the early
relationships which underlie this diversity form a focal universe, nor do theological views necessarily dismiss the
point within the study of biology. view that during the origins of life, life acquired those
This chapter attempts to describe and discuss the many characteristics explained by the natural laws of science.
theories concerning the origin of life and the possible ways The major theories accounting for the origin of life on
in which species have originated. Traditionally the study of Earth are:
the history of life has been fraught with allegations of (1) life was created by a supernatural being at a particular
indoctrination. Indoctrination may be defined as a cons- time (special creation);
cious effort to inculcate an unshakeable commitment to a (2) life arose from non-living matter on numerous occa-
belief or doctrine. Such an approach is not only anti- sions (spontaneous generation);
scientific but also intellectually dishonest, and efforts to (3) life has no origin (steady-state);
avoid this have been made in this text. A brief outline of the (4) life arrived on this planet from elsewhere (cosmozoan);
main theories on the origin of life is presented in this (5) life arose according to chemical and physical laws
chapter so that students are aware that there is diversity of (biochemical evolution).
opinion as to the nature of this event. Much of the evidence
on which these theories are based is metaphysical, that is to 24.1.1 Special creation
say it is impossible to repeat the exact events of the origin of
life in any demonstrable way. This is true of both scientific This theory is upheld by most of the world’s
and theological accounts. However, one theory, evolution, major religions and civilisations and attributes the origin of
is increasingly being seen not as a single metaphysical life to a supernatural event at a particular time in the past.
theory but as a collection of individual scientific hypotheses Archbishop Ussher of Armagh calculated in 1650 that God
each of which is capable of being tested, as described in created the world in October 4004 BC, and finished with
section A2.1. Man at 9.00 a.m. on the morning of the 23rd. He achieved
In this chapter, and in chapter 25, scientific facts have this figure by adding up the ages of all the people in the
been selected to produce a coherent account of the biblical genealogies from Adam to Christ (the ‘begats’).
processes underlying the origins and diversity of forms of Whilst the arithmetic is sound, it places Adam as having
life. Because of the necessity to be selective this account lived at a time when archaeological evidence suggests that
lacks absolute objectivity; indeed, this is inevitably true of there was a well-established urban civilisation in the
any account, be it historical, scientific or metaphysical. Middle East.
However, by stressing the limitations and assumptions The traditional Judaeo-Christian account of creation,
associated with the evidence presented here, this account given in Genesis 1:1—26, has attracted, and continues to
may have a degree of objectivity and tentativeness which attract, controversy. Whilst all Christians would agree that
characterises good scientific writing. It must be stressed the Bible is God’s word to Man, there are differences of
interpretation concerning the length of the ‘day’ mentioned
that the evidence presented in this chapter, and the
conclusions drawn from it, represent current views. These
in Genesis. Some believe that the world and all species
were created in six days of 24 hours duration. They reject
are constantly under review and their validity is limited by
any other possible views and rely absolutely on inspiration,
the knowledge available to us at any given time.
meditation and divine revelation. Other Christians do not
regard the Bible as a scientific textbook and see the Genesis
24.1 Theories of the origin of life account as the theological revelation of the Creation of all
living things through the power of God, described in terms
understandable to men in all ages. For them the Creation
Theories concerned with the origin of the
account is concerned with answering the question ‘Why?’
Earth, and indeed the Universe, are diverse and uncertain.
rather than ‘How?’. Whilst science broadly relies on
Steady-state cosmologists maintain that the Universe never
observation and experiment to seek truth, theology draws
had an origin. Other hypotheses suggest that it may have
its insights from divine revelation and faith.
begun as a ball of neutrons, exploded in a ‘big bang’,
emerged from one of several black holes, or may be the
‘Faith is the substance of things hoped for, the
857
evidence of things not seen ... by faith ... we 24.1. What did Van Helmont omit from his7
understand that the universe was created by God’s word, experiment? : oo
so that what can be seen was made out of what cannot be
seen.’ (Hebrews 11:1, 3) In 1688 Francesco Redi, an Italian biologist and physician
Faith accepts things for which there is no evidence in the living in Florence, took a more rigorous approach to the
scientific sense. This means that logically there can be no problem of the origin of life and questioned the theory of
intellectual conflict between scientific and theological spontaneous generation. Redi observed that the little white
accounts of creation, since they are mutually exclusive worms seen on decaying flesh were fly larvae. By a series of
realms of thought. Scientific truth to the scientist is experiments he produced evidence to support the idea that
tentative, but theological truth to the theist is absolute. life can arise only from pre-existing life, the concept of
Since the process of special creation occurred only once biogenesis.
and therefore cannot be observed, this is sufficient to put ‘Belief would be vain without the confirmation of
the concept of special creation outside the framework of experiment, hence in the middle of July, I put a snake,
scientific investigation. Science concerns itself only with some fish, some eels of the Arno and aslice of milk-fed
observable phenomena and as such will never be able to veal in four large, wide-mouthed flasks; having well
prove or disprove special creation. closed and sealed them, I then filled the same number of
24.1.2 Spontaneous generation flasks in the same way, leaving only these open.’ (Redi)
Redi reported his results as follows.
This theory was prevalent in ancient Chinese,
‘It was not long before the meat and the fish, in these
Babylonian and Egyptian thought as an alternative to
second vessels (the unsealed ones), became wormy and
special creation, with which it coexisted. Aristotle (384—
the flies were seen entering and leaving at will; but in the
322 BC), often hailed as the founder of biology, believed
closed flasks I did not see a worm, though many days had
that life arose spontaneously. On the basis of his personal
passed since the dead fish had been put in them.’
observations he developed this belief further in relating all
organisms to a continuum, a scala natura (ladder of life).
24.2 What do you consider was Redi’s
‘For nature passes from lifeless objects to animals in such basic assumption?
unbroken sequence, interposing between them, beings
which live and yet are not animals, that scarcely any
These experiments, however, did not destroy the idea of
difference seems to exist between neighbouring groups,
spontaneous generation and, whilst the old theory took a
owing to their close proximity.’ (Aristotle)
setback, it continued to be the dominant theory within the
In stating this he reinforced the previous speculations of secular community.
Empedocles on organic evolution. Aristotle’s hypothesis of Whilst Redi’s experiments appeared to refute the
spontaneous generation assumed that certain ‘particles’ of spontaneous generation of flies, the pioneer work in
matter contained an ‘active principle’ which could produce microscopy by Anton van Leeuwenhoek appeared to
a living organism when conditions were suitable. He was reinforce the theory with regard to micro-organisms.
correct in assuming that the active principle was present in Whilst not entering the debate between biogenesis and
a fertilised egg, but incorrectly extrapolated this to the spontaneous generation, his observations with the micro-
belief that sunlight, mud and decaying meat also had the scope provided fuel for both theories and finally stimulated
active principle. other scientists to design experiments to settle the question
‘Such are the facts, everything comes into being not only of the origin of life by spontaneous generation.
from the mating of animals but from the decay of earth In 1765 Lazzaro Spallanzani boiled animal and vegetable
. .. And among plants the matter proceeds in the same broths for several hours and sealed them immediately. He
way, some develop from seed, others, as it were, by then removed them from the source of heat. After being set
spontaneous generation by natural forces; they arise aside for several days, none of them, on examination,
from decaying earth or from certain parts of plants.’ showed any signs of life. He concluded from this that the
(Aristotle) high temperature had destroyed all forms of living
With the spread of Christianity, the spontaneous genera- organisms in his vessel and without their presence no life
Suggs anther eson wySpa

tion theory fell from favour, except among those who could appear.
24 3.
believed in magic and devil-worship, although it remained
as a background idea for many more centuries.
Van Helmont (1577-1644), a much-acclaimed and
successful scientist, described an experiment which gave growth of organisms. —
rise to mice in three weeks. The raw materials for the
experiment were a dirty shirt, a dark cupboard and a In 1860 Louis Pasteur turned his attention to the problem
handful of wheat grains. The active principle in this process of the origins of life. By this stage he had demonstrated the
was thought to be human sweat. existence of bacteria and found the solutions to the
858
economic problems of the silk and wine industries
. He had Tube 6-— Plug with cotton wool and cover top with
also shown that bacteria were ubiquitous
and that aluminium foil. Autoclave at 15 Ib pressure
non-living matter could easily become contamin
ated by for 20 min.
living matter if all materials were not adequately
sterilised. Pair D
Tube 7— Surround straight glass tubing with cotton
. 24, 4 ‘What were Pasteur’ Ssbasic assump:
wool and plug boiling tube. Autoclave at
_tions about the orgs oflife?
15 lb pressure for 20 min.
Tube 8 — Surround S-shaped glass tubing with cotton
In a series of experiments based upon those of Spall
an- wool and plug boiling tube. Autoclave at 15 Ib
zani, Pasteur demonstrated the theory of biogenesis
and pressure for 20 min.
finally disproved the theory of spontaneous generation
. (4) Place all boiling tubes in an incubator at 32 °C.
The validation of biogenesis however raised another (5) Examine all boiling tubes every two days for ten days.
problem. Since it was now clear that a living organism was Record observations in the form ofa table.
required in order to produce another living organism, (6) After ten days remove a drop of broth from each tube
where did the first living Organism come from? The using sterile techniques and examine under the high
steady-state hypothesis has an answer for this but all the power of the microscope. Record observations in the
other theories imply a transition from non- living to living at form of a table.
some stage in the history of life. Was this a primeval (7) Draw conclusions from these observations.
spontaneous generation?
24.5 Clearly state the hypothesis which
Experiment 24.1: To investigate the origin of o would: account for the|pbentance of Teo
micro-organisms in terms of / organisms in the nutrient broth.
spontaneous generation and :
biogenesis _ 24.6 List the variables ea ‘hon
i: may influencetheaah ofee in
The objectives of this experiment are to repeat the ae broth. _
experiments of Spallanzani and Pasteur and carry out
further guided experiments which take into account the | 24. 7 “Which variablediffersbeween each :
of thetubes1and2,3 and 4,5 and 6, and 7 and 82
Criticisms of their experimental techniques, and to evalu-
ate objectively the hypotheses of spontaneous generation 24, 8 Which variable differs between
onthe .
and biogenesis. _ pairsoftubes A,B,C andD? -
Materials
24.9 Which tubes repeat
theexperiments
| of Saat and Pasteur? —
8 x 30 cm? boiling tubes straight glass tubing
(24. 10 Which tubes actas‘controls?
boiling tube rack 0.5 xX 6cm
120 cm? nutrient broth air-lock (S-shaped) glass 24. 1 3c you consider that the experi-_ 1
5 cotton wool plugs tubing 0.5 x 10cm ene oee out above meet all the criteria of a _
aluminium foil access to: autoclave, _ scientific. investigation, and what ores:a oS -
water bath incubator set at 32 °C - would youattach toyour conclusions? .
Method
24.1.3 Steady-state theory
(1) Autoclave eight boiling tubes.
(2) Place 15 cm? of nutrient broth into each of these boiling This theory asserts that the Earth had no
tubes, labelled 1-8. origin, has always been able to support life, has changed
(3) Set up tubes 1-8 as outlined below. remarkably little, if at all, and that species had no origin.
Pair A Estimates of the age of the Earth have varied greatly
Tube 1 — Leave unplugged, do not heat. from the 4004 BC calculation of Archbishop Ussher to the
Tube 2-— Plug with cotton wool and cover top with present-day values of 5 000 x 10° years based on radioac-
aluminium foil. Do not heat. tive decay rates. Improved scientific dating techniques
Pair B (Appendix 5) give increasing ages for the Earth, and
Tube 3—Leave unplugged. Heat in a boiling water extrapolation of this trend provides advocates of this
bath for 10 min. theory with the hypothesis that the Earth had no origin.
Tube 4— Plug with cotton wool and cover top with Whilst generally discrediting the value of geochronology in
aluminium foil. Heat in a boiling water-bath giving a precise age for the Earth, the steady-state theory
for 10 min. uses this as a basis for supposing that the Earth has always
Pair C existed. This theory proposes that species, too, never
Tube 5 — Leave unplugged. Autoclave at 15 Ib pres- originated, they have always existed and that in the history
sure for 20 min. of a species the only alternatives are for its numbers to
vary, or for it to become extinct.
859
The theory does not accept the palaeontological The atmosphere is believed to have been totally different
evidence that presence or absence ofa fossil indicates the in those days. The lighter gases hydrogen, helium,
origin or extinction of the species represented and quotes, nitrogen, oxygen and argon would have escaped because
as an example, the case of the coelacanth, Latimeria. Fossil the gravitational field of the partially condensed planet
evidence indicates that the coelacanths died out at the end would not contain them. However, simple compounds
of the Cretaceous period, 70 million years ago. The containing these elements (amongst others) would have
discovery of living specimens off the coast of Madagascar been retained, such as water, ammonia, carbon dioxide
has altered this view. The steady-state theory claims that and methane, and until the earth cooled below 100 °C all
it is only by studying living species and comparing them water would have existed as vapour. The atmosphere
with the fossil record that extinction can be assumed and would appear to have been a ‘reducing atmosphere’, as
then there is a high probability that this may be incorrect. indicated by the presence of metals in their reduced form
The palaeontological evidence presented in support of the (such as iron(II)) in the oldest rocks of the Earth. More
steady-state theory describes the fossil’s appearance in recent rocks contain metals in their oxidised form (for
ecological terms. For example, the sudden appearance of a example iron(III)). The lack of oxygen in the atmosphere
fossil in a particular stratum would be associated with an would probably be a necessity, since laboratory experi-
increase in population size or movement of the organism ments have shown, paradoxically, that it is far easier to
into an area which favoured fossilisation. Most evidence generate organic molecules (the basis of living organisms)
for this theory is based on discredited aspects of evolution- in a reducing atmosphere than in an oxygen-rich atmos-
ary theory such as the gaps in the fossil record and is best phere.
studied, in detail, alongside it. In 1923 Alexander Oparin suggested that the atmos-
phere of the primeval Earth was not as we know it today
24.1.4 Cosmozoan theory but fitted the description given above. On theoretical
This theory does not offer a mechanism to grounds he argued that organic compounds, probably
account for the origin of life but favours the idea that it had hydrocarbons, could have formed in the oceans from more
an extraterrestrial origin. It does not therefore, constitute a simple compounds, the energy for these synthesis reactions
theory of origin as such, but merely transposes the problem probably being supplied from the strong solar radiation
to elsewhere in the Universe. (mainly ultra-violet) which surrounded the Earth before
The theory states that life could have arisen once or the formation of the ozone layer, which now blocks much
several times, at various times and in various parts of the of it out. Oparin argued that if one considered the
Galaxy or Universe. Its alternative name is the theory of multitude of simple molecules present in the oceans, the
panspermia. Repeated sightings of UFOs, cave-drawings surface area of the Earth, the energy available and the time
of rocket-like objects and ‘spacemen’ and reports of scale, it was conceivable that oceans would gradually
encounters with aliens provide the background evidence accumulate organic molecules to produce the ‘primeval
for this theory. Russian and American space probes have. soup’, in which life could have arisen. This was not a new
provided evidence that the likelihood of finding life within idea, indeed Darwin himself expressed a similar thought in
our Solar System is remote but cannot comment on the a letter he wrote in 1871:
nature of life outside our Solar System. Research into ‘It is often said that all the conditions for the first
meteoritic and cometary materials has revealed the production of a living organism are now present, which
presence of many pre-vital organic molecules, such as could ever have been present. But if, (and oh what a big
cyanogen and hydrocyanic acid, which may have acted as if) we could conceive of some warm little pond, with all
‘seeds’ falling on a barren Earth. There are several claims sorts of ammonia and phosphoric salts, light, heat,
that objects bearing resemblances to primitive forms of life electricity, etc., present that a protein compound was
on Earth have been found in meteorites but they have yet chemically formed ready to undergo still more complex
to gain any real credibility in scientific circles. changes, at the present day, such matter would be
constantly devoured or absorbed, which could not have
24.1.5 Biochemical evolution been the case before living creatures were formed.’
It is generally agreed by astronomers, geol- In 1953 Stanley Miller, in a series of experiments,
ogists and biologists that the Earth is some 4.5—5.0 x10” simulated the proposed conditions on the primitive Earth.
years old. In his experimental high-energy chamber (fig 24.1) he
Many biologists believe that the original state of the successfully synthesised many substances of considerable
Earth bore little resemblance to its present-day form and biological importance, including amino acids, adenine and
had the following probable appearance: it was hot (about simple sugars such as ribose. More recently Orgel at the
4 000-8 000 °C) and as it cooled carbon and the less Salk Institute has succeeded in synthesising nucleotides six
volatile metals condensed and formed the Earth’s core; the units long (a simple nucleic acid molecule) in a similar
surface was probably barren and rugged as volcanic experiment.
activity, continuous earth movements and contraction on It has since been suggested that carbon dioxide was
cooling, folded and fractured the surface. present in relatively high concentrations in the primeval
860
gas chamber
methane (CH,)
(suggested contents of
ee ee ae (NH,)
early atmosphere)
water vapour (H,O)
connection to vacuum pump and
gas supply hydrogen (H,)
(to eliminate original atmosphere
and introduce gases)
high voltage electrical discharge
(supplying energy source)
direction
of flow
cooling jacket
(condensation of hot vapour)
liquid trap
Ji (collects products of chemical reactions)
boiling water
(provides water vapour and
ensures circulation)
HEAT
atmosphere. Recent experiments using Miller’s apparatus Fig 24.1 Stanley Miller's apparatus in which he synthesised
but containing mixtures of carbon dioxide and water and amino acids from gases under conditions thought to have been
only traces of other gases have produced similar results to present in the primeval atmosphere. The gases and vapours
those of Miller. Oparin’s theory has been widely accepted, were circulated under pressure and exposed to a high voltage
for one week. At the end of the period the liquid products in
but major problems remain in explaining the transition the trap were analysed by paper chromatography. A total of
from complex organic molecules to simple living organ- 15 amino acids was isolated including glycine, alanine and
isms. This is where the theory of a process of biochemical aspartic acid
evolution offers a broad scheme which is acceptable to the
majority of contemporary biologists. However, there is no of self-replication into the coacervate, and an internal
rearrangement of the lipid-coated coacervate, may have
agreement as to the precise mechanism by which it may
have occured. produced a primitive type of cell. Increase in size of
coacervates and their fragmentation possibly led to the
Oparin considered that protein molecules were crucial to
formation of identical coacervates which could absorb
the transformation from inanimate to animate. Because of
more of the medium and the cycle could continue. This
the zwitterionic nature of protein molecules they are able
to form colloidal hydrophilic complexes which attract, and possible sequence of events would have produced a
primitive self-replicating heterotrophic organism feeding
become surrounded by, envelopes of water molecules.
on an organic-rich primeval soup.
These bodies may separate from the body of the liquid in
Whilst this account of the origin of life is widely accepted
which they are suspended (aqueous phase) and form a type
by many scientists, the astronomer Sir Fred Hoyle has
of emulsion. Coalescence of these structures produces a
recently argued that the probability of random molecular
separation of colloids from their aqueous phase, a process
interactions giving rise to life as described above is
known as coacervation (coacervus, clump or heap). These
colloid-rich coacervates may have been able to exchange ‘as ridiculous and improbable as the proposition that a
substances with their environment and selectively concen- tornado blowing through a junk yard may assemble a
trate compounds within them, particularly crystalloids. Boeing 747’.
The colloid composition of a coacervate would depend on
the composition of the medium. The varying composition 24.2 The nature of the earliest
of the ‘soup’ in different areas would lead to variation in the organisms
chemical composition of coacervates, producing the raw
material for ‘biochemical natural selection’. Current evidence suggests that the first organ-
It is suggested that substances within the coacervates isms were heterotrophs as these were the only organisms
may have undergone further chemical reactions and, by capable of utilising the external supplies of available energy
absorbing metal ions into the coacervates, formed en- locked up within the complex organic molecules present in
zymes. The alignment of lipid molecules (complex hydro- the ‘soup’. The chemical reactions involved in synthesising
carbons) along the boundary between the coacervates and food substances appear to have been too complex to have
the external medium would have produced a primitive cell arisen within the earliest forms of life.
membrane which conferred stability on the coacervates. As more complex organic molecules arose through
Thus the incorporation of a pre-existing molecule capable ‘biochemical evolution’, it is assumed that some of these
861
(a) steady state (b) special creation (c) spontaneous
were able to harness solar radiation as an energy source and generation
use it to synthesise new cellular materials. Incorporation of Origin none at specified times at any time
these molecules into pre-existing cells may have enabled
the cells to synthesise new cellular materials without the
need for them to absorb organic molecules, hence
becoming autotrophic. Increasing numbers of hetero-
trophs would have reduced the available food resources in c¢ d A ty a Oo ac
Species a b
the primeval soup and this competition for resources would
hasten the appearance of autotrophs. (d) cosmozoan (e) biochemical evolution
The earliest photosynthetic organisms, whilst utilising at specified time(s)
Origin infinity/at specified time
solar radiation as their energy source, lacked the biochem-
ical pathways to produce oxygen. At a later stage, it is evolution
evolution
believed that oxygen-evolving photosynthetic organisms
developed, similar to existing blue-green algae (section
3.2), and this resulted in the gradual build-up of oxygen in Species a ib te «@ A DaeiCammrcl
the atmosphere. The increase in atmospheric oxygen and
its ionisation to form the ozone layer would reduce the (f) special creation (or) (g) cosmozoan
ultra-violet radiation striking the Earth. Whilst decreasing Origin at specified times infinity /at specified time
the rate of synthesis of new complex molecules, the (implies)
decreasing radiation would confer some stability on
otherwise successful forms of life. A study of the spontaneous generation
(at a specific time)
physiology of present-day organisms reveals a great
diversity in biochemical pathways associated with energy
biochemical evolution
capture and release, which may mirror many of nature’s
early experiments with living organisms.
evolution
Despite the simplified account given above, the problem
of the origin(s) of life remains. All that has been outlined is
Species ap DEN EC a
speculative and, despite tremendous advances in biochem-
istry, answers to the problem remain hypothetical. The Fig 24.2 Diagrammatic representation of various theories of
above account is a simplified amalgam of present-day the origin of life and the formation of species
hypotheses. No ‘ruling hypothesis’ has yet achieved the
status of an all-embracing theory (section A2.1). Details of
the transition from complex non-living materials to simple
living organisms remain a mystery. 24.4 The theory of evolution
The term ‘evolution’ has a special place in the

24.3 Summary of the ‘theories’ of the study of the history of life. It has become the unifying
origin of life concept which underpins the whole study of biology.
Evolution implies an overall gradual development which is
Many of these ‘theories’ and the way they both ordered and sequential. In terms of living organisms it
explain the existing diversity of species cover similar may be defined as ‘the development of complex organisms
ground but with varying emphases. Scientific theories may from pre-existing simpler organisms over the course of
be ultra-imaginative on the one hand and ultra-sceptical on time’.
the other. Theological considerations too, may fit into this The concept of evolution did not begin with Darwin and
framework depending upon one’s religious views. One of the publication of On the Origin of Species. Long before
the major areas of controversy, even before the days of Darwin, Man’s attempts to explain the obvious diversity of
Darwin was the relationship between scientific and theolo- living organisms which surround him had, paradoxically,
gical views on the history of life. led him to consider the basic structural and functional
Diagrams (a)-(e) in fig 24.2 represent straightforward similarities which exist between organisms. Evolutionary
descriptions of theories, hypotheses or beliefs on the hypotheses had been proposed to account for this and these
history of life, whereas (f) represents an attempt to ideas have themselves ‘evolved’ since the time of Darwin as
combine certain aspects of three theories (b), (c) and (d) knowledge has advanced.
into an alternative acceptable to many people. The The historical background to the development of the
practices of science and religion are not therefore necess- theory of evolution, as outlined in table 24.1, shows that
arily mutually exclusive as witnessed by the number of the concept of continuity or gradual development of more
scientists who hold religious beliefs. complex species from pre-existing simpler forms had
862
Table 24.1 The history of evolutionary thought.
Ancient Chinese
Confucius Life originated from a single source through a gradual unfolding and branching
Greek and Mediaeval period

Diogenes All things are differentiations of the same thing and are the same thing
Empedocles Air, earth, fire and water are the four roots of all things. Life arose by the action of
the forces of
attraction and repulsion on the four elements. Explained origin of Universe, plants, animals and Man
(Produced the germ of the idea of organic evolution.)
Democritus Living things arose by spontaneous generation from the slime of the Earth
Anaxogoras Organisms sprang from atmospheric germs
Thales All life came from water
(640-546 BC)
Anaximander Plants, then animals, and finally Man arose from the mud of the emerging Earth
Aristotle Proposed theory of continuous and gradual evolution from lifeless matter, based on his observations
(384-322 BC) of animals. Recognised a‘scala natura’ for animals
Dark Ages All theories based on those above or acceptance of special creation
(400-1400 AD)
Age of speculation
(1400-1790)
John Ray Developed concept of species
(1627-1705)
Carl Linnaeus Formalised ‘binomial classification’ system. Suggested genera were created separately and species
(1707-78) were variants of them
Buffon Suggested different types of animals had different origins at different times. Recognised influence
(1707-88) of external environment. Believed in acquired inheritance
James Hutton Theory of uniformitarianism. Gave age of Earth in millions of years
(1726-97)
Age of formulation
(1790-1900)
Erasmus Darwin Life arose from one ‘filament’ made by God. Did not accept the preformation of Man. The filament
(1731-1802) evolved by acquired characteristics
Jean-Baptiste Lamarck Inheritance of acquired characteristics. Environment acts on organisms. Phenotype changes are
(1744-1829) passed on. Concept of use and disuse of organs
Georges Cuvier Established palaeontological evidence. Fossils the results of ‘catastrophes’ by which new species
(1769-1832) arose
William Smith Opposed Cuvier’s theory of catastrophism on basis of continuity of similar species in related
(1769-1838) strata
Charles Lyell Demonstrated the progressive history of fossil evidence
(1797-1875)
Charles Darwin Influenced by Lyell and Malthus. Established a theory of evolution by means of natural selection
(1809-82)
Alfred Russel Wallace Similar theory to Darwin, but excepted Man from his theory
(1823-1913)
Hugo de Vries Recognised existence of mutations which were heritable as a basis for discontinuous variation and
(1848-1935) regarded species as arising by mutation
August Weismann Showed that the reproductive cells of animals are distinct and therefore unaffected by the
(1834-1914) influences acting on somatic tissues
Gregor Mendel Work on genetics (published 1865) only came to light after 1900. Laws of inheritance
(1822-84)
Developments in twentieth century (neo-Darwinism)

W. L. Johannsen Phenotype characteristics are determined by genotype and environmental factors
T. Henry Morgan Developed chromosome theory of heredity on basis of cytological evidence
H. J. Muller Genotype can be altered by X-rays: induced mutation
(1927)
R. A. Fisher No difference between change investigated by geneticists and change shown in the fossil record
(1930)
G. W. Beadle and Demonstrated the genetic basis of biochemical synthesis (following A. E. Garrod (1909) and
and E. L. Tatum (1941) J. B. S. Haldane (1935))
J. Lederberg and Demonstrated value of using bacteria in studying changes in genotype
A. D. Hershey (1951)
J. D. Watson and Proposed molecular structure of DNA and its mechanism of replication
F. H. C. Crick (1953)
F. Jacob and J. Monod Proposed a mechanism for regulation of gene activity
(1961)
863
occurred to several philosophers and natural historians unpaid post as naturalist on the survey ship H.M.S. Beagle,
before the formal declarations of evolutionary hypotheses which spent the next five years at sea charting the East
were advanced in the early nineteenth century. Coast of South America. The Beagle returned to Falmouth
in October 1836 via the coast of Chile, the Galapagos
24.4.1 Lamarckian evolution Islands, Tahiti, New Zealand, Tasmania and South Africa.
For most of this time Darwin was concerned with studying
The French biologist Lamarck proposed, in geology, but during a five-week stay on the Galapagos
1809, a hypothesis to account for the mechanism of Islands he was struck by the similarities shown by the flora
evolution based on two conditions: the use and disuse of and fauna of the islands and mainland. In particular he was
parts, and the inheritance of acquired characteristics. intrigued by the characteristic distributions of species of
Changes in the environment may lead to changed patterns tortoises and finches (section 24.7.2). He collected a great
of behaviour which can necessitate new or increased use (or deal of biological data concerned with variation between
disuse) of certain organs or structures. Extensive use would organisms which convinced him that species were not
lead to increased size and/or efficiency whilst disuse would immutable. On his return home his work on the selective
lead to degeneracy and atrophy. These traits acquired breeding of pigeons and other domestic animals gave hima
during the lifetime of the individual were believed to be clue to the concept of artificial selection, but he was unable
heritable and thus transmitted to offspring. to appreciate how this could operate in the wild. An earlier
According to Lamarckism, as the theory came to be
Essay on the Principles of Population by the Reverend
known, the long neck and legs of the modern giraffe were Thomas Malthus, published in 1778, had highlighted the
the result of generations of short-necked and short-legged consequences of the reproductive potential of Man.
ancestors feeding on leaves at progressively higher levels of Darwin applied this to other organisms and saw that
trees. The slightly longer necks and legs produced in each
despite this the numbers within populations remained
generation were passed on to the subsequent generation,
relatively constant. Having collated a vast amount of
until the size of the present-day giraffe was reached. The
information he began to realise that under the intense
webbed feet of aquatic birds and the shape of flat fish could competition of numbers in a population, any variations
be explained similarly. In aquatic birds the constant
which favoured survival in a particular environment would
spreading of the toe bones and the skin between them in
increase that individual’s ability to reproduce and leave
order to swim to find food and escape predators gave rise to
fertile offspring. Less favourable variations would be at a
their webbed feet. Likewise adaptations resulting from fish
disadvantage and organisms possessing them would there-
lying on their sides in shallow water were proposed to
fore have their chances of successful reproduction de-
explain the shape of flat fish. Whilst Lamarck’s theory
creased. These data provided Darwin with the framework
helped prepare the way for acceptance of the concept of to formulate, by 1839, a theory of evolution by natural
evolution, his views on the mechanism of change were selection, but he did not publish his findings at that time.
never widely accepted.
Indeed Darwin’s greatest contribution to science was not so
However, Lamarck’s emphasis on the role of the
much to show that evolution occurs but how it might occur.
environment in producing phenotypic changes in the
In the meantime, another naturalist, Alfred Russel
individual was correct. For example, body-building exer-
Wallace, who had travelled widely in South America,
cises will increase the size of muscles, but these acquired
Malaya and the Eastern Indian archipelago, and also read
traits, whilst affecting the phenotype, are non-genetic, and
Malthus, had come to the same conclusions as Darwin
having no influence on the genotype cannot be inherited.
regarding natural selection.
To demonstrate this, Weismann cut off the tails of mice
In 1858, Wallace wrote a 20-page essay outlining his
over many successive generations. According to Lamarck-
theory and sent it to Darwin. This stimulated and
ism, the enforced disuse of tails should have led to progeny
with smaller tails. This was not the case. Weismann encouraged Darwin and in July 1858, Darwin and Wallace
postulated that somatic (body) acquired characteristics presented papers on their ideas at a meeting of the Linnean
(resulting in phenotypic changes) did not directly affect the Society in London. Over a year later, in November 1859,
germ (gamete) cells which were the means by which Darwin published On the Origin of Species by Means of
characteristics are passed on to the next generation. His Natural Selection. All 1 250 printed copies were sold on the
theory of the ‘Continuity of the Germ-Plasm’ was a day of publication and it is said that this book has been
historical necessity before the inheritance of genetic second only to the Bible in its impact on Man’s thinking.
characteristics by sexual reproduction could be accepted.
24.5 Natural selection
24.4.2 Darwin, Wallace and the origin of
Darwin and Wallace proposed that natural
species by natural selection
selection is the mechanism by which new species arise from
Charles Darwin was born in 1809, the son of a pre-existing species. This hypothesis/theory is based on
wealthy doctor, and like many great people he had an three observations and two deductions which may be
undistinguished academic career. In 1831 he accepted an summarised as follows.
864
Observation 1: Individuals within a population have a great him of the importance of intraspecific variation. Likewise
reproductive potential. the adaptive significance of the interspecific variation seen
Observation 2: The numbers of individuals in a population in Galapagos finches (genus Geospiza) gave Darwin a clue
remain approximately constant. to his second deduction. Data collected by Wallace in the
Deduction 1: Many individuals fail to survive or reproduce. Malayan archipelago provided further evidence of varia-
There is a ‘struggle for existence’ within a population. tion between populations. Darwin and Wallace, however,
Observation 3: Variation exists within all populations. were unable to account for the sources of the variation.
Deduction 2: In the ‘struggle for existence’ those indi- This was not to be clarified until Mendel’s work on the
viduals showing variations best adapted to their environ- particulate nature of inheritance demonstrated how gene-
ment have a ‘reproductive advantage’ and produce more tic variation is conserved.
offspring than less well-adapted organisms. Deduction 2: Since all individuals within a population
Deduction 2 offers a hypothesis called natural selection exhibit variation and a ‘struggle for existence’ has been
which provides a mechanism accounting for evolution. clearly established, it follows that some individuals possess-
ing particular variations will be more suited to survive and
24.5.1 Evidence for natural selection reproduce. The key factor in determining survival is
adaptation to the environment. Any variation, however
Observation 1: It was Malthus who highlighted slight, be it physical, physiological or behavioural, which
the reproductive potential of Man and observed that gives One organism an advantage over another organism
human populations are able to increase exponentially will act as a selective advantage in the ‘struggle for
(section 12.6.3). The capacity for reproduction is basic to existence’. (The term ‘selective advantage’ is less emotive
all living organisms, and is a fundamental drive which than that coined by the social philosopher, Herbert
ensures continuance of the species. This applies to other Spencer, who described natural selection as ‘survival of the
organisms as shown in table 24.2. If every female gamete fittest’. The term ‘fit’ has irrelevant human connotations,
was fertilised and developed to maturity, the Earth would which have given an erroneous impression of natural
be totally overcrowded in a matter of days. selection.) Favourable variations will be inherited by the
next generation. Unfavourable variations are ‘selected out’
Table 24.2 Reproductive potential of selected species. or ‘selected against’, their presence conferring a selective
disadvantage on that organism. In this way natural
Crassostrea virginica American oyster 1.0 x 10° eggs selection leads to increased vigour within the species and,
per season in phylogenetic terms, ensures the survival of that species
Lycoperdon sp. giant puff ball TOE 10* spores (assuming environmental conditions remain constant).
Papaver rhoeas poppy capsule 6.0 x 10° seeds The whole of Darwin’s and Wallace’s hypothesis of natural
Carcinus maenas shore crab 4.0 x 10° eggs selection is summed up most succinctly in Darwin’s own
per season words:
‘As many more individuals of each species are born than
Observation 2: All population sizes are limited or checked can possibly survive, and as, consequently, there is a
by various environmental factors, such as food availability, frequently recurring struggle for existence, it follows that
space and light. Populations tend to increase in size until any being, if it vary however slightly in any manner
the environment supports no further increase and an profitable to itself, under the complex and sometimes
equilibrium is reached. The population fluctuates around varying conditions of life, will have a better chance of
this equilibrium, as discussed in section 12.6.3. Hence surviving and thus be naturally selected. From the strong
population sizes generally remain approximately constant principle of inheritance, any selected variety will tend to
over a period of time related to the length of the organism’s propagate its new and modified form.’ (Darwin, 1859)
life cycle. Many misconceptions have grown up around the theory
Deduction 1: The continuous competition between indi- of evolution as outlined by Darwin and they may be
viduals for environmental resources creates a ‘struggle for summarised as follows.
existence’. Whether this competition occurs within a (1) Darwin made no attempt to describe how life origin-
species (intraspecific competition) or between members of ated on the Earth: his concern was with how new
different species (interspecific competition) may be im- species might arise from pre-existing species.
material in affecting the size of the individual population (2) Natural selection is not simply a negative, destructive
(section 12.6.5), but it will still imply that certain organisms force, but can be a positive mechanism of innovation
will fail to survive or reproduce. within a population (section 25.5). The ‘struggle for
Observation 3: Darwin’s study of beetles whilst an existence’ described by Darwin was popularised by the
undergraduate at Cambridge, his subsequent journey in coining of unfortunate terms such as ‘survival of the
the Beagle and his knowledge gained through the selective fittest’ and ‘elimination of the unfit’ by the philosopher
breeding of certain characteristics in pigeons convinced Herbert Spencer and the press of the day.
865
(3) The misconception that Man was ‘descended from the Evidence for past evolution comes from many sources
apes’ by some process of linear progression was based on geology, such as fossils and stratigraphy.
over-sensationalised by the press and offended both the Evidence for a mechanism is found in the experimental and
religious and secular communities. The former saw this observational data of the natural selection of heritable
as an insult to their belief that Man was created in the characteristics, such as the selection of shell colour in
‘image of God’, whilst the latter were outraged by the Cepaea (section 25.5.1), and the mechanism of inherit-
apparent undermining of Man’s ‘superior’ position ance demonstrated by Mendelian genetics, as in Mendel’s
within the animal kingdom. work on peas. Finally, evidence for the action of these
(4) The apparent contradiction between the Genesis processes occurring today is provided by studies of present
six-day Creation account and that of a progressive populations, such as speciation in the herring gull (section
origin for species was exacerbated by the meeting of the 25.8.4), and the results of artificial selection and genetic
British Association for the Advancement of Science in engineering, as in the cultivation of wheat and the
June 1860. Bishop Samuel Wilberforce of Oxford production of monoclonal antibodies.
vehemently attacked the conclusions of Darwin as There are no laws of evolution, only well-corroborated
outlined in On the Origin of Species but not being a hypotheses which collectively add together to form a well-
biologist his address lacked accuracy. In concluding, he attested theory. The premature acceptance of current con-
turned to Professor Thomas Henry Huxley, a propo- cepts as dogmatic truths at Advanced level, as at any level
nent of Darwin’s theory, and asked whether he claimed of scientific inquiry, may stifle intellectual growth and the
his descent from a monkey through his grandfather or search for truth. The uncritical acceptance of evolutionary
grandmother. Huxley replied by expounding the more theory is a case in point. Some of the events presented as
important ideas of Darwin and correcting the mis- evidence for evolutionary theory can be reproduced under
conceptions of Bishop Wilberforce. In conclusion he laboratory conditions, but that neither implies nor confirms
implied that he would prefer to have a monkey for an that they did take place in the past; it merely indicates the
ancestor than ‘to be connected with a man who used possibility that these events occurred. Contemporary
great gifts to obscure the truth’. This unfortunate scientific debates on evolution are not concerned that
controversy has continued as the Genesis versus evolution takes place but that it takes place according to
Evolution debate. Professor R. J. Berry has summa- natural selection of randomly generated mutations.
rised the extremes of the debate as:
(a) those who are awed by scientists and believe that
the Bible has been disproved;
24.7 Evidence for the theory of evolution
(b) those who cling to the inspiration of Scripture and
Evidence associated with current views on the
their own interpretations of it, and shut their eyes
theory of evolution, is provided from many sources, the
to the fact that God’s-work can be studied by
main ones being palaeontology, geographical distribution,
scientific methods.
classification, plant and animal breeding, comparative
anatomy, adaptive radiation, comparative embryology and
comparative biochemistry.
24.6 Modern views on evolution
Much of the evidence presented in this chapter was un-
available to Darwin and Wallace at the time of publica-
The theory of evolution as proposed by
tion of their papers on the origin of species by natural
Darwin and Wallace has been extended and elaborated in
selection. Whilst great scientists are often characterised
the light of contemporary evidence from genetics, mole-
more by their powers of induction than those of deduction
cular biology, palaeontology, ecology and ethology and is
based on observation and experiment, Darwin and Wallace
known as neo-Darwinism (neo, new, adding to the notion
appear to have had a judicious blend of both. Darwin sums
of). This may be defined as the theory of organic evolution
up both of these approaches in his statement:
by the natural selection of genetically determined charac-
teristics. ‘In October 1838, that is, 15 months after I had begun my
The term ‘evolution’ may mean the result, or the systematic enquiry, I happened to read, for amusement,
process, of the above and different types of evidence Malthus on Population, and being well prepared to
support different aspects of this theory. In order to accept appreciate the struggle for existence which everywhere
neo-Darwinian evolutionary theory, as defined above, for goes on from long-continued observation of the habits of
the historical development of life it is necessary to: animals and plants, it at once struck me that under these
(1) establish the fact of change through time (past circumstances favourable variations would tend to be
evolution); preserved, and unfavourable ones to be destroyed. The
(2) demonstrate a mechanism which produces evolution- result of this would be the formation of new species.
ary change (natural selection of genes); Here, then, I had at last got a theory by which to work.’
(3) observe evolution happening today (‘evolution in The evidence presented here largely, but tentatively,
action’). supports the theory of evolution by natural selection as
866
Table 24.3 Types of fossils, their formation and examp
les
Fossil Fossilisation process

Examples
Entire organism Frozen into ice during glaciation
Woolly mammoths found in Siberian
permafrost
Encased in the hardened resin (amber) of coniferous trees Insect exoskeletons found in Oligocene rocks in
Baltic coast
Encased in tar ‘Mummies’ found in asphalt lakes of California
Trapped in acidic bogs: lack of bacterial and fungal activity ‘Mummies’ found in bogs and peat in
prevents total decomposition
Scandinavia
Hard skeletal Trapped by sedimentary sand and clay which form Bones, shells and teeth (very common in
materials sedimentary rocks, e.g. limestone, sandstone and silt British Isles)
Moulds and casts Hard materials trapped as above. Sediments harden to rock. Gastropods from Portland Stone, Jurassic.
The skeleton dissolves leaving its impression as a mould of Casts of giant horsetails (Calamites) of Carbon-
the organism. This can be infilled with fine materials iferous forests. Internal casts of mollusc shells
which harden to form a cast. Great detail is thus preserved showing muscle attachment points
Petrifaction Gradual replacement by water-carried mineral deposits, Silica replacements of the echinoderm
such as silica, pyrites, calcium carbonate or carbon. Slow Micraster
infilling as organism decomposes producing fine detail
Impressions Impressions of remains of organisms in fine-grained sedi- Feathers of Archaeopteryx in Upper Jurassic.
ments on which they died Jellyfish in Cambrian found in British
Columbia. Carboniferous leaf impressions
Imprints Footprints, trails, tracks and tunnels of various organisms Dinosaur footprints and tail scrapings indicate
made in mud are rapidly baked and filled in with sand and size and posture of organisms
covered by further sediments
Coprolites Faecal pellets prevented from decomposing, later com- Cenozoic mammalian remains
pressed in sedimentary rock. Often contain evidence of food
eaten, e.g. teeth and scales
ee
outlined in section 24.5. It draws on data obtained from classified into the same taxonomic groups (phyla and
many sources, and in ail cases is interpreted in terms that classes) as living species, but whether they represent the
assume the validity of the concept of evolution. Much of ancestors of present-day forms can only be debated, not
the evidence is also supportive of other sources of proved. /
evidence. Circular arguments and exceptions to the ‘ The oldest fossil-bearing rocks contain very few types of
evidence abound and alternative interpretations can be fossilised organisms and they all have a simple structure.
found, but the broad concept of evolution is backed up by a Younger rocks contain a greater variety of fossils with
wealth of scientific evidence which, at this level, is difficult increasingly complex structures. Throughout the fossil
to present in a form which is comprehensible yet not record many species which appear at an early stratigraphic
indoctrinatory. level disappear at a later level. This is interpreted in
evolutionary terms as indicating ihe times at which species
originated and became extinct.
24.7.1 Palaeontology
Geophysical evidence suggests that geographical regions
f “Palaeontology is the study of fossils. Fossils and climatic conditions have varied throughout the Earth’s
are any form of preserved remains thought to be derived history. Since organisms are adapted to particular environ-
from a living organism. They may include the following: ments, the constantly changing conditions may have
entire organisms, hard skeletal structures, moulds and favoured a mechanism for evolutionary change that
casts, petrifactions, impressions, imprints and coprolites / accounts for the progressive changes in the structure of
(fossilised faecal pellets) (table 24.3). f organisms as shown by the fossil record. Ecological
7 Fossil evidence alone is inadequate to uphold an considerations also fit in with the fossil evidence. For
NC
evolutionary theory, but it supports a theory of progressive example, plants appeared on land before animals, and
increase in complexity of organisms and denies the fixity of insects appeared before insect-pollinated flowers.
species//Fossils were well known before evolution was One of the major criticisms of using fossil evidence in
generally accepted. They were interpreted either as the support of an evolutionary theory is the lack of a continu-
remains of former creations or as artefacts inserted into the ous fossil record. Gaps in the fossil record (‘missing links’)
rocks by God. Most of the remains found so far can be are taken as strong evidence against a theory of descent by
867
s
Fig 24.3 Photograph of

trilobite fossil in
Cambrian rocks
modification. However, there are several explanations for proposed a process called ‘punctuated equilibria’ which
the incompleteness of the fossil record. These include the accounts for the sudden appearance of species. This
facts that: depends upon the fact that evolutionary rates are variable
(1) dead organisms decompose rapidly; and that certain new species arise rapidly with the
(2) dead organisms are eaten by scavengers; palaeontological consequence of an incomplete fossil
(3) soft-bodied organisms do not fossilise easily; record. These apparent ‘jumps’ in the evolutionary
(4) only a small fraction of living organisms will have died sequence have given rise to the term ‘saltatory evolution’
in conditions favourable for fossilisation; (saltare, to jump). Darwin himself considered this possibil-
(5) only a fraction of fossils have been unearthed. ity and stated as much in the Origin of Species:
Support for an evolutionary process increases as more and ‘I do not suppose that the process (speciation) . . . goes
more possible ‘missing links’ are discovered, either as on continuously; it is far more probable that each form
fossils, such as Seymouria (amphibia/reptile), Archaeop- remains for long periods unaltered, and then again
teryx (reptile/bird) and Cynognathus (reptile/‘mammal), or undergoes modification.’
as living organisms representing groups with close struc-
tural similarities, such as Peripatus (fig 24.18) and The fossil history of the horse
Latimeria. The history of the horse provides one of the best examples
Alternatively, there exists the possibility that new of phylogeny based on an almost complete fossil record
species appeared so suddenly that intermediate forms in found in North American sedimentary deposits from the
the lineage do not exist. Eldredge and Gould have early Eocene to the present.
868
The earliest recognisable perissodactyls (odd-toed, The history of the horse does not show a gradual
hoofed mammals) appeared about 54 x 10° years ago and transition regularly spaced in time and locality, and neither
present-day perissodactyls include horses, tapirs and is the fossil record totally complete. It would appear that
thinoceroses. The oldest recognisable horse-like fossils several offshoots occurred from the line represented in fig
belong to a genus called Hyracotherium which was widely 24.4, but they all became extinct. All modern horses
distributed throughout North America and Europe during appear to be descended from Pliohippus. The modern
the early Eocene. By the beginning of the Oligocene it was genus Equus arose in North America during the Pleis-
extinct everywhere except North America. It was a small tocene and migrated into Eurasia and Africa where it gave
animal, lightly built and adapted for running. The limbs rise to zebras and asses as well as the modern horse.
were short and slender and the feet elongated so that the Paradoxically, having survived in North America for
digits were almost vertical. There were four digits in the millions of years, the horse became extinct there several
forelimbs and three digits in the hindlimb. The incisors thousand years ago, at a time which coincided with the
were small and the molars had low crowns with rounded arrival of Man. Cave-paintings from other parts of the
cusps covered in enamel. world suggest that the earliest use for the horse was as a
The probable course of development of horses from source of food. The horse was absent from North America
Hyracotherium to Equus involved at least twelve genera until its reintroduction by the Spaniards almost 500 years
and several hundred species. The major trends seen in the ago.
development of the horse were concerned with locomotion
and feeding. They represent adaptations to changing
environmental conditions and may be summarised as 24.7.2 Geographical distribution
follows: All organisms are adapted to their environ-
(1) increase in size, ment to a greater or lesser extent. If the abiotic and biotic
(2) lengthening of limbs and feet, factors (section 12.2) within a habitat are capable of
(3) reduction of lateral digits, supporting a particular species in one geographical area,
(4) increase in length and thickness of the third digit, then one might assume that the same species would be
(5) straightening and stiffening of the back, found in a similar habitat in a similar geographical area, for
(6) better-developed sense organs, example lions in the savannah of Africa and the pampas of
(7) increase in size and complexity of the brain associated South America. This is not the case. Plant and animal
with point (6) above, species are discontinuously distributed throughout the
(8) increase in width of incisors, world. Ecological factors often account for this discon-
(9) replacement of premolars by molars, tinuous distribution, but evidence from the successful
(10) increase in tooth length, colonisation of habitats by plant and animal species
(11) increase in crown height of molars, introduced there by Man suggests that factors other than
(12) increased lateral support of teeth by cement, those of ecological adaptation are involved. Rabbits are
(13) increased surface area of cusps by exposure of enamel not endemic (naturally occurring) species in Australia, yet
ridges. their rapid increase in numbers following their introduction
A dominant genus from each geological epoch of the by Man indicates the suitability of the Australian habitat.
Cenozoic has been selected to show the progressive Similar examples of this principle are illustrated by the
development of the horse in fig 24.4. However there is no spread of domestic animals and plants by Man, such as
evidence that the forms illustrated are direct relatives of sheep, corn, potatoes and wheat. A rational explanation
each other. for the discontinuous distribution of organisms is based on
The significance of the fossil sequence shown in fig 24.4 is the concept of species originating in a given area and their
that it supports a theory of progressive change based on subsequent dispersal outwards from that point. The extent
homologous structures such as limbs and teeth. Each of the of the dispersal will depend upon the success of the
species shown in fig 24.4 represents a stage of development organisms, the efficiency of the dispersal mechanism and
which was successful for several million years (as judged by the existence of natural barriers such as oceans, mountain
the abundance of fossils) before becoming extinct. The ranges and deserts. Wind-blown spores and seeds and
extinction of a species did not, however, represent the flying animals would appear to have the best adaptations
disappearance of the family line. The fossil evidence for dispersal over land and sea.
reveals that another closely related species always In contrast to, and despite the general principle of
superseded its extinction. As all the species in the sequence organisms being naturally confined to certain parts of the
show structural and ecological similarities, this gives world, many related forms are found in widely separated
support to a theory of descent with modification. Other regions, for example the three remaining species of lungfish
fossils found in the same rock strata suggest changing (order Dipnoi) are found separately in tropical areas of
climatic conditions which, together with other evidence, South America (Lepidosiren), Africa (Protopterus) and
indicates that each species was adapted to prevailing Australia (Neoceratodus); camels and llamas (family
conditions. Camelidae) are distributed in North Africa, Asia and
869
‘
Body form (all heights are Bones of right forelimb Mode oflife, climate and structural modification
Epoch and age Genus
of oldest rocks ground to shoulder)
hock
carpals ce
splintbones Adapted to life in dry grasslands. Very efficient
at running. Metacarpals and metatarsals
cannon lengthened. Hoof formed from broadened
Pleistocene Equus bone phalanx 3 covering soft pad and all covered by
1X 10° yr claw. Teeth with large surface area. Enamel
pastern exposed where cement worn away. Premolars
3rd digit { replaced by molars. Grind food.
Increased reliance on speed. Digits 2 and 4 very

much reduced. Thickening of metacarpals and
Pliocene Pliohippus metatarsals (hindlimb) for support. Phalanx 3
7X 10° yr forms hoof. High-crowned teeth for eating
grass.
Very dry conditions: prairies. Speed more

important. Reduction of digits 2 and 4.
Miocene Merychippus Running on digit 3. Increase in length of
26 X 10° yr remaining metacarpal and metatarsal. Faller
with longer neck. Teeth longer with cement on
crown.
Dry conditions: forests and prairies. Speed

Oligocene Mesohippus
important to escape enemies. Only three digits
38 X 10° yr very obvious. Third digit much enlarged.
up to 0.6m
a
oS
Zim
Size of fox. Lived on soft ground near streams.
Eocene Hyracotherium Four digits in forelimb and three digits in
54 X 10° yr hindlimb increase surface area for support.
Low-crowned molar teeth adapted to browsing
about 0.4m on soft lush vegetation.
]
3
ee)
metacarpals
(numbered as shown)
Fig 24.4 Evolution of the modern horse
870
South America; and racoons are widely found in North and
Fig 24.5 (a) Relative positions of South America, Africa and
South America and a small area of south-east Asia. Fossil Australia during early stages of continental drift, indicating
evidence indicates that the distribution of these organisms proximity of areas where lungfish may have originated.
was not always as seen today and that in the past they were (b) Present distribution of species of lungfish
more widely distributed.
(@)
Whilst none of this evidence has any immediate
significance for evolutionary theory, it does point to the
fact that the distribution of land masses was not always as it
is today, as explained below. Africa
It used to be believed that the world had always been as it
now is and that the present continents and oceans had
never changed positions. Early geologists, such as Hutton
and Lyell (table 24.1), accounted for the existence of South America
sedimentary rocks in terms of the periodic rise and fall of
the sea. Later it was suggested that there were once two
large continental masses, one in the Northern Hemisphere
called Laurasia and one in the Southern Hemisphere called
Gondwanaland, linked by extensive land bridges across
which animals and plants could migrate and disperse.
Subsequent geological research has modified this idea and
()%S
favours the hypothesis of continental drift, based on the oo
concept of plate tectonics. The hypothesis of continental

drift was first proposed by Snider in 1858 but developed by
Taylor in America and Wegener in Germany in the late
1800s. Wegener proposed that, during Carboniferous
times, Laurasia and Gondwanaland formed one large land
mass called Pangaea (Greek, all earth) which floated on the
denser molten core of the Earth. It is now believed,
Lepidosiren Protopterus Neoceratodus
though, that continents have drifted apart as a result of
convection currents within the Earth spreading upwards
and outwards, dragging plates on which the continents In the case of the camels and llamas it is believed that
float. This hypothesis would account for the continuous they arose from a common ancestor which fossil evidence
movements of land masses and the present distribution of suggests had its origin in North America. During the
species such as those of the lungfishes (fig 24.5). Pleistocene this ancestor spread southwards into South
wage
Camelidae believed to have

originated here during
Pleistocene
i he world showingj istributi

distribution ] Camelidae,
of present members of the family j the came Is in North Africa and
ee peers South Gehohica: During Pleistocene times the Camelidae were distributed throughout North and South
dpionts and much of Asia and North Africa. This distribution is based on fossil evidence. Solid black lines indicate possible
migration routes. (Based on Matthews (1939) Climate of evolution, Vol. 1, 2nd ed., NY Acad. of Sci.)
871
s
America via the Isthmus of Panama, and northwards into organisms are believed to have been carried there by ocean
Asia before changes in sea-level separated it from North currents, whilst terrestrial organisms may have been
America (fig 24.6). Throughout this time it is thought that carried clinging to logs or floating masses of vegetation.
progressive changes within the Camelidae occurred, Birds, bats and flying insects would have fewer problems of
producing the two genera Camelus and Lama at the dispersal to these islands.
extremes of their Pleistocene migration. Forms intermedi- The Galapagos Islands are situated in the Pacific Ocean
ate between the present camels and llamas exist in the fossil on the equator almost 1 000 km west of Ecuador and form
record throughout North America, Asia and North Africa. an archipelago described further in section 25.8.3. When
The fossil record indicates that other animals of the Darwin visited the islands in 1835, he noticed the similarity
Camelidae in other parts of the world became extinct at the of the species found there to those on the nearest mainland,
close of the last Ice Age. a fact he had also observed on the Cape Verde Islands off
Another example of discontinuous distribution as a the coast of West Africa. However, the plant and animal
result of geographical isolation is provided by the mono- species on oceanic islands were noticeably larger in most
tremes and marsupials of Australasia. Australasia is cases. This may be accounted for by the lack of competition
believed to have broken away from the other land masses from larger, and more dominant, advanced species which
during the late Jurassic, just after the appearance of were absent from the islands, but which co-habited with
primitive mammals. The mammals are divided into three smaller related species on the mainland. For example, the
orders: Monotremata, Marsupialia and Eutheria. In giant tortoise (Geochelone elephantopus), nearly 2 m long
Australasia only the monotremes and marsupials de- and weighing 260 kg, feeding on the plentiful vegetation
veloped. Here they coexisted and underwent adaptive found on the islands presumably attained this size due to
radiation to produce the characteristic Australasian fauna the absence of competition from various mammalian
represented by the monotremes Tachyglossus and Zaglos- species which existed on the mainland. Darwin noticed too
sus (the spiny anteaters) and Ornithorhynchus (the that iguana lizards on the Galapagos Islands were abundant
duck-billed platypus), and 45 genera of marsupials. and again much larger than related mainland species.
Elsewhere in the world the more advanced eutherian Lizards are terrestrial reptiles, but on the Galapagos
(placental) mammals also developed. As they spread out Islands, where two species were found, one was aquatic.
over the continents it is believed that they ousted the more The aquatic form, Amblyrhyncus cristatus, fed on marine
primitive monotremes and marsupials from their ecological algae and showed adaptations for locomotion in water such
niches, except where geographical barriers disrupted their as a laterally-flattened tail and well-developed webs of skin
dispersal, as into Australasia. between the toes of all four limbs (fig 24.7). Competition
These points may be summarised as: for food, space and a mate within the terrestrial form is
(1) species originated in a particular area; thought to have exerted a selection pressure on the lizards
(2) species dispersed outwards from that area; and favoured those showing variations with aquatic
(3) dispersal could only occur for most species where land adaptations. This mechanism of environmental factors
masses were close enough together to permit dispersal; operating on a variable genotype is called natural selection
(4) the absence of more advanced organisms from a region and is described above. It could have been the process
usually indicates the prior separation of that region
from the area of origin of those organisms.
Whilst none of the evidence presented above indicates the
mechanism by which species are thought to have origin-
ated, it does suggest that various groups have originated at
various times and in various regions. Fossil evidence
reveals the ways in which these organisms have undergone
gradual modification, but again gives no indication of the
possible mechanism.
Evidence for a possible mechanism of the origin of
species by natural selection is supplied by the distribution
of plants and animals on oceanic islands. Both Wallace and
Darwin were struck by the amazing diversity of species
found on such islands, such as the Hawaiian and the
Galapagos groups. Geological evidence indicates that
these islands were formed by oceanic volcanic activity
which thrust them up above sea level, so that they have
never had any direct geographical links with any land mass.
Plant species must have arrived on the islands by wind
dispersal as spores and seeds, or water dispersal as floating
seeds and masses of vegetation. Aquatic and semi-aquatic Giant aquatic lizard of the Galapagos Islands
872
(a) Types offinch ; Beak shape Food source Habitat Number of species
large ground finch (ancestral) typical main land type: short crushing seed coastal
and straight
ground finches
ta
various, but short and seeds/insects coast/lowlands 3
straight as above
SS
cactus ground finches long slightly curved, split- nectar of prickly-pear cactus lowland 2
tongue
= “es
insectivorous tree finches parrot-like seeds/insects forest 3
vegetarian tree finch curved, parrot-like fruit/buds/soft fruit forest 1

a
warbler finch slender insects in flight forest 1

-
woodpecker finches large, straight, (uses cactus larvae insect forest 2

spine or stick to poke insects
SS out of holes in wood)
a —s
which gradually gave rise to the aquatic species. It was, Fig 24.8 (a) Adaptive radiation of Darwin's finches. (After
however, the diversity of adaptive structure shown by the Lack) (b) A male cactus finch (Geospiza scandens)
13 species of finches found within the archipelago which (b) gaygme
had the greatest influence on Darwin’s thinking on the
mechanism of the origin of species. Only one type of finch
existed on the mainland of Ecuador and its beak was
adapted to crushing seeds. On the Galapagos Islands, six
major beak types were found, each adapted toa particular
method of feeding. The various species, their feeding
methods and numbers of species are summarised in
fig 24.8.
Darwin postulated that a group of finches from the
mainland colonised the islands. Here they flourished, and
the inevitable competition produced by increase in num-
bers, and the availability of vacant ecological niches,
favoured occupation of niches by those organisms showing
the appropriate adaptive variations. Differences between
species relate to small differences in body size, feather
colour and beak shape. Several species of finch are found
on all the bigger islands. The ground and warbler finches,
thought to be the most primitive types, are found on most
islands. The tree and vegetarian/tree finches are missing
from the outlying islands, and the woodpecker finches are
confined to the central group of islands. The actual species
distribution is interesting and has been explained by Lack
on the basis of adaptiv2 radiation and geographical
isolation. For example, on the central islands there are
873
many species of several different types of finch, such as
ground, tree warbler and woodpecker, rather than several
species of the same type. Even where several species of
only one type of finch are present, as on the outlying
islands, each species differs in its ecological requirements.
This fits in with the Gaussian exclusion principle (section
12.6) which states that two or more closely related species
will not occupy the same area unless they differ in their
ecological requirements.
24.7.3 Classification
The system of classification described in
Appendix 3 was proposed by Linnaeus before the time of
Darwin and Wallace, but has implications for the origin of
species and evolutionary theory/Whilst it is possible to
conceive that all species, both living and extinct, were
created separately at a specific time or had no origin, the
structural similarities between organisms, which forms the
basis of a natural system of phylogenetic classification,
suggest the existence of an evolutionary process. These
similarities and differences between organisms may be
explained as the result of progressive adaptation by
organisms within each taxonomic group to Ee
environmental conditions over a period of time./
Z Numerical taxonomists, working mainly from compara-
tive phenotypic characters have found it possible to
construct a phenetic classification system (Appendix 3)
which is consistent, to the extent of present knowledge,
with the concept of evolution. These systems of classifica-
tion are capable of standing in their own right as a basis for
biological organisation, but they also strongly suggest that
an evolutionary process has occurred,
24.7.4 Plant and animal breeding

One of the earliest features of Man’s civilisa-
tion was his cultivation of plants and domestic animals from
ancestral wild stocks. By selecting those members of a
species which showed a favourable variation, such as
increased size or improved flavour, and artificially breeding
them by selective mating, selective propagation or pollina- Fig 24.9 The result of selective breeding. The wild pig
tion, the desired characteristics were perpetuated. Con- (a) is native to Europe, Asia and Africa but has been
tinued selective breeding by Man has produced the selectively bred to produce a variety of breeds, of which the
varieties of domestic animals and plants of agricultural English Large White pig (b) with its high quality of meat yield,
importance seen today. It is known from archaeological is an example
remains that early Man was proficient in rearing cattle, pigs
and fowl, and cultivating cereal crops and certain veg- economically important varieties of animals and plants
etables. Until the revelation of Mendel’s work on genetics which have been studied most by plant and animal breeders
the theoretical basis of inheritance and breeding was not (fig 24.9). Some specific examples of phenotypic character-
clear, but this had not limited Man’s practical endeavours. istics which have been artificially selected are shown in
In terms of genetics, Man is preserving those genes which table 24.4. A recently developed form of artificial selection
are considered desirable and eliminating those which are is the selection for resistance to antibiotics, pesticides and
undesirable for his purposes. This selection exploits herbicides shown respectively by pathogens, pests and
naturally occurring gene variation, together with any weeds. A vicious circle is produced as new strains of
fortuitous mutations which occur from time to time. organisms become immune to the ever-increasing number
Whilst varieties of dogs, cats, birds, fish and flowers have of chemical substances produced to contain and control
been produced for sporting or decorative purposes, it is them.
874
Table 24.4 Selected phenotypic characteristics and
24.7.5 Comparative anatomy
examples of them.
ii
ee
Comparative study of the anatomy of groups
Phenotypic characteristic Example of animals or plants (morphology) reveals that certain
structural features are basically similar. For example, the
Hardiness Sweetcorn grown in England basic structure of all flowers consists of sepals, petals,
Size Potato, cabbage
Increased yield stamens, stigma, style and ovary; yet the size, colour,
Milk, eggs, wool, fruit
Earlier maturity Cereal crops (two per season) number of parts and specific structure are different for each
Lengthened season Strawberries individual species. Similarly, the limb-bone pattern of all
Taste/eating quality Apples, seedless grapes tetrapods from amphibia to mammals has the same
Harvesting ease Peas structural plan: it is called the pentadactyl limb (fig 17.11).
Length of storage Beans/peas for freezing
Increased ecological
This basic structure has been modified in several ways as
Protein from plants, e.g. soyabean
efficiency illustrated in fig 24.10. In each case, the particular structure
Resistance to disease Rust and mildew (fungi)-resistant is adapted to a certain method of locomotion ina particular
wheat environment.
——— SS Organs having a similar basic structure, a similar
topographic relationship as structures in other species, the
Since characteristics can be ‘produced’ by Man’s ability same histological appearance and a similar embryonic
to selectively breed, as in the case of breeds of dogs or development are said to be homologous, a term introduced
pigeons, Darwin used this as evidence for a mechanism by in 1843 by Richard Owen.
which species might arise ‘naturally’. In the latter case the Homologous structures showing adaptations to different
environment rather than Man was believed to act as the environmental conditions and modes of life are examples
agent of selection. Artificially selected forms probably of adaptive radiation. The ecological significance of these
would not have arisen in the ‘wild’; in most cases they are processes is considered in section 24.7.6. The specific
unable to compete successfully with closely related non- functions that these structures carry out may vary in
domesticated forms. different organisms. These differences reflect the particu-
} carpals q
Horse
' metacarpals (running)
(fused with)
radius humerus radius
i ‘knee’ carpals
metacarpal rudimentary metacarpals
phalanges (1 cannon bone) (2 splint bones)
fetlock
ulna radius pastern
hoot A
phalange
ulna 3
-} carpals enlarged displaced humerus

metacarpals carpals carpal :
P 1 radius
metacarpals
ulna
5 f) phalanges 1 hal
phalange s Ft
7 Man phalanges
Seal (swimming) (manipulating)
5439 Mole (digging) metacarpals
Fig 24.10 Adaptations of the pentadactyl limb shown by mammals
875
becomes
x
lar ways the organisms are adapted to their environments Pericarp. The ovary wall in flowering plants
a variety
and modes of life. Other examples of homology are given modified, following fertilisation of the ovules, in
).
below. of ways to aid seed dispersal (figs 20.28 and 24.12
has
Branchial arches/Ear ossicles. Certain bones of the jaw in Whilst homology does not prove that evolution
of
fish can be traced through other vertebrates, where they occurred, the existence of homology within a group
nt from a
are involved in jaw suspension, to mammals where they organisms is interpreted as evidence of their desce
rela-
appear as the ear ossicles, the malleus, incus and stapes common ancestor and indicates close phylogenetic
(fig 24.11). tionships.
Halteres. The hind pair of wings typical of most insects have Linnaeus used homology as the basis for his system of
been modified in the Diptera into little rods, the halteres, classification. The more exclusive the shared homologies,
which serve as gyroscopic organs helping to maintain the closer two organisms are related and hence the lower
balance in flight. the rank of the taxonomic group in which they are placed.
dried stigma
hyomandibular : — pericarp wing seed
cranium attachment for jaws to
cranium ‘pepper-poU’
: uadrate seed pericarp
upper jaw is) q bones of
articular jaw
j di
(wind dispers
Poppy (wind al)l
lower jaw Sycamore (wind dispersal)
(a) Fish
stapes-transmits
sound from
tympanic
: membrane to
cranium and inner ear
upper jaw drying
quadrate jaw pod >
fused together
:
articular { articulation << ___fettilised
ovary wall
seeds
lower jaw
Gorse (mechanical Coconut (water
(b) Amphibia dispersal) dispersal)
malleus
incus
stapes succulent
m
auditory : mesocarp
canal 1 stony hooks
ec) oval window endocarp
seed
round window
Cherry Goosegrass
(c) Mammal Eustachian tube (animal dispersal) (animal dispersal)
Fig 24.11 Relative positions and functions of bones of the Fig 24.12 Variation in pericarp structure for different
mammalian ear ossicles as seen in fish and amphibia methods of seed dispersal
876
For example, butterflies and moths belong to the
same
order (Lepidoptera) whereas wasps and bees belon
g to
another order (Hymenoptera). a ant
Certain homologous structures in some species have no
apparent function and are described as vestigial organs
.
The appendix of Man, although not concerned with
digestion, is homologous with the functional appendix of mx.
herbivorous mammals. Likewise, certain apparently non-

functional bones in snakes and whales are thought to be sucking
e.g. butterfly, labrum reduced;
homologous with the hip bones and hindlimbs of quad- mandibles lost; maxillae long
wy
ruped vertebrates. The vertebrae of the human coccyx are forming sucking tube; labium
thought to represent vestigial structures of the tail reduced
possessed by our ancestors and embryos. It would be very

difficult to explain the occurrence of vestigial organs
without reference to some process of evolution.
24.7.6 Adaptive radiation licking and biting

e.g. honey bee, labium
long to lap up nectar;
When a group of organisms share a homolo- mandibles chew pollen
gous structure which is differentiated to perform a variety and mould wax
of different functions, it illustrates a principle known as
adaptive radiation. Adaptive radiation may be demons-
trated within all taxonomic groups higher than the species.
For example, all organisms belonging toa particular class
share a number of diagnostic characteristics. Additionally,
, variations between different species within the class enable
them to have modes of life adapted to particular habitats.
For instance, the mouthparts of insects consist of the same primitive state:
basic structures: a labrum (upper lip), a pair of mandibles, biting and chewing
e.g. grasshopper, strong
a hypopharynx (floor of mouth), a pair of maxillae and a mandibles; maxillae for
labium (fused second pair of mandibles, lower lip). Insects manipulating food
are able to exploit a variety of food materials, as shown in

fig 24.13, because some of the above structures are
enlarged and modified, others reduced and lost. This
produces a variety of feeding structures. /
The relatively high degree of adaptive radiation shown
by insects reflects the adaptability and utility of the basic
features of the group. It is this ‘evolutionary plasticity’
which has permitted them to occupy such a range of
ecological niches. This is one of a group of related criteria
used to describe the biological success of a taxonomic
group, such as a phylum or class. Other criteria include the
evolutionary age of the group and the number of species
within the group. piercing and sucking
e.g. female mosquito,
The presence of a structure or physiological process in labrum and maxillae form
an ancestral organism, which has become greatly modified tube; mandibles form
piercing stylets; labium
in more advanced, apparently related organisms, may be grooved to hold other
interpreted as indicating a process of descent by modifica- parts
tion which is the basis of evolutionary theory as defined in
section 24.4.2. The significance of adaptive radiation is that
Fig 24.13 Adaptive radiation of insect mouthparts;

a, antennae; c, compound eye; Ib, labium; Ir, labrum;
md, mandibles; mx, maxillae
877
flying phalanger
<—_—_———_ primitive marsupials
red kangaroo tasmanian wolf
banded anteater
See x
tasmanian devil
wombat
marsupial mole
Fig 24.14 Adaptive radiation of marsupials in Australia (from a variety of sources)
878
it suggests the existence of divergent evolution based
on 24.7.7 Comparative embryology
modification of homologous structures.
Similar structures, physiological processes or modes of A study of the embryonic development of the
life in organisms apparently bearing no close phylogenetic vertebrate groups by Von Baer (1792-1867) revealed
links but showing adaptations to perform the same striking structural similarities occurring in all the groups,
functions are described as analogous. Examples include the particularly during cleavage, gastrulation and the early
eyes of vertebrates and cephalopod molluscs, the wings of stages of differentiation (section 21.5). Haeckel (1834
insects and bats, the jointed legs of insects and higher —1919) suggested that this had an evolutionary significance.
vertebrates, the presence of thorns on plant stems and He formulated the principle that ‘ontogeny recapitulates
spines on animals, and the existence of vertebrate phylogeny’, that is the developmental stages through which
neuroendocrines, such as acetylcholine, 5-hydroxy- an organism passes repeats the evolutionary history of the
tryptamine and histamine, in nettle stings. Analogous group to which it belongs. Whilst this principle over-
structures only bear superficial similarities. For example, generalises the situation, it is attractive and has a degree of
the wings of insects are supported by toughened veins demonstrable validity. Examination alone of the embryos
composed of cuticle, whereas both bats and birds have and foetal stages of all the vertebrate groups reveals that it
hollow bones for support. Likewise the embryological is impossible to identify the group to which they belong.
development of the cephalopod and vertebrate eyes is Fig 24.15 shows that it is only in the later stages of
different. The former produces an erect retina with development that they begin to assume some similarity to
photoreceptors facing the incoming light, whereas the their adult form. At comparable stages the vertebrate
latter has an inverted retina with photoreceptors separated embryos all possess the following.
from incoming light by their connecting neurones (fig (1) External branchial grooves (visceral clefts) in the
16.33). Thus the vertebrate eye has a blind spot, which is pharyngeal region andaseries of internal paired gill
absent in cephalopods. pouches. These join up in fishes to form the gill slits
The existence of analogous structures suggests the involved in gaseous exchange. In the other vertebrate
occurrence of convergent evolution. Convergent evolution groups the only perforation that develops in adults
may be explained in terms of the environment, acting becomes the Eustachian tube and auditory canal
through the agency of natural selection, favouring those involved in hearing.
variations which confer increased survival and reproduc- (2) Segmental myotomes are evident in the tail-like
tive potential on those organisms possessing them. structure which is retained in certain species only.
The significance of divergent evolution, suggesting an (3) There is a single circulation which includes a two-
evolutionary process, and convergent evolution, suggest- chambered heart showing no separation into right and
ing an evolutionary mechanism, is highlighted by the left halves, a situation retained completely only in
fishes.
parallel evolution of marsupial and placental mammals.
As development proceeds in the vertebrate embryo,
Both groups are thought to have undergone convergent
changes occur which produce the characteristics of fish,
evolution and come to occupy identical ecological niches in
amphibian, reptile, bird or mammal depending upon the
different parts of the world (fig 24.14 and table 24.5).
embryo’s parentage. The interpretation placed on these
Table 24.5 Examples of parallel evolution shown by
marsupial and placental mammals.
Marsupial mammals Placental mammals

(Australasia) (elsewhere)
Marsupial mole Mole

Marsupial mouse Mouse
Banded anteater Anteater
Wombat Prairie dog
Kangaroo Antelope
Bandicoot Rabbit
Flying phalanger Flying squirrel
Koala Sloth
Tasmanian wolf Hyena
Fig 24.15 Stages in embryological development as shown

by examples from three vertebrate classes tortoise rabbit
879
observations is that these embryos, and hence the groups to show spiral cleavage and their blastopore becomes the
which they belong, had a common ancestor. There seems mouth of the adult. This pattern of development is seen in
little point in an organism having developmental structures the annelids, molluscs and arthropods. Deuterostomes
which are apparently non-functional in the adult unless show radial cleavage and their blastopore becomes the
they are the remaining stages of ancestral structures. anus of the adult. The echinoderms and chordates show
However the principle of recapitulation cannot be accepted this pattern of development. These differences are shown
entirely since no living organisms can show all the features in fig 24.16. It is evidence such as this which has helped
of their proposed evolutionary ancestors. What appears to clarify problems of the phylogenetic affinities of the
be probable is that organisms retain the inherited develop- echinoderms. The adult structure of echinoderms suggests
mental mechanisms of their ancestors. Hence at various that they are an invertebrate phylum, but their deuterosto-
stages in development it is likely that an organism will show mic embryological development confirms their affinities
structural similarities to the embryos of its ancestors. with the chordate line of development. This example
Subsequent adaptations to different environmental condi- illustrates the principle that phylogenetic relationships
tions and modes of life will modify later stages of the should not be decided purely on evidence of adult
developmental process. Observation reveals that the closer homologous structures.
the organisms are classified on the basis of common adult Evidence of the progressive development of various
homologous structures the longer their embryological groups on the basis of embryological evidence can be seen
development will remain similar. Organisms showing within the plant kingdom, but examples are less well
adaptations to certain modes of life and environments not documented than for the animal kingdom. The early
typical of the major group to which they belong show fewer gametophyte of mosses and ferns, as represented by the
similarities to other members of the group during their protonema produced by germination of the spores, has a
embryonic development. This is clearly seen in the similar structure, physiology and pattern of growth to the
development of the parasitic platyhelminths (flatworms) filamentous green algae from which they are therefore
Fasciola and Taenia, where a series of larval stages showing thought to have developed. The principle of alternation of
adaptations to secondary hosts exist which do not appear in generations in plant life cycles, and the homologous
the development of the free-living turbellarian platyhel- variations upon it reflecting adaptations to various environ-
minths, such as Planaria. Similarly, the terrestrial earth- mental conditions, may be interpreted as examples of
worm Lumbricus does not possess the ciliated trochopore homology and provide further evidence for evolutionary
larva which is typical of more primitive annelids. This relationships between plant groups.
evidence highlights the limitations of Haeckel’s principle The gymnosperms represent a group of plants which
of recapitulation. show features intermediate between those plants adapted
Study of the embryological development of major to a terrestrial existence and those plants which still require
groups of organisms reveals structural similarities evident water for the transfer of gametes. In the order Cycadales
in the embryonic and larval stages which are not apparent the male gametophyte resembles the light dry microspore
in the adult stages. These observations are interpreted as (pollen grain) of the angiosperms in that it is distributed by
suggesting phylogenetic relationships between various wind. As the male gametophyte develops, a pollen tube is
groups of organisms and the implication underlying this is formed as in angiosperms, but instead of this conveying a
that an evolutionary process exists. On the basis of the non-motile male gamete to the archegonium the terminal
cleavage patterns of the zygote and the fate of the (antheridial) cell gives rise to two flagellated antherozoids
blastopore, triploblastic animals may be divided into two (sperms) which swim to the ovule to bring about fertilisa-
groups, the protostomes and deuterostomes. Protostomes tion (fig 24.17). The cycads therefore appear to represent
an intermediate group between the lower plants and the
mouth Ce cilia angiosperms and this suggests that a phylogenetic conti-
nuum exists within the plant kingdom. The existence of a
Ly = => group of organisms possessing features common to two
Coy ees other groups showing different levels of complexity or
gn blastopore adapted to different environments, may be interpreted as
cleavage \ ¢
anus a
(-) embryo blastula stage typical deuterostome —
J like larva tube nucleus
©
= OS
tube Slight flagella
spiral
mouth
cilia
ee antheridial cell
(2
cleavage
anus
prothallial ee
blastopore
(a) ripe pollen (b) pollen tube (c) mature
typical protostome —like larva grain germinating antherozoid
Fig 24.16 Early developmental stages of deuterostomes and Fig 24.17 Development of motile male gametes from pollen
protostomes grain in Cycas
880
Fig 24.18 The primitive arthropod Peripatus
suggesting phylogenetic continuity between the three light on evolutionary ideas. The occurrence of similar
organisms based on the descent of one group (such as the molecules in a complete range of organisms suggests the
angiosperms) from the other (the lower plants) via the existence of biochemical homology in a similar way to the
intermediate form (the cycads). Many of these intermedi- anatomical homology shown by organs and tissues. Again,
ate forms are extinct and it is only by studying the fossil this evidence for an evolutionary theory is supportive of
record that a progressive developmental sequence can be other evidence rather than confirmatory in its own right.
deduced. In many cases intermediate forms have not been Most of the research which has been carried out on
found. These are equivalent to the ‘missing links’ that comparative biochemistry has involved analyses of the
appear as gaps in the fossil record. It may be that these links primary structure of widely distributed protein molecules,
do not exist according to the hypothesis of punctuated such as cytochrome c and haemoglobin, and more recently
equilibria (section 24.7.1). However if one accepts the of nucleic acid molecules, particularly ribosomal RNA.
adage ‘Natura non facit saltum’ (‘Nature does not make Slight changes in the genetic code as a result of gene
leaps’), their absence may be explained by the possibility mutation produce subtle variations in the overall structure
that they did not fossilise, have not yet been found, or even of a given protein or nucleic acid. This forms a basis for
did not exist. In the case of the phylogenetic link between determining phylogenetic relationships if the following
the annelids and the arthropods there is one group of assumption is made: the fewer the differences in the
organisms, the Onychophora, characterised by the genus molecular structure, the fewer the mutations which have
Peripatus, which has features of both annelids and occurred and the greater the affinity between organisms
arthropods (fig 24.18). The annelid features include a body containing the molecule. Large differences in the molecu-
wall containing circular and longitudinal muscles, non- lar structure represent large differences in the DNA and
jointed parapodia-like limbs, segmental nephridia-like predictably this situation exists in organisms showing fewer
excretory tubules and a double ventral nerve cord. The anatomical homologies.
arthropod features include a chitinous cuticle, spiracles and Cytochromes are respiratory proteins situated in the
tracheae and an open blood system. Another ‘living fossil’ mitochondria of cells and are responsible for the transfer of
intermediate form is represented by the Dipnoi (lungfish) electrons along the respiratory pathway which produces
which suggest a link between fish and amphibians. water and liberates the energy required to synthesise ATP
Whilst much of this evidence suggests that some form of molecules (section 11.5.4). Cytochrome c is one such
evolutionary process has occurred, it must be appreciated protein from the pathway. It is a conjugated protein
that there is no conclusive proof that it did occur. composed of an iron-containing prosthetic group sur-
rounded by a polypeptide chain containing between 104
and 112 amino acids, depending upon species. Modern
24.7.8 Comparative biochemistry techniques of computerised mass spectrometry have
As techniques of biochemical analysis have enabled the primary structure of the cytochrome c
become more precise, this field of research has shed new polypeptide chain to be worked out for a range of
881
Table 24.6 Cytochrom e c amino acid sequences for 21 species.
Oe a
Amino acid sequence
Species ~~oS 80 ‘Oo(sS)
So = wo w 52.68
7 8) ae N Ww ax Nn Oo XI Co \o SS — NO Ww - Nn
Man L Zz
Rhesus monkey Hrd
Horse
Pig, bovine, sheep
Dog
Grey whale
Rabbit
Kangaroo
Chicken, turkey
Penguin
Pekin duck
Snapping turtle
Bullfrog
Tuna
Screwworm fly
Silkworm moth el
Wheat
Fungus (Neurospora)
Fungus (baker’s yeast)
Fungus (Candida)
Bacterium (Rhodospirillum) ess)see
(walle)
es)
1S)
S)
(S)
SS)
(6)
(SS)
ZANS)
74
74,
a)
pe HHH
Se4e POUMCnNNSZESSSELESSEEE
ZeZASsoogse
HeseHeeeSes
MI
ee
Ue
lcs
Cara
octet
aCe
ae
Ca I
KO
SR
HR
ne CeCe
PFPUMMUP RRAAKR
KR
RH
RAARRRAAARK
KARR
mMmMmMtmmmmh
MMMM
MMM ST
ee
eet
Cree
ea
eae ZT SUD RA
UUU
UU
UU
LALLA RARAR
KAKA
RRA
ARR
RAR
RARAARAR
RAARAAR
DAARK
AKed
ZZLALACALALALA
AMAMrOmMmMOmOM
Momo B26HD
eee
ee
oe
eee
URDU
UU
UCU
DU
UU >
2260 << lANAAAAAAAD
AQQV>
DDD
D>
DDO
>DD
MAAAAAAAAANAAAAAAANAAAA
UU 4444H5 ARAAK
HRARR
RAARARA
RAR
ARRAAK
tot
Sel
ate
rete
etal
shat
Tel
ste
Petey
rel
neta
ote
MeN
Bad
)e|
eet
ted
| RAAR
KARR
AAA
4444944444444
Oy PP
PEPE
ee
Key to amino acids

A alanine F phenylalanine K lysine ie proline T threonine
C_ cysteine G glycine L_ leucine Q glutamine V_ valine
D aspartic acid H_ histidine M methionine R arginine W tryptophan
E glutamic acid I isoleucine N asparagine S_ serine Y tyrosine
After Dayhoff, M. O. and Eck, R. V. (1967-8) Atlas of protein sequence and structure, National Biomedical Research Foundation, Silver Spring, Md.
organisms, including bacteria, fungi, wheat, screwworm Table 24.7 Similarities and differences between the
fly, silkworm, tuna, penguin, kangaroo and primates. The polypeptide chains of haemoglobin in four primate species.
similarity in cytochrome c amino acid sequence between 21
organisms studied in this way is surprisingly high. In 20 out Polypeptide chains
of the 21 organisms studied, ranging from the athlete’s-foot a-haemoglobin (-haemoglobin
fungus to Man, the amino acids in positions 78-88 were (141 amino (146 amino
Species acids) acids) y-haemoglobin
identical (table 24.6). The amino acid sequence for
cytochrome c of Man and chimpanzee is identical and Man ate te +
differs from the rhesus monkey by only one amino acid. Chimpanzee + + 1
Computer studies, based on amino acid sequences of Gorilla 1 1 1
cytochrome c, have produced plant and animal phylogene- Gibbon 3 3 zy
tic trees which show close agreement with phylogenetic
trees based on anatomical homologies. Haemoglobin is composed of four polypeptide chains, made up of a, B,
and y polypeptides. + indicates no difference in amino acid sequence
Similar results have been obtained from the study of the from that of Man, figures indicate number of amino acid differences.
globin proteins, haemoglobin and myoglobin, involved in
oxygen transport and storage. The similarities and dif- present in serum, act as antigens when injected into the
ferences between the haemoglobin molecules of four bloodstream of animals that lack these proteins. This
primate species are shown in table 24.7. The relationships causes the animal to produce antibodies against them
between the various globins, based on amino acid which results in an antigen/antibody interaction (section
sequences, and their occurrence in organisms is shown in 14.14). This immune reaction depends upon the host
fig 24.19. Variations in the amino acid sequence of animal recognising the presence of foreign protein struc-
cytochrome c and the globins are thought to have arisen by tures in the serum. Human serum injected into rabbits
mutations of ancestral genes. sensitises them to human serum and causes them to
Immunological research has also produced evidence of produce antibodies against human serum proteins. After a
phylogenetic links between organisms. Protein molecules, period of time, if human serum is added to a sample of
882
myoglobin a-chain y-chain B-chain 6 -chain work has clarified many phylogenetic
(jawed (foetal (primates)
uncertainties
(Man and h l
vertebrates) mammals) apes) anon ssisLne mammals.
The separation of animal phyla into protostomes and
deuterostomes on the basis of embryological development
has been reinforced by analysis of the phosphate-
containing storage molecules found in muscle and used in
the synthesis of ATP. Protostomes, represented by
B -chain of mammals annelids, molluscs and arthropods contain arginine phos-
phate, whilst deuterostomes represented by echinoderms
and chordates, contain creatine phosphate.
A final example of biochemical homology is provided by
ancestral £-type chain the presence of similar or identical hormones in vertebrates
where they carry out a range of different functions. For
example, a hormone similar to mammalian prolactin
ancestral haemoglobin chain
occurs in all vertebrate groups where it is produced by the
pituitary gland. Although it has been reported that there
may be 90 distinct effects of prolactin, these can be
arranged under two broad headings, reproduction and
ancestral globin chain osmoregulation (table 24.9).
Fig 24.19 Supposed origins of myoglobin and vertebrate Table 24.9 Action of prolactin in vertebrates.
globin polypeptide chains. All five types are found in Man.
(After V. M. Ingram, (1963) Haemoglobins in genetics and Group Reproduction Osmoregulation
evolution, Columbia University Press.)
Bony fish Secretion of skin mucus _ Increases urine produc-
es : : ‘ tion
sensitised rabbit serum, antigen/antibody complexes form re: é (as ;
: a, Amphibia Secretion of ‘egg jelly Increases skin per-
which settle out as a Yaenenee that can be measured. meability to water
Adding aa from BALE! ; animals to'samples of Reptiles Suppresses egg Stimulates water loss in
rabbit serum containing antibodies against human serum production turtles
produces varying amounts of precipitate. pe that Birds Production of ‘crop milk’ Increases water uptake
the amounts of fore osale are directly related to the Mammals Mammary development. ADElike activity
amounts of ‘foreign’ protein present, this method can be Padilactaton
used to establish affinities between animal groups as shown
in table 24.8.
This technique of comparative serology has been used 24.7.9 Conclusion
extensively to corroborate phylogenetic links. For ex-
Neo-Darwinian evolutionary theory is based
ample, zoologists were uncertain as to the classification of
on evidence from a broad range of sources and supported
the king crab (Limulus). When various arthropod antigens
by a mass of otherwise unrelated observations. This
were added to Limulus serum the greatest amount of
constitutes to the scientist the strongest type of evidence for
precipitate was produced by arachnid antigens. This
the ‘validity’ of the theory. Evolution is widely accepted
evidence reinforced morphological evidence, and Limulus
amongst scientists but there is still much work to be done in
is now firmly established in the class Arachnida. Similar
refining the theory and its application to all observed
circumstances.
Table 24.8 Amounts of precipitate produced by adding
All scientific accounts, hypotheses and theories of the
serum from the following mammals to rabbit serum
history of life are tentative and, as long as Man remains
containing anti-human antibodies against human serum
objective in his search for truth, will remain so.
(amount of precipitate produced with human serum taken
Since evolution forms a focal point within the study of
as 100%).
biology it would be remiss to conclude this chapter without
relating evolution to the perspective of the natural world.
oe ae To do this it is fitting to quote from Darwin’s final
Berane 92% paragraph of the Origin of Species,
Gibbon 79% ‘There is a grandeur in this view of life, with its several
Baboon 75% powers, having been originally breathed by the Creator
Spider monkey a into a few forms or into one; and that, whilst this planet
ae 7% has gone cycling on according to the fixed law of gravity,
Pig 8% from so simple a beginning endless forms most beautiful
a
a and most wonderful have been and are evolving.’
883
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a
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in
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i 7 ' i f .
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S. i
7s ey > 7 ae
wih a mf a >, 7
Seyi. Pia. Se
ie CORON Tal
“—
‘ pray Neh tes Caley
en Sy —
bedix vn me. Nt;
ie 4c
» = vine aby ay '
ee as
hs ee
¥
,
~ a '
ee ar SA
Pah iC
awe
; 4 > on tes
va ‘=
2 aang bp arr
“Seyi, retire wa ete *
pen i
sins! (OE ak
ah alanny: > War
on Ss:!, wee,
as
eR
von
amen
nes.
are)vac » ga
- it Pye
ES On sae ay? aL
- Pp
78.4" i fae eae
3h 6 T fy mari
abe age beat 97
Awaisial?, GSP 20
iene
a
y> e ras F yy) .
' ae ee e : &*
=
Sa ere Th
oe s ~-*
taps. + mre
oe aad a
¥
“a
° teeea rei
oa nh 7a, itt
Chapter Twenty-five
NSE FGM ee 0 Fa ei wt 1 vy nas
Mechanisms of speciation
The previous chapter described how Darwin came to

duce before dying contribute to the future of the species.
appreciate that heritable variations occurred in natural
The fate of an individual organism is relatively insignificant
populations as well as in artificial breeding situations. He in the history of a species.
perceived that these heritable variations were significant in
evolutionary theory but had no means of accounting for the
mechanism by which variations could appear and charac- 25.1.1 Gene pool
teristics remain discrete. It was only with the reappearance A gene pool is the total variety of genes and
of the work of Mendel on inheritance, and the appreciation alleles present in a sexually reproducing population, and
of its importance in the understanding of evolutionary in any given population the composition of the gene pool
theory, that many of these problems could be resolved. may be constantly changing from generation to generation.
Modern explanations of variation between organisms are a New combinations of genes produce unique genotypes
blend of evolutionary theory based on the work of Darwin which, when expressed in physical terms as phenotypes,
and Wallace and genetic theory based on principles undergo environmental selection pressures which continu-
expounded by Mendel. Variation, inheritance and evolu- ally select and determine which genes pass on to the next
tionary theory may now be explained by evidence from a generation.
branch of biology known as population genetics. A population whose gene pool shows consistent change
from generation to generation is undergoing evolutionary
25.1 Population genetics
change. A static gene pool represents a situation where
genetic variation between members of the species is inade-
quate to bring about evolutionary change.
A population is a group of organisms of the
same species usually found in a clearly defined geograph-
ical area. Darwin was concerned how natural selection 25.1.2 Allele frequency
worked at the level of the individual organism in bringing The appearance of any physical characteristic,
about evolutionary change. Following the rediscovery of for example coat colour in mice, is determined by one or
Mendel’s demonstration of the particulate nature of more genes. Several forms of each gene may exist and these
inheritance, the importance of the genotype became are called alleles (table 23.2). The number of organisms in a
significant in the study of variation, inheritance and population carrying a particular allele determines the allele
evolutionary change. Bateson, the scientist who intro- frequency (which is sometimes, less correctly, referred to
duced the term ‘genetics’ in 1905, saw genetics as as the gene frequency). For example, in humans the
‘the elucidation of the phenomena of heredity and frequency of the dominant allele for the production of
variation’. pigment in the skin, hair and eyes is 99%. The recessive
It is the study of population genetics which forms the basis allele, which is responsible for the lack of pigment, a
of modern views of evolutionary theory, a theory called condition known as albinism, has a frequency of 1%. It is
neo-Darwinism, or the synthetic theory of evolution. usual in population genetic studies to represent gene or
Genes acting independently, or in conjunction with allele frequencies as decimals rather than percentages or
environmental factors, determine the phenotypic charac- fractions. Hence this dominant allele frequency is 0.99 and
teristics of organisms and produce variation within popula- the recessive albino allele frequency is 0.01. Since the total
tions. Phenotypes adapted to the environmental conditions population represents 100% or 1.0 it can be seen that:
or ‘ecological framework’ are ‘selected for’, whereas dominant allele frequency + recessive allele frequency =1
nonadaptive phenotypes are ‘selected against’ and even- 0.99 a 0.01 =1
tually eliminated. Whilst natural selection operating on the
phenotypic characteristics of individual organisms deter- In terms of Mendelian genetics the dominant allele would
mines the fate of its genotype, it is the collective genetic be represented bya letter, say N (for normal pigmenta-
response of the whole population that determines not only tion), and the recessive allele would be represented by n
the survival of the species but also the formation of new (the albino condition). In the example above, N = 0.99 and
species. Only those organisms which successfully repro- n = 0.01.
885
Population genetics has borrowed two symbols from the in producing evolutionary change, and when changes occur
mathematics of probability, p and gq, to express the the Hardy-Weinberg equation provides a means of study-
frequency with which a pair of dominant and recessive ing the change and of measuring its rate.
alleles appear in the gene pool of the population.
Therefore, 25.1.4 The Hardy-Weinberg equation
pt+q=1 Whilst the Hardy-Weinberg equation pro-
where p = dominant allele frequency, and q = recessive vides a simple mathematical model of how genetic
allele frequency. equilibrium can be maintained in a gene pool, its major
In the case of pigmentation in humans, p = 0.99 and q = application in population genetics is in calculating allele
0.01, and genotype frequencies.
since pit ga Starting with two homozygous organisms, one dominant
0.99 + 0.01 = 1 for allele A and one recessive for allele a, it can be seen that
all offspring will be heterozygous (Aa).
The value of the above equation lies in the fact that if the
frequency of either allele is known, the frequency of the Let A = dominant allele
other may be determined. For example, if the frequency of a = recessive allele
the recessive allele is 25% then gq = 25% or 0.25. Parental phenotypes homozygous xX homozygous
dominant recessive
Since Dek iG i= 1 AA x aa
p+0.25 =1 Meiosis
Dp o—1l—0:25 Gametes (n) ®@® x @©®
p =0.75 Random fertilisation
F, genotypes (2n) Aa Aa Aa Aa
That is, the frequency of the dominant allele is 0.75 or F, phenotypes all heterozygous
75%.
If the presence of the dominant allele A is represented by
25.1.3 Genotype frequencies the symbol p and the recessive allele a by the symbol q, the
The frequencies of particular alleles in the nature and frequency of the genotypes produced by
crossing the F, genotypes above are seen to be:
gene pool are of importance in calculating genetic changes
in the population and in determining the frequency of
F,, phenotypes heterozygous x heterozygous
genotypes. Since the genotype of an organism is the major F, genotypes (2n) Aa x Aa
factor determining its phenotype, calculations of genotype Meiosis
frequency are used in predicting possible outcomes of Gametes (n) Or @n as ® @
particular matings or crosses. This has great significance in Random fertilisation A a
horticulture, agriculture and medicine. (p) (4)
The mathematical relationship between the frequencies A AA Aa
of alleles and genotypes in populations was developed (p) | (p’) (pq)
independently in 1908 by an English mathematician G. H. a Aa aa
Hardy and a German physician W. Weinberg. The (q) | (pq) (q’)

F, genotypes (2n) AA 2Aa aa
relationship known as the Hardy-Weinberg equilibrium is
based upon a principle which states that
(p’) (2pq) Ca)
F, phenotypes homozygous heterozygous, homozygous
‘the frequency of dominant and recessive alleles in a dominant, recessive
population will remain constant from generation to
generation provided certain conditions exist.’ Since A is dominant, the ratio of dominant to recessive
These conditions are: genotypes will be 3:1, the Mendelian monohybrid cross
(1) the population is large; ratio. From the cross shown above it can be seen that the
(2) mating is random; following genotypes can be described in terms of the
(3) no mutations occur; symbols p and q:
(4) all genotypes are equally fertile, so that no selection
occurs; p= homozygous dominant
(5) generations do not overlap; 2pq = heterozygous
(6) there is no emigration or immigration from or into the ge homozygous recessive
population, that is, there is no gene flow between
populations. The distribution of possible genotypes is statistical and
Any changes in allele or genotype frequencies must based on probability. Of the three possible genotypes
therefore result from the introduction of one or more of the resulting from such a cross it can be seen that they are
conditions above. These are the factors that are significant represented in the following frequencies:
886
AA 2Aa aa the frequency of the dominant allele in the population is
dawaihty wh 0.99 or 99%.
In terms of genotype frequency the sum of the three Since
genotypes presented in the above population equal one,
0.99
or, expressed in terms of the symbols p and q, it can be S
p (0.99)?
seen that the genotypic probabilities are:
= 0.9801,
p+ 2pq+q=1 the frequency of the homozygous dominant genotype in the
(In mathematical terms p + g = 1 is the mathematical population is 0.9801, or approximately 98%.
equation of probability and p’ + 2pq + q* = 1 is the Since
binomial expansion of that equation (that is (p + q))).
p = 0.99 and q = 0.01,
To summarise, since
2pq = 2 X (0.99) x (0.01)
p = dominant allele frequency = 0.0198,
q = recessive allele frequency the frequency of the heterozygous genotype is 0.0198, or
p’ = homozygous dominant genotype approximately 2% of the population carry the albino allele
2pq = heterozygous genotype
either as heterozygotes or albino homozygotes.
q = homozygous recessive genotype These calculations reveal a surprisingly high value for the
it is possible to calculate all allele and genotype frequencies frequency of the recessive allele in the population
using the expressions: considering the low number of individuals showing the
homozygous recessive genotype.
allele frequency p+q =1, and Heterozygous individuals showing normal phenotypic
genotype frequency p* + 2pq + q’= 1. characteristics but possessing a recessive gene capable of
producing some form of metabolic disorder when present
However, in most populations it is only possible to in homozygous recessives are described as carriers.
estimate the frequency of the two alleles from the Calculations based on the Hardy-Weinberg equation show
proportion of homozygous recessives, as this is the only that the frequency of carriers in a population is always
genotype that can be identified directly from its pheno- higher then would be expected from estimates of the
type. occurrence of the disorder in the phenotype. This is shown
For example, one person in 10 000 is albino, that is to say in table 25.1.
that the albino genotype frequency is 1 in 10 000. Since the
albino condition is recessive, that person must possess the
homozygous recessive genotype and in terms of probability Table 25.1 Some metabolic disorders and the frequencies of
it can be seen that homozygous recessive and heterozygous genotypes.
1 Approximate
7 ~ 10 000 frequency of
homozygous reces-
Frequency of
‘carrier’ heterozygous
0.0001 Metabolic disorder sive genotype (q’) genotype (2pq)
Knowing that g° = 0.0001 the frequencies of the albino albinism 1 in 10000 1 in 50
allele (q), the dominant pigmented allele (p), the homozy- (lack of pigmenta- (in Europe)
gous dominant genotype (p’) and the heterozygous tion in body)
genotype (2pq) may be determined in the following alkaptonuria 1 in 1000 000 1 in 503
manner. (urine turns black
upon exposure to
Since
air)
gq = 0.0001 amaurotic family 1 in 40 000 1 in 100
q= V0.0001 idiocy
= 0.01, (leads to blindness
and death)
the frequency of the albino allele in the population is 0.01 diabetes mellitus 1 in 200 Lainie
or 1%. (failure to secrete
Since insulin)
phenylketonuria 1 in 10000 1in 50
pereg
=1 (may lead to mental (in Europe)
Dae GakG retardation if not
=1-0.01 diagnosed)
0.99,
887
action. It clearly shows the influence of an environmental
25.1 Cystic fibrosis of the ypanofcre1 asin = selection mechanism on changes in allele frequency, a
occurs in the population with a frequenc mechanism which disrupts the genetic equilibrium pre-
2 000. Calculate the frequency of the carrier geno- dicted by the Hardy-Weinberg principle. It is mechanisms
such as these that bring about the variations in populations
which lead to evolutionary change.
25.1.5 implications of the Hardy-Weinberg

equation
The Hardy-Weinberg equation shows that a large propor- 25.2 Factors producing changes in
tion of the recessive alleles in a population exist in carrier populations
heterozygotes. In fact, the heterozygous genotypes main-
tain a substantial potential source of genetic variability. As The Hardy-Weinberg equilibrium principle
a result of this, very few of the recessive alleles can be states that given certain conditions the allele frequencies
eliminated from the population in each generation. Only remain constant from generation to generation. Under
the alleles present in the homozygous recessive organism these conditions a population will be in genetic equilibrium
will be expressed in the phenotype and so be exposed to and there will be no evolutionary change. However the
environmental selection and possible elimination. Many Hardy-Weinberg equilibrium principle is purely theoretic-
recessive alleles are eliminated because they confer al. Few natural populations show the conditions necessary
disadvantages on the phenotype. This may result from the for the Hardy-Weinberg equilibrium to exist (section
death of the organism prior to breeding or genetic death, Dor):
that is the failure to reproduce. Not all recessive alleles, The four major sources of genetic variation within a gene
however, are disadvantageous to the population. For pool were described in detail in section 23.8.4, and they are
example, in human blood groups the commonest phenoty- crossing-over during meiosis, independent segregation
pic characteristic in the population is blood group O, the during meiosis, random fertilisation and mutation. The
homozygous recessive condition. This phenomenon is also first three sources of variation are often collectively
clearly illustrated in the case of sickle-cell anaemia. This is referred to as sexual recombination, and they account
a heritable disease of the blood common in certain for gene reshuffling. These processes however, whilst pro-
populations in Africa, India, certain Mediterranean coun- ducing new genotypes and altering genotype frequen-
tries and amongst North American negroes. Homozygous cies, do not produce any changes in the existing alleles,
recessive individuals usually die before reaching adulthood hence the allele frequencies within the population remain
thereby eliminating two recessive alleles from the popula- constant. Many evolutionary changes, however, usually
tion. Heterozygotes, on the other hand, do not suffer the occur following the appearance of new alleles and the
same fate. Studies have revealed that the sickle-cell allele major source of this is mutation.
frequency has remained relatively stable in many parts of Other situations in which the conditions for the Hardy—
the world. In some African tribes the genotype frequency is Weinberg equilibrium principle do not exist are when there
as high as 40%, and it was thought that this figure was is non-random breeding, when the population is small and
maintained by the appearance of new mutants. Investiga- leads to genetic drift, when genotypes are not equally
tions have revealed that this is not the case, and in many fertile so there is genetic load, and when gene flow occurs
parts of Africa where malaria is a major source of illness between populations. These situations are discussed
and death, individuals possessing a single sickle-cell allele below.
have increased resistance to malaria. In malaria regions of
Central America the selective advantage of the heterozy-
25.2.1 Non-random breeding
gous genotype maintains the sickle-cell allele in the
population at frequencies between 10 and 20%. The main- Mating in most natural populations is non-
tenance of a fairly constant frequency for a recessive random. Sexual selection occurs whenever the presence of
allele which may be potentially harmful is known as one or more inherited characteristics increases the likeli-
heterozygote advantage. In the case of North American hood of bringing about successful fertilisation of gametes.
negroes who have not been exposed to the selection effect There are many structural and behavioural mechanisms in
of malaria for 200-300 years the frequency of the sickle-cell both plants and animals which prevent mating from being
allele has fallen to 5%. Some of this loss may be accounted random. For example, flowers possessing increased size of
for by increased gene flow resulting from black-white petals and amounts of nectar are likely to attract more
marriages, but an important factor is the removal of the insects and increase the likelihood of pollination and
selection pressure for the heterozygote due to the absence fertilisation. Colour patterns in insects, fishes and birds,
of malaria in North America. As a result of this the and behavioural patterns involving nest-building, territory
recessive allele has slowly been eliminated from the possession and courtship, all increase the selective nature
population. This is an example of evolutionary change in of breeding.
888
An experimental investigation with Drosophila illus- frequency. Continual mating within a small population
trated the effect of non-random mating on genotype and decreases the proportion of heterozygotes and increases
allele frequencies. A culture of fruit flies containing equal the number of homozygotes. Examples of the founder
numbers of red-eyed and white-eyed males and females principle were shown by studies carried out on the small
was set up and within 25 generations all white-eyed fruit populations of religious sects in America who emigrated
flies were eliminated from the population. Observation from Germany in the eighteenth century. Some of these
revealed that both red-eyed and white-eyed females sects have married almost exclusively amongst their own
preferred mating with red-eyed males. Thus sexual members. In these cases they show allele frequencies which
selection, as a mechanism of non-random mating, ensures are uncharacteristic of either the German or American
that certain individuals within the population have an populations. In the case of the Dunkers, a religious sect in
increased reproductive potential so their alleles are more Pennsylvania, each community studied was made up of
likely to be passed on to the next generation. Organisms about 100 families, a population so small as to be likely to
with less favourable characteristics have a decreased lead to genetic drift. Blood group analyses produced the
reproductive potential and the frequency of their alleles following results:
being passed on to subsequent generations is reduced.
Blood group A
Pennsylvanian population 42%
25.2.2 Genetic drift West German population 45%
Dunker population 60%
This refers to the fact that variation in gene
frequencies within populations can occur by chance rather These values would appear to be the result of genetic drift
than by natural selection. Random genetic drift or the occurring within small populations.
Sewall Wright effect (named after the American geneticist Whilst genetic drift may lead to a reduction in variation
who realised its evolutionary significance) may be an within a population it can increase variation within the
important mechanism in evolutionary change in small or species as a whole. Small isolated populations may develop
isolated populations. In a small population not all the characteristics atypical of the main population which may
alleles which are representative of that species may be have a selective advantage if the environment changes. In
present. Chance events such as premature accidental death this way genetic drift can contribute to the process of
prior to mating of an organism which is the sole possessor speciation.
of a particular allele would result in the elimination of that
allele from the population. For example, if an allele has a 25.2.3 Genetic load
frequency of 1% (that is g = 0.01) in a population of
1000 000 then 10000 individuals will possess that allele. The existence within the population of dis-
advantageous alleles in heterozygous genotypes is known
In a population of 100 only one individual will possess that
allele so the probability of losing the allele from a small as genetic load. As mentioned in section 25.1.5, some
recessive alleles which are disadvantageous in the homozy-
population by chance is much greater.
gous genotype may be carried in the heterozygous
Just as it is possible for an allele to disappear from a
genotype and confer a selective advantage on the pheno-
population it is equally possible for it to drift to a higher
type in certain environmental conditions, such as the
frequency simply by chance. Random genetic drift, as its
sickle-cell trait in regions where malaria is endemic. Any
name implies, is unpredictable. In a small population it can
increase in recessive alleles in a population as a result of
lead to the extinction of the population or result in the
deleterious mutations will increase the genetic load of the
population becoming even better adapted to the environ-
population.
ment or more widely divergent from the parental popula-
tion. In due course this may lead to the origin of a new 25.2.4 Gene flow
species by natural selection. Genetic drift is thought to
have been a significant factor in the origin of new species on Within the gene pool of a given cross-
islands and in other reproductively isolated populations. fertilising population there is a continual interchange of
A phenomenon associated with genetic drift is the alleles between organisms. Providing there are no changes
founder principle. This refers to the fact that when a small in allele frequency as a result of mutation, gene reshuffling
population becomes split off from the parent population it will confer genetic stability or equilibrium on the gene
may not be truly representative, in terms of alleles, of the pool. If a mutant allele should arise it will be distributed
parent population. Some alleles may be absent and others throughout the gene pool by random fertilisation.
may be disproportionally represented. Continuous breed- Gene flow is often used loosely to describe the
ing within the pioneer population will produce a gene pool movement of alleles within a population as described
with allele frequencies different from that of the original above, but strictly speaking it refers to the movement of
parent population. Genetic drift tends to reduce the alleles from one population to another as a result of
amount of genetic variation within the population, mainly interbreeding between members of the two populations.
as a result of the loss of those alleles which have a low The random introduction of new alleles into the recipient
889
population and their removal from the donor population
affects the allele frequency of both populations and leads to mutation causes
the appearance of
increased genetic variation. Despite introducing genetic allele
variation into populations, gene flow has a conservative
effect in terms of evolutionary change. By distributing
mutant alleles throughout all populations, gene flow
ensures that all populations of a given species share a B allele is strongly
common gene pool, that is it reduces differences between selected for and
populations. The interruption of gene flow between passes to next
generation in
populations therefore is a prerequisite for the formation of increasing number
new species.
The frequency of gene flow between populations
depends upon their geographical proximity, and the ease
with which organisms or gametes can pass between the two
populations. For example, two populations may be
situated so close together that interbreeding is continuous
and they may be considered in genetic terms as being one
population since they share a common gene pool, for meiosis gametes two organisms with
example two snail populations in adjacent gardens sepa- random fertilisation the mutant allele mate
rated by a privet hedge.
It is relatively easy for flying animals and pollen grains to
be actively or passively dispersed into new environments.
Here they may interbreed or cross with the resident homozygous mutant allele
population, thereby introducing genetic variation into that expressed in the phenotype
population. Fig 25.1 Diagram showing the increased rate of spreading

of a mutant allele (a) through a population if linked to a
dominant allele (B) which is strongly selected for
25.3 Selection
This is a mechanism that can be thought of as differential advantages they exhibit when expressed as
occurring at two interrelated levels, at the level of the phenotypes. Selection pressure can then be seen as a means
organism and at the level of the alleles. of increasing or decreasing the spread of an allele within the
Selection is the process by which those organisms which gene pool and these changes in allele frequency can lead to
appear physically, physiologically and behaviourally better evolutionary change. Major changes in genotype arise
adapted to the environment survive and reproduce; those from the spread of mutant alleles through the gene pool.
organisms not so well adapted either fail to reproduce or The extent and timing of selection will depend upon the
die. The former organisms pass on their successful nature of the mutant allele and the degree of effect it has
characteristics to the next generation, whereas the latter do upon the phenotypic trait. If the allele is dominant, it will
not. Therefore selection can be seen to operate through the appear in the phenotype and be selected for or against. If
processes of differential mortality and differential repro- the allele is recessive, as is the case with most mutants, it
ductive potential. Selection has an adaptive significance in will not undergo selection until it appears in the homozy-
perpetuating those organisms most likely to ensure survival gous state. The chances of this occurring immediately are
of the species and depends upon the existence of slight and the allele may be ‘lost’ from the gene pool before
phenotypic variation within the population. appearing in the homozygous condition. An allele which is
When a population increases in size, certain environ- recessive in a given environment may persist until changes
mental factors become limiting, such as food availability in in the environment occur where it may have a dominant
animals and light in the case of plants. This produces effect. These effects would probably appear first in the
competition for resources between members of the heterozygote and selection would favour its spread
population. Those organisms exhibiting characteristics throughout the population, as in the case of sickle-cell
which give them a competitive advantage will obtain the anaemia.
resource, survive and reproduce. Organisms without those A recessive mutant allele may spread rapidly through a
characteristics are at a disadvantage and may die before population if it occupies a position (locus) on a chromo-
reproducing. Both environmental limiting factors and some very close (linked) to a functionally important
population size operate together to produce a selection dominant allele which is strongly selected for. In this
pressure which can vary in intensity. ‘linked’ condition the chances of the mutant. allele
Therefore, selection is the process determining which combining with another mutant allele to produce the
alleles are passed on to the next generation by virtue of the homozygous condition are increased (fig 25.1).
890
The influence of a given mutant allele can vary. Those mortality. From these results it is possible to calculate the
mutations affecting alleles controlling important functions intensity of selection pressure.
are likely to be lethal and removed from the population If 614 babies died at birth or within one month this
immediately. Evolutionary change is generally brought represents a mortality of 4.5%. Even at the optimum birth
about by the gradual appearance of many mutant alleles weight 1.8% of babies died. Hence the selection pressure
which exert small progressive changes in phenotypic for weight at birth for babies of 3.6 kg is 4.5% —1.8% =
characteristics. 2.7% or 0.027. At a birth weight of 1.8 kg there is a 34%
There are three types of selection process occurring in mortality giving an intensity of selection pressure at this
natural and artificial populations and they are called weight of approximately 30% or 0.3. It should be pointed
stabilising, directional and disruptive. They may be best out, however, that advances in paediatric medicine have
explained in terms of the normal distribution curve considerably reduced post-natal mortality since 1946.
associated with the continuous phenotypic variation found Stabilising selection pressures do not promote evolution-
in natural populations (fig 25.2). ary change but tend to maintain phenotypic stability within
the population from generation to generation.
25.3.1 Stabilising selection
This operates when phenotypic features coin- 25.3.2 Directional selection
cide with optimal environmental conditions and competi- This form of selection operates in response to
tion is not severe. It occurs in all populations and tends to gradual changes in environmental conditions. It operates
eliminate extremes from the population. For example, on the range of phenotypes existing within the population
there is an optimum wing length for a hawk of a particular and exerts selection pressure which moves the mean
size with a certain mode of life in a given environment. phenotype towards one phenotypic extreme. Once the
Stabilising selection, operating through differential repro- mean phenotype coincides with the new optimum environ-
ductive potentials will eliminate those hawks with wing mental conditions stabilising selection will take over.
spans larger or smaller than this optimum length. This kind of selection brings about evolutionary change
Karn and Penrose carried out a study on the correlation by producing a selection pressure which favours the
between birth weight and post-natal mortality on 13 730 increase in frequency of new alleles within the population.
babies born in London between 1935 and 1946. Of these Directional selection forms the basis of artificial selection
614 were still-born or died within one month of birth. where the selective breeding of phenotypes showing
Fig 25.3 shows that there is an optimum birth weight of desirable traits increases the frequency of those pheno-
about 3.6 kg. Babies heavier or lighter than this are at a types within the population (section 25.4). In a series of
selective disadvantage and have a slightly increased rate of experiments, D. S. Falconer selected the heaviest mice
Stabilising Directional Disruptive /\

0 0 N N 0 N
|
| | |
! | | |
| | |
|
frequency !
3
|
!
5
A
pees 3
|
Generations
0
|
frequency
ee | har 4
2 Generations
| >
:
0 0
- ——_>
frequency | Selection . Selection
pressure
<_< — >
phenotypic characteristic phenotypic characteristic phenotypic characteristic

1 ! ] Generations
Fig 25.2 Diagrams showing the three types of selection operating within populations. O indicates the original coincidence
between optimum phenotype and optimum environmental conditions; N indicates the new position of coincidence of optimum
phenotype and optimum environmental conditions. Organisms possessing characteristics in the shaded portions of the normal
distribution are at a selective disadvantage and are eliminated by selection pressure. (The numbers 1-3 indicate the order of
generations.)
891
100, 25.3.4 Intensity of selection pressure
The intensity of selection pressure within a
population varies at different times and in different places
and may be produced by changes in external or internal
75,
factors. External factors may include an increase in
numbers of predators or pathogens or competition from
other species (interspecific competition) for food and
survival
Percentage breeding space in the case of animals, and light, water and
50+
mineral salts in the case of plants. Changes in climatic
l ! ie l a J | conditions or the state of the habitat in which organisms
live may exert new selection pressures. Internal factors
(2
aes Deel S26 41 4.5 50 e5:4
Birth weight/kg such as a rapid increase in the size of the population can
result in increased competition for environmental re-
Fig 25.3 The relationship between percentage survival and
birth weight in human babies. (After M. N. Karn & L. S. sources (intraspecific competition). As the population size
Penrose (1951) Ann. Eugen., London, 16, 147-64.) increases, so do the numbers of parasites and predators.
Pathogens, too, are more easily transmitted from organism
to organism as the host population rises and diseases spread
from a population at six weeks and let them inbreed. He very rapidly. All of these factors may not only affect the
also selected the lightest mice and allowed them to inbreed. intensity of the selection pressure but also the direction of
This selective breeding according to mass resulted in the the pressure. ‘New’ phenotypes (and genotypes) are
production of two populations, one increasing in mass and selected for, and poorly adapted organisms are eliminated
the other decreasing (fig 25.4). After termination of from, the population. The organisms‘to be eliminated first
selective breeding neither group returned to the original are those at the non-adaptive extremes of the phenotypic
population mass of approximately 22 g. This suggested that range.
the artificial selection of phenotypic characteristics led to One result of increased selection pressure is that it may
some genotypic selection and some loss of alleles from each cause organisms to become specialised to certain modes of
population. Many classic examples of natural directional life or narrower environmental conditions. This may be a
selection can be seen in evidence today and they are disadvantage for the future of that species. Increased
discussed in section 25.5. uniformity and dependency by a species increases the
likelihood of that species becoming extinct should environ-
25.3.3 Disruptive selection mental conditions change. The fossil record contains many
extinct organisms that were bizarre and overspecialised.
This is probably the rarest form of selection
but can be very important in bringing about evolutionary 25.2 How might a knowledge of selection
change. Fluctuating conditions within an environment, say pressure and mode of life be useful in the eradication
associated with season and climate, may favour the
presence of more than one phenotype within a population.
of anmamed parasite?
Selection pressures acting from within the population as a
result of increased competition may push the phenotypes
away from the population mean towards the extremes of
the population. This can split a population into two sub-
populations. If gene flow between the subpopulations is
prevented, each population may give rise to a new species.
24
In some cases this form of selection can give rise to the
appearance of different phenotypes within a population, a 22
phenomenon known as polymorphism (poly, many; mor-
phos, form), and is discussed in section 25.5.1. Within a 20
species organisms with different phenotypes, or ecotypes,
may show adaptations to particular environmental condi- weight/g
Body
tions (section 25.6.2). When a species occupies an
extremely large geographical range, organisms distributed
along it may show local changes in phenotypic characteris-
tics which are intermediate between those at the extremes 2 4 6 8 10
of the range. This continuous gradation of characteristics Number of generations
along a geographical range is usually a phenotypic response Fig 25.4 Changes in weight in two mouse populations in
to climate and/or edaphic (soil) variables and is known as a successive generations undergoing selection for body weight.
cline (section 25.6.3). (After D. S. Falconer (1953) J. Genetics, 51 470-501.)
892
From what has been said it can be seen that increased quality meat but lack the hardiness and foraging ability of
selection pressure is a conservative mechanism selecting
the Cheviot and Scotch Blackface. The latter examples
for the phenotype best adapted to the prevailing environ-
show that no single breed has all the characteristics
mental condition (the optimum phenotype). necessary for the best economic yield under all conditions
A reduction in the intensity of selection pressure usually and therefore a planned programme of selective breeding is
has the opposite effects to those described above. It may be often practised to increase the quality of the breed and the
produced by an absence of predators, pathogens, parasites yield.
and competing species or an increase in optimum environ- In artificial selection Man is exerting a directional
mental conditions. These conditions are usually found selection pressure which leads to changes in allele and
when an organism is introduced into a new environment. It genotype frequencies within the population. This is an
is conditions such as these which are believed to have evolutionary mechanism which gives rise to new breeds,
favoured the diversity of finch species found on the strains, varieties, races and subspecies. In all cases these
Galapagos Islands. groups have isolated gene pools, but they have retained the
basic gene and chromosomal structure which is character-
istic of the species to which they still belong.
25.4 Artificial selection
Man has practised artificial selection in the

25.4.1 Inbreeding
form of the domestication of animals and plants since the This involves selective reproduction between
earliest times of his civilisation. Darwin used evidence from closely related organisms, for example between offspring
artificial selection to account for the mechanism whereby produced by the same parents, in order to propagate
changes in species could arise in natural populations, that is particularly desirable characteristics. Inbreeding is a
natural selection. The basis of artificial selection is the particularly common practice in the breeding of ‘show’
isolation of natural populations and the selective breeding animals such as cats and dogs. It was used by livestock
of organisms showing characteristics or traits which have breeders to produce cattle, pigs, poultry and sheep with
some usefulness to Man. In the case of cattle, the Hereford high yields of milk, meat, eggs and wool respectively, but
and Aberdeen Angus breeds have been selected for the for reasons stated below inbreeding is not now widely
quality and quantity of their meat, whereas Jersey and practised.
Guernsey cows are favoured for their milk yield. Hamp- Prolonged inbreeding can lead to a reduction in fertility
shire and Suffolk sheep mature early and produce a good and this is a particular problem in the breeding of livestock.
greeoe :
eRe y
: Pomel
i vi
5 An example of hybridid vigour. Photograph (a) shows two parental maize j w hen interbred produce thethe —
} stalks which
by hybridisation
oe shown in eect ofthe photograph. The ear shown in the centre of the photograph (b) was produced
by D. F. Jones, Connecticut
of parental stocks with ears A and B as shown on the left and right of the photograph. (Photograph
Agricultural Experiment Station. )
893
s
Intensive inbreeding reduces the variability of the genome strains, and in certain plants between closely related
(the sum of all the alleles of an individual) by increasing species. The progeny are known as hybrids, and have
the number of homozygous genotypes at the expense of the phenotypes showing characteristics which are superior to
number of heterozygous genotypes. In order to overcome either of the parental stocks. This phenomenon is known as
these problems breeders resort to outbreeding after several hybrid vigour or heterosis. Hybrids produced from crossing
generations of inbreeding. For example, a dairy farmer homozygous parental stocks from different populations are
may use his own bull and successive generations of his own called F, hybrids and show advantages such as increased
cows to produce cows with a high milk yield. Before the fruit size and number, increased resistance to disease and
cattle begin to show signs of decreased resistance to disease earlier maturity. In maize (sweet corn), hybridisation has
and reduced fertility, the farmer will use another bull or increased the grain yield of the F, hybrids by 250% over the
artificially inseminate his breeding cows with semen parental stocks (fig 25.5). In the case of double-cross
acquired from a cattle-breeding centre. This introduces hybridisation, the hybrids produced by crossing two inbred
new alleles into the herd, thereby increasing the hetero- strains are themselves crossed. The resulting hybrid
zygosity of the breeding population. produces ears having the quality and yield which more than
covers the costs involved in a two-year breeding program-
me (fig 25.6).
Increased vigour results from the increased heterozy-
gosity which arises from gene mixing. For example, whilst
25.4.2 Outbreeding each homozygous parent may possess some, but-not all, of
This is particularly useful in plant breeding, the dominant alleles for vigorous growth, the heterozygote
but is being used increasingly in the commercial production produced will carry all the dominant alleles, as shown in
of meat, eggs and wool. It involves crossing individuals fie 25.27:
from genetically distinct populations. Outbreeding usually Increased vigour in certain varieties may not result
takes place between members of different varieties or simply from the increased prominence of dominant alleles,
Fig 25.6 The phenotypes produced by double-cross hybridisation in maize. TI j |

crossing the hybrids of the inbred strain (shown on theleft) Se ee
a peti Gimmie ia giana
894
Fig 25.7 A simple genetic explanation of increased vigour in
25.4.3 Artificial selection in Man
F, hybrids
Recent advances in Man’s knowledge of the
FFgghhliljj x FFGGHHiiJJ
structure of the gene, the genetic code, the mechanisms of
Meiosis heredity and the prenatal diagnosis of genetic defects, have
Gametes (n) FehlJ «@GHid) opened up the possibilities of selecting or eliminating
Random fertilisation certain characteristics in Man. The science of eugenics is
F, genotypes (2n) FfGgHhliJj concerned with the possibilities of ‘improving’ the ‘quality’
F, phenotypes This carries a dominant allele for of the human race by the selective mating of certain
each gene individuals. This is a very emotive topic and raises all sorts
of objections. Aldous Huxley in his book Brave New
but also from some form of interaction between particular World, published in 1932, fictionalised the day when
combinations of alleles in the heterozygote. eugenics would be taken to its extreme possibilities and
If F; phenotypes are continually inbred the vigour will particular types of individuals would be produced accord-
decrease as the proportion of homozygotes increases (fig ing to the needs of society at that time. Whilst these ideas
25:85 are repugnant to societies in which the freedom and rights
Selective hybridisation can induce changes in chromo- of the individual are paramount, there are strong argu-
some number (chromosomal mutation), a phenomenon ments for the exercise of limited forms of eugenic practice.
known as polyploidy, which can lead to the production of In medicine, genetic counselling is becoming more accep-
new species. An example of this is described in section table as a means of informing couples with family histories
eo 92: of genetic abnormalities about the possible risks involved
e a
. nee id vigour
vi
i .8 Maize stalks of eight generations. The seven stalks on the right demonstrate loss of hybrid as a resul It of
oaeaits from the hybrid Siouh on the left. The last three generations show reduced loss of vigour as a result of their
becoming homozygous. (Photograph by D. F. Jones, Connecticut Agricultural Experiment Station.)
895
in having children. By applying the Hardy-Weinberg (a)
equation it is possible to calculate the number of carriers of
metabolic disorders such as phenylketonuria or abnormali-
ties of the blood, such as thalassaemia, sickle-cell anaemia
or haemophilia. Known carriers can be advised as to the
likelihood of marrying another carrier and the possibilities
of producing offspring affected by the disorder. Such forms
of preventive medicine offer advice rather than dictate
policy. Any scientific advances which reduce suffering must
receive sympathetic appreciation. The dangers of eugenics
lie in their possible abuse.
(5)
25.5 _—Natural selection
Natural selection, as postulated by Darwin
and Wallace, represented a hypothesis based on historical
evidence. For Darwin, the time span involved in the
evolutionary change of a population was such that it could
not be observed directly. Recent changes accompanying
the industrial, technological and medical revolutions have
produced such strong directional and disruptive pressures
that we can now observe the results of dramatic changes in
genotypic and phenotypic characteristics of populations
within days. The introduction of antibiotics in the 1940s
Fig 25.9 Polymorphic forms of peppered moth, Biston
provided a strong selection pressure for strains of bacteria betularia. (a) The normal form, Biston betularia typica; (b) the
that have the genetic capability of being resistant to the melanic form, Biston betularia carbonaria. (From E. B. Ford
effects of the antibiotics. Bacteria reproduce very rapidly, (1973) Evolution studied by observation and experiment,
producing many generations and millions of individuals Oxford Biology Readers, 55, Oxford University Press.)
each day. Random mutation may produce a resistant
organism in the population which will thrive in the absence trunks of trees. The normal form of the moth is extremely
of competition from other bacteria which have been well camouflaged as its colouration merges with that of the
eliminated by the antibiotic. As a result, new antibiotics lichens growing on the trunks. With the spread of the
have to be developed to eliminate the resistant bacteria, industrial revolution sulphur dioxide pollution from the
and so the cycle continues. Other examples of the effects of burning of coal killed off the lichens growing on trees in
chemicals in producing selection pressure have been seen industrial areas, exposing the darker bark which was
with DDT on body-lice and mosquitoes and the effect of further darkened by soot deposits (fig 25.10).
the anticoagulant warfarin on rats. Following the develop- Kettlewell, in the 1950s released known numbers of
ment of resistant strains they spread very rapidly through- marked light and dark forms into two areas, one a polluted
out the population. area near Birmingham where 90% of the population was
Perhaps the classic example of evolutionary change is the black form, and the other an unpolluted area in Dorset
provided by the response of moth species to the directional where the dark form was rarely found. On recapturing the
selection pressure produced by the atmospheric pollution moths using a light trap he obtained the following results:
which accompanied the industrial revolution. Within the
last 100 years darkened forms of about 80 species of moths Birmingham — Dorset
have appeared in varying frequencies throughout the Percentage marked dark form 34.1 6.3
United Kingdom. This is a phenomenon known as Percentage marked light form 15.9 12:5
industrial melanism. Up to 1848 all reported forms of the Kettlewell demonstrated using cine-film that robins and
peppered moth (Biston betularia) appeared creamy-white thrushes feed on the moths. This is a form of natural
with black dots and darkly shaded areas (fig 25.9). In 1848 a selection known as selective predation, and it acts as a
black form of the moth was recorded in Manchester, and by selection pressure on the distribution of the melanic and
1895, 98% of the peppered moth population in Manchester non-melanic forms.
was black. This black ‘melanic’ form arose by a recurring The results show that the melanic form of the moth,
random mutation, but its phenotypic appearance had a Biston betularia carbonaria, has a selective advantage in
strong selective advantage in industrial areas for reasons industrial areas over the lighter form, Biston betularia
put forward and tested by Dr H. B. D. Kettlewell. typica, whereas the lighter form has the selective advantage
The moths fly by night and during the day they rest on the in non-polluted areas.
896
Subsequent research has demonstrated that the coloura-
defined as the existence of two or more forms of the same
tion of the dark form is due to the presence of a dominant
species within the same population, and can apply to
melanic allele. Fig 25.11 shows a recent distribution of the
biochemical, morphological and behavioural characteris-
two forms in the British Isles.
tics. There are two forms of polymorphism, transient
The presence of melanic forms in non-industrial areas of
polymorphism and balanced, or stable, polymorphism.
the east of England is explained by the distribution of
A classic quantitative study of balanced polymorphism
melanic forms by prevailing westerly winds. Since the
was carried out by Cain, Currey and Shepherd on the
introduction of the Clean Air Act in 1956 the proportion of
common land snail Cepaea nemoralis. The shells of this
non-melanic forms has increased slightly as the selection
species may be yellow (and appear green with the living
pressure on these forms has been reduced in industrial
snail inside), brown, or various shades including pale fawn,
areas. pink, orange and red. The lip of the shell may be dark
brown, pink or white and the whole shell may have up to
25.5.1 Polymorphism five dark brown bands following the contours of the shell
Polymorphism plays a significant role in the (fig 25.12). Both colouration and banding pattern are
process of natural selection. It demonstrates many of the determined genetically. The colours are determined by
principles outlined earlier in the chapter regarding the multiple alleles with brown being dominant to pink and
relationship between genotype frequency within the both being dominant to yellow. Banding is recessive.
population and variations in selection pressure. It is Studies have revealed that the snails are predated upon
by thrushes which carry the snails to a nearby stone which
they use as an ‘anvil’ to crack open the shell; the snail inside
is then eaten. By studying the proportions of types of shell
found near an anvil with those in the immediate habitat,
Cain, Currey and Shepherd demonstrated that selective
forces were at work within the population. In areas where
the background was fairly uniform, such as grass and
woodland litter, the yellow and brown unbanded shells had
a selective advantage as fewer of these shells were found
near the anvil (fig 25.13). In areas where the ground cover
was tangled and mottled, as in rough pasture or hedgerows,
the darker banded shells had a selective advantage. The
)
© Biston betularia typica
© Biston betularia
carbonaria
Fig 25.10 Melanic and non-melanic forms of Biston betularia

on tree trunks in (a) an area near Birmingham, and (b) an Fig 25.11 The distribution of melanic and non-melanic forms
area in Dorset. (Courtesy of Dr H. B. D. Kettlewell, of Biston betularia in the British Isles in 1958. (After H. B. D.
Department of Zoology, University of Oxford.) Kettlewell (1978) Heredity, 12, 51-72.)
897
Fig 25.12 Colour and banding pattern variation in the shells of Cepaea nemoralis. The extremes of colour and banding are
shown as a progression from yellow unbanded (top left) to brown banded (bottom right). Photographs (a) and (b) show top and
side views of the same shells. (After Tribe, Tallan & Erant (1978) Basic Biology Course, Book 12, Cambridge University
Press.)
898
Fig 25.13 Unbanded shells of
Cepaea nemoralis against a back-
ground of leaf litter. The shell on the
extreme right is yellow, the shell at
the top of the photograph is pink and
the two shells on the left are brown.
(After E. B. Ford (1973) Evolution
Studied by observation and experi-
ment, Oxford Biology Reader, 55,
Oxford University Press.)
forms suffering the greatest predation in any area were heterozygotes which forms the basis of the polymorphism.
those which were visually conspicuous to the thrushes. A The added linkage of genes controlling certain physio-
large population of polymorphic snails may include several logical effects is also thought to contribute to the mainten-
areas with a range of backgrounds. Seasonal effects also ance of the balanced polymorphism. The existence of a
produce changes in background colour and pattern. number of distinct inherited varieties coexisting in the
Although predation of conspicuous forms is continuous same population at frequencies too great to be explained
there is no overall selective advantage for any form, hence by recurrent mutations, as in the case of Cepaea, is called
the numbers of each form within a population remain fairly genetic polymorphism.
constant from year to year.
The balance in numbers of each form may not be Transient polymorphism
determined purely by colour and banding pattern. There is This arises when different forms, or morphs, exist in a
evidence to suggest that physiological effects may help to population undergoing a strong selection pressure. The
maintain the polymorphic equilibrium. In some areas frequency of the phenotypic appearance of each form is
where the soil is calcareous and dry and the background determined by the intensity of the selection pressure, such
cover is light, the dominant forms are not always those with as the melanic and non-melanic forms of the peppered
the least conspicuous colour and banding pattern. The moth. Transient polymorphism usually applies in situations
genetic basis for the polymorphism shown by Cepaea is where one form is gradually being replaced by another.
thought to rely on the existence of a special form of gene
linkage. The genes for colour and banding pattern are Balanced polymorphism
linked and form a super-gene which acts as a single genetic This occurs when different forms coexist in the same
unit and is inherited as such. These genes determine population in a stable environment. It is illustrated most
characteristics which have such a selective advantage that clearly by the existence of the two sexes in animals and
they are maintained within the population. It is the variety plants. The genotypic frequencies of the various forms
of allelic forms of these genes, maintained by the exhibit equilibrium since each form has a selective
899
Table 25.2 Alternative ways of defining a species.
advantage of equal intensity. In Man, the existence of the
A, B, AB and O blood groups are examples of balanced
e
e
Definition
polymorphism. Whilst the genotypic frequencies within Biological aspect
different populations may vary, they remain constant from Breeding A group of organisms capable of interbreeding
generation to generation within that population. This is and producing fertile offspring
because none of them has a selective advantage over the Ecological A group of organisms sharing the same
others. Statistics reveal that white men of blood group O ecological niche; no two species can share the
have a greater life expectancy than those of other blood same ecological niche
groups but, interestingly, they also have an increased risk Genetic A group of organisms showing close similarity
in genetic karyotype
of developing a duodenal ulcer which may perforate and
lead to death. Red-green colour blindness in Man is Evolutionary A group of organisms sharing a unique collec-
rile
tion of structural and functional characteristics
another example of polymorphism, as is the existence of
workers, drones and queens in social insects and pin and
ee
SS
thrum-eyed forms in primroses. races along the coast of California. One race, the ‘sun’ race,
is found on exposed southerly facing grassy slopes, whilst
25.6 The concept of species the ‘shade’ race is found in shaded oak woodlands and
redwood groves. These races differ in the size of their
A species represents the lowest taxonomic petals, a characteristic which is determined genetically.
group which is capable of being defined with any degree of
precision. It may be defined in a variety of ways and some 25.6.3 Clines
of these are summarised in table 25.2. A species exhibiting a gradual change in
Organisms belonging to a given species rarely exist
phenotypic characteristics throughout its geographical
naturally as a single large population. It is usual for a
range is referred to as a cline. More than one cline may be
species to exist as small interbreeding populations, called
exhibited by a species and they may run in opposite
demes, each with its own gene pool. These populations may directions as shown by fig 25.14.
occupy adjacent or widely dispersed geographical areas.
Species exhibiting marked phenotypic variation within a
Spatial separation of populations means that the species
population according to their degree of geographical
may encounter a variety of environmental conditions and
isolation are known as polytypic species. One classic form
degrees of selection pressure. Mutation and selection
of a polytypic species is illustrated by gulls belonging to the
within the isolated populations may produce the following
genus Larus (section 25.8.4).
degrees of phenotypic variation within the species.
All cases of phenotypic variation described above
25.6.1 Geographical races represent varying degrees of genetic dissimilarity which
may interfere with the breeding potential of members of
Populations which are distributed over a wide the populations if brought together.
geographical range or have occupied well-separated geo-
graphical habitats for a long period of time may show 25.7 Speciation
considerable phenotypic differences. These are usually
based on adaptations to climatic factors. For example, the This is the process by which one or more
gypsy moth (Hymantria dispar) is distributed throughout species arise from previously existing species. A single
the Japanese Islands and eastern Asia. Over this range a species may give rise to new species (intraspecific specia-
variety of climatic conditions are encountered, ranging tion), or, as is common in many flowering plants, two
from subarctic to subtropical. Ten geographical races have different species may give rise to a new species (inter-
been recognised which differ from each other with regard specific hybridisation). If intraspecific speciation occurs
to the timing of hatching of their eggs. The northern races whilst the populations are separated it is termed allopatric
hatch later than the southern races. The phenotypic speciation. If the process occurs whilst the populations are
variations shown by the ten races are thought to be the occupying the same geographical area it is called sympatric
result of climatic factors producing changes in gene speciation.
frequencies within their gene pools. The evidence that
these variations are genetically controlled is shown by the 25.8 Intraspecific speciation
fact that under identical environmental conditions the
different races still hatch at different times. There are several factors involved in intra-
specific speciation, but in all cases gene flow within
25.6.2 Ecological races (ecotypes) populations must be interrupted. As a result of this each
Populations adapted to ecologically dissimilar subpopulation becomes genetically isolated. Change in
habitats may occupy adjacent geographical areas; for allele and genotype frequencies within the populations, as
example the plant species Gilia achilleaefolia occurs as two a result of the effects of natural selection on the range of
900
Table 25.3 Isolating mechanisms (after Dobzhansky).
Prezygotic mechanisms (barriers to the formation of hybrids)

Seasonal isolation Occurs where two species mate or flower
W at different times of the year; for
E
example in California Pinus radiata
flowers in February whereas Pinus
attenuata flowers in April
Ecological isolation Occurs where two species inhabit similar
regions but have different habitat
small
preferences; for example Viola arvensis
grows on calcareous soils whereas Viola
tricolor prefers acid soils
Fig 25.14 Phenotypic variation in size and fur Behavioural isolation Occurs where animals exhibit courtship
colour shown
by the wood mouse (Apodemus) in Scotland patterns, mating only results if the
courtship display by one sex results in
phenotypes produced by mutation and sexual recom acceptance by the other sex; for example
bina- certain fish, bird and insect species
tion, lead to the formation of races and subspecies.
If the Mechanical isolation Occurs in animals where differences in
genetic isolation persists over a prolonged period of
time genitalia prevent successful copulation
and the subspecies then come together to occupy the same
and in plants where related species of
area they may or may not interbreed. If the breeding flowers are pollinated by different
is
successful they may still be considered to belong to animals
the
Same species. If the breeding is unsuccessful, then
Postzygotic mechanisms (barriers affecting hybrids)
speciation has occurred and the subspecies may now be
considered to be separate species. This is the way in which Hybrid inviability Hybrids are produced but fail to develop
to maturity; for example hybrids formed
it is believed evolutionary change can be brought about.
between northern and southern races of
An initial factor in the process of speciation may be the the leopard frog (Rana pipiens) in
reduction in the intensity of selection pressure within the North America
population. This may lead to increased intraspecific Hybrid sterility Hybrids fail to produce functional
variability. These new phenotypes may enable the popula- gametes; for example the mule (2n =
tion to increase its geographical range if the phenotypes 63) results from the cross between the
horse (Equus equus, 2n = 60) and the
show adaptations to environmental conditions found at the
ass (Equus hemionus, 2n = 66)
extremes of the range. Providing there is no reduction in
Hybrid breakdown F, hybrids are fertile but the F,
gene flow throughout the population, the species, whilst generation and backcrosses between F,
exhibiting the localised phenotypic variation (ecotypes), hybrids and parental stocks fail to
will still share the same gene pool and continue to exist as a develop or are infertile, for example
single species. This is the situation found in a cline. hybrids formed between species of
Speciation will only occur as a result of the formation of cotton (genus Gossypium)
barriers which lead to reproductive isolation between
members of the population. Reproductive isolation is separation. This inability of organisms or their gametes to
brought about by some form of what the geneticist meet leads to reproductive isolation. Adaptations to new
Theodosius Dobzhansky called isolating mechanism. conditions or random genetic drift in small populations
lead to changes in allele and genotype frequencies.
25.8.1 Isolating mechanisms Prolonged separation of populations may result in them
becoming genetically isolated even if brought together. In
An isolating mechanism is a means of produc-
this way new species may arise. For example, the variety
ing and maintaining reproductive isolation within a
and distribution of the finch species belonging to the family
population. This can be brought about by mechanisms
Geospizidae on the islands of the Galapagos archipelago
acting before or after fertilisation. Dobzhansky suggested a
are thought to be the result of allopatric speciation. David
classification of isolating mechanisms which has been
Lack suggested that an original stock of finches reached the
modified and is shown in table 25.3.
Galapagos Islands from the mainland of South America
and, in the absence of competition from endemic species
25.8.2 Allopatric speciation
(representing relaxed selection pressure), adaptive radia-
Allopatric (allos, other; patria, native land) tion occurred to produce a variety of species adapted to
speciation is characterised by the occurrence, at some particular ecological niches. The various species are
stage, of spatial separation. Geographical barriers such as believed to have evolved in geographical isolation to the
mountain ranges, seas or rivers, or habitat preferences, point that when dispersal brought them together on certain
may produce a barrier to gene flow because of spatial islands they were able to coexist as separate species.
901
of two
25.8.3 Sympatric speciation Fig 25.16 The Galapagos Islands and the distribution
e follow ing allopa tric
species of finch illustrating coexistenc
Genetic differences may accumulate allopatri- speciation
cally in populations which have been geographically
isolated for a much shorter period of time. If these
populations are brought together, hybrids may form where Equator
these overlap. For example, both the carrion crow (Corvus
corone) and the hooded crow (Corvus corone cornix) are |
found in the British Isles. The carrion crow is completely \ Indefatigable
black and is common in England and southern Scotland. h
Aopen atham
: -
The hooded crow is black with a grey back and belly and is Albemarle
Hh
\
found in the north of Scotland. Hybrids formed from the
mating of carrion and hooded crows occupy a narrow S Charles ta), /
< =< NEAR wt & Hood
region extending across central Scotland (fig 25.15). These =a *Ndisiributioniof
distribution of
hybrids have reduced fertility and serve as an efficient Camarhyncus psittacula Camarhyncus pauper
reproductive barrier to gene flow between the populations
of the carrion and hooded crows. on Albemarle Island are approximately equal, but on
In time, selection against cross-breeding may occur, Charles Island C. psittacula has a longer beak. This
leading to speciation. Since such speciation occurs finally in difference is significant enough to ensure that the two
the same geographical area, this is called sympatric (sym, species, which feed on different foods, appear unattractive
together; patria, native land) speciation. to each other during the breeding season. In this way the
Sympatric speciation does not involve geographical species remain distinct and are able to coexist.
separation of populations at the time at which genetic
isolation occurs. It requires the development of some form 25.8.4 Ring species
of reproductive isolating mechanism which has arisen by This is a special form of sympatric speciation
selection within a geographically confined area. This may
which occurs at the point where two populations at the
be structural, physiological, behavioural or genetic.
extremes of acline meet up and inhabit the same area, thus
Sympatric speciation is more commonly thought of as
‘closing’ the ring. For example, gulls of the genus Larus
providing an explanatory mechanism of how closely related
form a continuous population between latitudes 50-80 °N,
species, which probably arose from a common ancestor by
encircling the North Pole. A ring of ten recognisable races
temporary isolation, can coexist as separate species within
or subspecies exist which principally differ in size and in the
the same geographical area. For example, in the Galapagos
colour of their legs, back and wings. Gene flow occurs
archipelago the finch Camarhyncus pauper is found only on
freely between all races except at the point where the ‘ends
Charles Island, where it coexists with a related form C.
of the ring’ meet at the British Isles. Here, at the extremes
psittacula which is widely distributed throughout the
of the geographical range, the gulls behave as distinct
central islands (fig 25.16). The finch species appear to
species, that is the herring gull (Larus argentatus) and the
choose their mates on the basis of beak size. The range of
lesser black-backed gull (L. fuscus). These have a different
beak sizes of C. pauper on Charles Island and C. psittacula
appearance, different tone of call, different migratory
“J
HOOPED CROW
0 species
patterns and rarely interbreed. Selection against cross-
breeding is said to occur sympatrically.
Sympatric speciation without geographical isolation in
sexually reproducing species is unlikely. However, in
ZONE aaOF HYBRIDISATION species
asexually reproducing organisms, including vegetatively
=
= propagated higher plants, a single mutant so different from
=
= its parent population as to be genetically isolated could give
species
B
rise to a new species sympatrically. An example is
CARRION CROW
polyploidy in Spartina (section 23.9.2).
Fig 25.15 Hybrid barrier as a means of preventing gene flow

_ Ten subspecies of the Larus argen- _
between two populations. The maintenance of the two crow tatus-fuscus population form a continuous ring |
species is shown to be due to the existence of a zone of extending from the British Isles through Scandinavia, _
hybridisation extending across Scotland as shown in (a). The _ Russia, Siberia, across the Bering Straits, through _
existence of hybrid barriers between adjacent populations is Alaska and Canada and backto the British Isles. If _
common and functions as follows. Where the geographical the subspecies inhabiting the Bering Straits and |
ranges of A and B overlap, mating produces a hybrid with
lowered fertility. A will interbreed freely with AB and AB with B Alaska was eliminated what predicted effects might —
but the existence of AB prevents free interbreeding of A and this have on the population? - .)
B populations
902
25.9 interspecific hybridisation
This is a form of sympatric Speciation which
occurs when a new species is produced by the cross
ing of
individuals from two unrelated species. Fertile
hybrids
usually only appear in cases of interspecific hybri
disation
as a result of a form of chromosome mutation
known
as allopolyploidy (section 23.9.2). An example of
this was
demonstrated by Karpechenko in the case of hybri
ds
formed between the cabbage and the radish. The
genetic
changes involved in this hybridisation are show
n in
fig 25.17.
cabbage x radish
(Brassica oleracea) (Raphanus sativus)
2n = 18 2n= 18
Meiosis
Gametes n=9 n=9
F, hybrids 2n = 18
During meiosis in the F, hybrids chromosomes from each parent cannot
pair
together to form homologous chromosomes. The F, hybrids are therefore
sterile. Occasionally non-disjunction ofthe F, hybrids produces gametes
with
the diploid set of chromosomes (2n = 18).
Gametes 2n= 18 x 2n=18
F, hybrids 4n = 36 (tetraploid)
(Rhaphanobrassica)
The F, hybrids are fertile. Homologous pairing can occur in meiosis as two
sets of parental chromosomes are present. Diploid gametes (2n = 18), are
produced which possess 9 chromosomes from the parental cabbage and 9
chromosomes from the parental radish.
Fig 25.17 Stages involved in the hybridisation of the

cabbage and the radish
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Answers and discussion
Chapter 14 14.14 TP = y + OP
= —950 kPa + 1400 kPa
14.1 (a) Solution B_ (b) Solution A (c) From B to = 450 kPa.
A (d) Note that different beetroots may have different values of OP
Solution A (e) (ii) — 1000 kPa. If you find this confus
ing and w.
remember — 1000 is nearer zero than —2000. Zero
is the
maximum, or highest, w that can occur. (f) Soluti 14.15 (a) The cells of the intact scape are turgid and their walls
on B
(g) The higher the OP of a solution the lower the yw (in are therefore tending to expand with turgor pressure. The
practice —OP = yw for a solution at atmospheric pressure). thick walls of the epidermal cells are less capable of stretching
14.2 The hypertonic solution. Remember that the cell wall is than the thin walls of the cortex cells and therefore exert a
freely permeable to solutions. restraining influence on expansion of the cortical cells. The
14.3 Zero. The protoplast is not exerting pressure against the cell latter are under compression. Cutting the epidermis removes
wall. the restraint, each cortical cell expands slightly, and there is
an overall increase in volume of the cortex which causes the
14.4 Prokaryotes (the bacteria and blue-green algae) and fungi.
strip to curve outwards.
Although prokaryotes lack vacuoles, they have the same
(b) Distilled water has a higher water potential than the scape
Osmotic properties as plant cells and fungi.
cells. Water therefore enters the tissue from the distilled
14.5 At the start y™ = TP — OP, wy" = —OP = -1200 kPa; water by osmosis, inflating the cortical cells even further and
TP = 0 because the cell is at incipient plasmolysis; OP = causing outward curvature.
2000 kPa; so p“" = 0 kPa —2000 kPa = —2000 kPa. At (c) The concentrated sucrose solution has a lower water
the start, wp of the solution is greater than w of the cell. Water potential than the scape cells. Water therefore leaves the
therefore moves from the solution into the cell by osmosis. tissue by osmosis, causing greater shrinkage of the cortical
The only term that changes significantly as water enters the cells than the epidermal cells and a bending inwards of the
cell is TP. As water enters, TP, and hence yw, increases until tissue.
it prevents any further net entry of water. At this point p of (d) The dilute sucrose solution must have the same water
the cell equals w of the solution, and the cell is turgid. potential as the scape cells. There is therefore no net gain or
Therefore, at equilibrium py“ = —1200kPa. y = TP — OP loss of water by solution or tissue.
so TP = yp + OP = —1200 kPa + 2000 kPa = 800 kPa. (e) Water potential. An outline of the experiment is as
14.6 OP“ = 1 100 kPa; p pure water = 0; p“" = Oat equilibrium. follows.
TP = y + OP = 0 kPa + 1100 kPa = 1100 kPa. Prepare a dilution series of sucrose solutions from 1 M to
14.7 (a) y™" at transfer = y pure water = 0, p sucrose solution distilled water (such as distilled water, 0.2 m, 0.4 M, 0.6 M,
= —800 kPa, so the difference in wy between cell and 0.8 mM and 1.0 M). The typical curvature of freshly cut
external solution = 800 kPa. dandelion scapes should be recorded by drawing and then
(b) Water would leave the cell (from higher to lower yw). two pieces of scape placed in each solution in separate
(c) TP would decrease. labelled petri dishes (two pieces are preferred so that an
average can be obtained). Observe and accurately record
14.8 (a) Cell B (b) From cell B to cell A curvatures (such as by drawing) after equilibrium has been
(c) The cells must have the same y at equilibrium and this will reached (about 30 min). The solution which induces no
be the average of the two initial ws, that is — 1000 kPa.
change in curvature has the same wp as the average dandelion
(d) Cell A at equilibrium: TP = py + OP scape cell immediately after the cut was made.
— 1000 kPa + 2000 kPa
1000 kPa 14.16 Outlines of two suitable experiments are as follows.
Cell B at equilibrium: TP w + OP Effect of temperature. Cut cubes of fresh beetroot, wash to
— 1000 kPa + 1400 kPa remove the red pigment from broken cells, and place in
= 400 kPa. beakers of water at different temperatures over a range, say,
from 20-100 °C. The appearance of red pigment in the water
14.9 1060 kPa. For intermediate values between those shown in
would indicate destruction of the differential permeability of
table 14.4, plot a graph of molarity of sucrose solution against
the tonoplast (vacuole membrane) and plasma membrane,
osmotic pressure.
attended by diffusion of the pigment from the cell sap to the
14.10 Average OP of beetroot cells is about 1400 kPa. water. The time taken for the appearance of a standard
14.11 w beetroot is about —940 kPa. amount of pigment would give an indication of the rapidity of
14.12 A more accurate result can be obtained by taking the mean breakdown of membrane structure. The colour could be
value of two or more replicates. Some indication of the measured in a colorimeter or simply by using the eye.
variation that can be expected between strips is given in table Effect ofethanol. Method as above, using a range of ethanol
14.6. concentrations instead of a range of temperatures.
14.13 To prevent evaporation of water, with subsequent increase 14.17 (a) Leaves contain a very large number of stomata for
in concentration of sucrose solutions, and possible drying up gaseous exchange and there is little resistance to movement
of beetroot strips. of water vapour through these pores.
905
s
Table 14.22(ans)
(b) Leaves have a large surface area (for trapping sunlight and
exchanging gases). The greater the surface area, the greater ES
—
will be the loss of water by transpiration. Dicots Monocots

14.18 Light intensity increases as the Sun rises, reaching a Piliferous layer quickly lost Piliferous layer remains
maximum at midday when the Sun attains its highest point in Many protoxylem groups
the sky. Air temperature rises similarly, but it takes about Few protoxylem groups (few
arms of metaxylem) 2-8, (polyarch)
two hours for the heating effect of the Sun to be reflected in a commonly four (tetrarch) or
rise in air temperature (mainly because the soil has to heat up 5 (pentarch)
first and then radiate heat to the air). The initial rise in Pith not common. Xylem Central pith. Xylem forms
transpiration rate between 3 am and 6 am, before air forms a solid cylinder at the rods around the pith
temperature rises, is due to opening of the stomata in the centre of the root
light. From 6 am onwards transpiration rate is closely Cambium often present, Cambium rarely present;
therefore no secondary
correlated with temperature for reasons explained in the making secondary thickening
thickening
text. It is not closely correlated with light intensity, possible
eee OS_N—
presumably because the stomata are now fully open and any
further increase in light intensity has no effect.
(b) There are two distinct phases of uptake. In the initial rapid
During the afternoon, light intensity decreases as the Sun
phase, K” diffuses into the free spaces of the root. It enters
sinks, followed by a drop in temperature with the same lag of
through the cell walls of the piliferous layer, which are
about two hours. Transpiration rate decreases both as a
exposed to the solution, and diffuses through the apoplasm,
result of decreasing temperature and decreasing light
possibly entering the intercellular spaces where these contain
intensity, but it is much more closely correlated with a
solution. The results show that this phase is more or less
decrease in the latter, probably because this induces stomatal
independent of temperature and therefore not dependent on
closure. By about 7.30 pm it is dark and the stomata are
metabolism (enzyme-controlled reactions would proceed
probably closed. Any remaining transpiration is probably
faster at 25°C than 0 °C). This first phase is a passive process.
cuticular and still influenced by temperature.
The second phase is temperature-dependent, and does not
14.19 See table 14.19(ans). occur at 0 °C when the rate of metabolism is very low. This
Table 14.19(ans) suggests that the process is dependent on metabolism, and its
inhibition by KCN indicates that it is dependent on
Dicots Monocots respiration. The second phase is therefore an active transport
Vascular bundles arranged in Vascular bundles scattered
across cell membranes into cells.
a ring 14.25 Rise in respiratory rate is accompanied by arise in KCl
Endodermis (starch sheath) Endodermis absent uptake. Once KCI is available, it is therefore apparently
present taken up by active transport, the energy being supplied by an
Separate pith and cortex Ground tissue not split into increased respiratory rate.
pith and cortex
Pericycle forms a cap to Pericycle surrounds vascular 14.26 KCN inhibits respiration and thence active transport of
vascular bundle bundle KCl into the carrot discs.
Metaxylem has many vessels Metaxylem has few vessels 14.27 Much of the phosphate inside the root was in the free space
(commonly two or three) and could therefore diffuse out to the water outside,
Protoxylem present Most protoxylem breaks reversing passive uptake.
down to leave a lysigenous
canal
14.28 No. The endodermis is a barrier to movement of water and
solutes through the apoplast pathway (see section 14.5.2,
Cambium present and No cambium, therefore no
secondary thickening may secondary thickening (very apoplast pathway).
occur later (some herbaceous few exceptions where 14.29 Autoradiography reveals the location of the ion in thin
plants, such as Helianthus, monocotyledon stems develop sections. Treat one plant with an inhibitor of active transport
develop secondary vascular a tree-like form)
bundles; perennials develop
(such as low temperature or KCN) and have an untreated
wood, rays, annual rings, control plant; allow them both to take up the radioactive ion.
cork) In the treated plant ions will move only passively by way of
the cell walls. Autoradiography should show that the
radioactive ion tends to penetrate the root only as far as the
14.20 (a) Hollow cylinder
(b) Solid rod/cylinder providing support endodermis, whereas the control should show much greater
(c) Solid rod/cylinder providing support movement of ions to the tissue inside the endodermis.
(d) Solid cylinder 14.30 (a) See table 14.30(ans).
14.21 (1) Long tubes formed by joining of neighbouring cells, Table 14.30(ans).
with breakdown of cross-walls between them.
(2) No living contents, so less resistance to flow.
(3) Tubes have high tensile strength so do not collapse. Plant part Percentage Percentage
(4) Fine tubes are necessary to prevent water columns from distribution when distribution when
physically collapsing. upper leaf treated lower leaf treated
14.22 See table 14.22(ans). apical region of shoot 4.8 3.6
14.23 wy soil solution > root hair cell > cell C > cell B> cell A> C-treated leaf 48.4 53.9
xylem sap other leaves 1.0 0.8
14.24 (a) There is a rapid initial uptake of potassium (K’) at both stem 3) De)
temperatures (during the first 10-20 min). After 20 min there pod 38.7 23.4
is a continuous gradual uptake of K” at 25 °C but no further roots 350) 12.8
uptake at 0 °C. Uptake at 25 °C is inhibited by KCN.
906
Similarities in “C export from upper and lower
leaves. return to normal of heartbeat and blood pressure.
Similar proportions of assimilates are exported by
both leaves 14.37 In tissues which are respiring actively the partial pressure of
(compare results for ‘“C-treated leaf’) and simila
r propor- carbon dioxide will be high. This leads to a reduced pH and to
tions pass to the other leaves. The major destinatio
n of the the displacement of the oxygen dissociation curve to the
assimilates in both cases is the pod.
Differences in “C export from upper and lower leaves. right. Analysis of fig 14.64 shows that as this occurs it
The pattern of export from a given leaf is facilitates the delivery of increased quantities of oxygen from
affected by its the blood to the cells which can be used by respiratory
position on the plant. The upper leaf expor
ts a higher processes for the production of energy. Looked at another
proportion of its assimilates to the pod and apical
region than way, haemoglobin takes up oxygen less readily initially but
the lower leaf, which correspondingly exports
a greater when saturated releases it quickly.
proportion to the roots.
(b) Once a pod is formed, it becomes a sink of consi Increased metabolic activity increases the temperature ina
derable part of the body. This produces a reduction in the affinity of
importance for carbon compounds, particularly
from the oxygen for haemoglobin and an increased dissociation of
leaves inits proximity. There are two other important
sinks in oxygen. Thus the dissociation curve is again shifted to the
the mature plant, namely the apical region of the
shoot and right. This is physiologically advantageous as more oxygen is
the roots. The lower leaves export mainly to the roots
and the delivered to the active regions. The oxygen dissociation
upper leaves mainly to the apex.
curve is not exactly the same for all animals. For example,
14.31 0.72 s. The answer is obtained as follows:
compared with Man, the curve for small mammals is
100 cm = 1000 mm = 1000000 um = 10° um displaced to the right. Small mammals possess a much higher
therefore sucrose moves at 10° um h',
metabolic rate than Man and therefore it is appropriate that
= 1 um in 1/10° h, oxygen should be released much more readily.
= 200 um in 200/10°h = 2/10h = (2 x 3 600)/10's =2 x 0.36s
14.38 The position of the curve of the foetus relative to that of its
= 0.72 s.
mother means that its blood has a greater affinity for oxygen
14.32 2500 sieve plates per metre: than the maternal blood. This has to be so, as the foetus must
1m = 10° ym, obtain all of its oxygen from its mother’s blood at the
400 um = 4 x 10° um, placenta. So, at any given partial pressure of oxygen, the
10/(4
6 ee 4
x 10°) =an 10/4 = 2500. foetal blood will take up oxygen from the maternal blood and
14.33 Oxygenated blood can be delivered to the tissues rapidly will always be more saturated with oxygen than the maternal
enough to satisfy the increased metabolic demands of the blood.
body. Various organs such as the kidney and the capillaries 14.39 This means that the blood has a high affinity for oxygen and
rely on a high blood hydrostatic pressure for effective and that it is able to combine with it at the low oxygen tensions
efficient functioning. experienced at high altitude. This is another good example of
14.34 This means that oxygenated blood of the systemic physiological adaptation.
circulation reaches the body capillaries at a much higher 14.40 (1) Carboxyhaemoglobin reaches the lungs and takes up
pressure. This is essential for the efficient function of organs oxygen and forms oxyhaemoglobin,
and tissue fluid formation and permits active chemical (2) Oxyhaemoglobin is a weaker base than carboxyhaemo-
processes and a high body temperature to be maintained. It is globin and releases hydrogen ions.
essential that a much lower pressure is developed in the (3) Hydrogen ions combine with hydrogencarbonate ions in
pulmonary artery in order to prevent rupture of the delicate the erythrocyte so forming carbonic acid.
pulmonary capillaries. (4) Carbonic acid dissociates into carbon dioxide and water.
14.35 Local vasodilation in the wounded area enables more (5) As a result of the loss of hydrogencarbonate ions from the
blood carrying oxygen and nutrients to arrive there and speed erythrocyte, further hydrogencarbonate ions diffuse into the
up the process of repair and replacement. Increased body erythrocyte from the plasma.
blood pressure prepares the body of the animal to respond to (6) More carbonic acid is formed which dissociates into more
any further stress more readily and efficiently. carbon dioxide and water.
14.36 Before the race. Adrenaline is secreted in anticipation of (7) Carbon dioxide diffuses out of the erythrocyte and is
the race. This stimulates vasoconstriction throughout the eventually excreted from the body via the lungs.
body in all but the most vital organs. Hence blood pressure is
raised. Heart rate is also increased. Extra blood is passed to
the general circulation from the spleen.
Chapter 15
During the race. Increased metabolic activity takes place 15.1 Locomotion is primarily associated with the need to search
during the race, especially in the skeletal muscles. Increased for food (and is closely associated with the development of a
carbon dioxide levels in these regions promote local nervous system). Green plants are autotrophic, that is make
vasodilation. The increased body temperature further en- their own organic requirements, so do not need to search for
hances vasodilation. However the general increase in carbon food.
dioxide level in blood is noted by the chemoreceptors of the
15.2 See table 15.2(ans) (next page).
aorta and carotid bodies which in turn stimulate the
vasomotor centre to promote vasoconstriction. This in- 15.3. Growth could be inhibited on the contact side, or stimulated
creases blood pressure and therefore speeds up blood flow. on the opposite side, or a combination of these effects might
Heart rate is also increased and a more complete emptying of occur. In fact, growth is slowed down (cells become less
the ventricles occurs. Towards the end of the race the muscles elongated) on the contact side and growth on the opposite
will be respiring anaerobically and producing lactic acid side is stimulated 40-200 fold.
(section 11.3.8). Strong contractions of the muscles knead 15.4 Various methods are possible. A simple experiment is
the veins and promote faster venous return to the heart. illustrated in fig 15.4(ans).
Recovery. The oxygen debt is paid off and lactic acid removed 15.5 (a) Spirogyra (or any other filamentous green alga).
from the blood system. Tissues subside in activity and the (b) The bacteria are aerobic and positively aerotactic.
carbon dioxide level decreases. Consequently there is a Therefore they swim towards oxygen along a gradient from
907
coleoptile mica
low oxygen concentration to high oxygen concentration. The
highest oxygen concentrations are around the edges of the
Ly
cover-slip, where oxygen is diffusing into the water from the
atmosphere, and adjacent to the algal filament where oxygen
is being released as a waste product of photosynthesis.
Table 15.2(ans).
SSS ooo
Example Advantage
Shoots and coleoptiles Leaves exposed to the light which is the

positively phototropic source of energy for photosynthesis hormone stimulates growth only on side not
CONTROL -
Roots negatively Exposed roots more likely to grow normal growth interrupted by mica
phototropic towards soil or equivalent suitable
substrate Fig 15.8(ans) Repetition of Boysen-Jensen’s experiments in
Shoots and coleoptiles Shoots of germinating seeds will grow uniform light. Three experiments are shown; treatment left,
negatively geotropic | upwards through soil towards light result right, in each case
Roots positively Roots penetrate soil
geotropic
Rhizomes, runners Helps plants colonise new areas of soil (c) Leave the slide in the dark for about 30 min and
diageotropic re-examine. All the bacteria should now be around the edges
Dicotyledonous leaves Flat surface of leaf will gain maximum of the cover-slip because the alga cannot photosynthesise in
diageotropic exposure to sunlight (at right-angles to the dark.
incident radiation) 15.6 (a) The stimulus of light is detected by the coleoptile tip.
Lateral roots Large volume of soil exploited and the Some kind of signal is transmitted from the tip (the receptor)
plagiogeotropic arrangement of roots provides support to the region behind the tip (the effector).
(similar to guy-ropes supporting a (b) Experiment c was a check on the result from experiment b
tent) which could have been the result of injury to the coleoptile.
Branches Larger volume of space occupied for 15.7 Further evidence of the existence of a signal, presumably a
plagiogeotropic exploitation of light chemical transmitter substance (hormone), has been
Hyphae positively Grow towards food obtained. It cannot pass through an impermeable barrier. It
chemotropic moves mainly down the shaded side of the coleoptile. In
Pollen tubes positively Grow towards ovule, where fertilisa- experiment b mica prevented this movement. Light therefore
chemotropic tion takes place either, inhibits production of the hormone, causes its
Roots and pollen tubes Water essential for all living inactivation (stimulates its breakdown) or causes it to be
positively processes redistributed laterally.
hydrotropic 15.8 See fig 15.8(ans).
Tendrils positively Essential for their function of support 15.9 The coleoptile tip produces a chemical which diffuses into
haptotropic the agar. It can stimulate growth in the region behind the tip
Sundew tentacles Enables plant to imprison insects and restores normal growth (experiment a). There is little or
positively haptotropic which walk over the tentacles (section no lateral transmission of the chemical (experiment b) under
O72) conditions of uniform illumination or darkness.
Pollen tubes Another mechanism ensuring that 15.10 The coleoptile would have grown to the left.
negatively aerotropic initial growth of the pollen tube is 15.11 A100 ppm B10ppm Clppm D0.1 ppm
towards the tissue of the style (away E 0.01 ppm_ F zero
from air) 15.12 A relatively high auxin concentration stimulates growth of
coleoptiles (or shoots) but inhibits growth of roots. This
supports the conclusions drawn from experiment 15.1.
semi-transparent cover, such as tissue paper of 15.13 See section 16.5.6.
increasing thickness; remove for examination with 15.14 (a) Abscisic acid can be transported away from root tips,
binocular microscope after about 30 min undergo lateral transport in root tissues in response to
Sans
LIGHT gravity, and inhibit growth.
(b) IAA is probably not involved in the geotropic response of
maize since it is apparently not transported away from the
root tip.
15.15 (a) starch
(b) maltose
blackened dish
(c) maltase
(d) The main food reserve of cereal seeds is starch, stored in
the endosperm.
algae concentrated at 15.16 Storage proteins are digested (hydrolysed) to provide
preferred light intensity amino acids, the basic units of proteins. These are reassemb-
led to produce enzymes (which are always proteins), such as
Fig 15.4(ans) Experiment to demonsirate preferred light a-amylase, which are then used to digest the food stores of
intensity of Euglena or Chlamydomonas the endosperm.
908
15.17 The amylase activity could be associated
with micro-
Organisms present on the fingers or with saliva which has —_ —_ — — _—-— —=—-s
been transferred from mouth to fingers. Note the impor —_——_

- Zs “far-red light
tance, therefore, of not handling the seeds after their surfac
e
sterilisation in this kind of experiment.
15.18 Incubate seeds with radioactive (C-labelled) amino
acids.
This results in production of labelled amylase. Altern
atively,
incubation of seeds with inhibitors of protein synthesis (such
as cycloheximide) prevents synthesis of amylase and
no
amylase activity is then recorded.
15.19 Dissection of the seeds into aleurone and non-al
eurone Floral
stage
portions should show that the initial appearance of labelled red light
amylase is in the aleurone layer. Alternatively, separa
te
incubation of endosperm with aleurone layers and endo-
sperm without aleurone layer, with starch-gibberellin agar
would result in amylase production only in the former
(difficult to do in practice).
15.20 One of the best bioassays for gibberellin (quick, reliable
0 10 20 30 40 50
and sensitive) involves incubating embryo halves of barley
Duration of light/s
grains with the substance being assayed. After two days the
amount of reducing sugar present is proportional to the Fig 15.26(ans) Effects of red light and red/far-red light
amount of gibberellin present. interruptions of long night on flowering of cocklebur
15.21 (a) The amino acid is retained by the young leaf and does
not move very far from the point of application. In the old
leaf some of it is exported via the veins and midrib. 15.25 See chapter 9.
(b) The young leaf would use the amino acid to make protein 15.26 The graph is shown in fig 15.26(ans).
in growth. The old leaf is no longer growing and so is The opposite effects of red and far-red light are demon-
exporting nutrients to other parts of the plant such as roots strated. Red light exposure of 30s, at the intensity used in the
and young leaves. experiment, completely nullifies the inductive effect of along
(c) Amino acids are retained by, or move towards, tissues night. The effectiveness of red light increases with time of
treated with kinetin. (The reasons for this are unknown, but exposure up to 30s. The red light effect is reversed by far-red
presumably connected with the maintenance or stimulation light, although a longer exposure (50 s) was needed to
of normal cell activity by kinetin.) completely reverse the effect. These results suggest that
phytochrome is the photoreceptor involved.
15.27 There are several possible methods. Fig 15.27(ans)
lanolin paste lanolin paste containing [AA illustrates one simple solution. Boxes represent light-proof
covers, used as appropriate to give short days.
apical bud
short days ——»> flowers long days ——» flowers

lateral bud
apical bud
Normal growth Remove apical Apical dominance
bud — apical restored by [AA leaves —
short days
dominance broken
Fig 15.23(ans) Experiment to show role of IAA in apical

dominance
15.22 One solution would be to take a plant where applied

gibberellin is known to affect stem growth and remove its
long days ——» no flowers short days —» no flowers
source of auxin by removing the shoot apex. Gibberellin
should then prove ineffective. It is important to demonstrate
that the response can be restored by addition of auxin (such
as IAA in lanolin paste) as injury might be the reason for lack
of response to gibberellins, or another chemical might be
involved. Such experiments do demonstrate a total depen- long days
dence on auxin.
15.23 (a) Auxin (IAA)
(b) See fig 15.23(ans).
15.24 Small leaves offer less resistance to passage through the soil
(leaves of grasses remain inside the coleoptile). The hooked
plumule of dicotyledonous plants protects the delicate apical Fig 15.27(ans) Experiment to determine whether leaves or
meristem from soil particles. Elongated internodes ensure floral apex are sensitive to the photoperiod that stimulates
the maximum chance of reaching light. flowering
909
ty
15.28 Lateral bud inhibition or apical dominance is largely intensity activating the cones. At night the light intensi
By lookin g slightl y to
controlled by auxins. (See apical dominance, section 1553533) would be too low to activate the cones.
one side of the object the reflected light from it will not strike
the fovea but a point on the retina to the side of it where there
Chapter 16 are rods. At night these will be activated by the low light
intensity and an image will be produced in the brain.
16.1 (a) A steep concentration gradient of Na’ ions exists 16.11 The object will appear yellow. Each retina will distinguish
between the outside and inside of the axon and Na’ ions one colour only. In one eye, green cones will be stimulated by
rapidly diffuse down this gradient. light of 530 nm and, in the other, red cones will be stimulated
(b) The relatively high negative potential within the axon by light of 620 nm. Mixing will occur in the brain due to equal
encourages the inward movement of the positively charged stimulation by these colours and the object will appear to be
Na’ ions. the colour of the average of the combined wavelengths,
16.2 If the outflow of positive K” ions from the axon balanced the 530 +620
that is nm = 575nm, which corresponds to yellow.
inflow of positive Na’ ions into the axon there would be no 2
change, or perhaps only a slight decrease, in the resting
potential.Such a slight change would be insufficient to reach
the threshold required to produce an action potential.
Chapter 17
16.3 (a) Normal sea water 17.1 The femur is basically a hollow bone. When a compression
(b) One-half sea water force is exerted on one side of the bone, the other side is
(c) One-third sea water subjected to tension. Along the central axis of the bone the
The amplitude of the action potential spike in (a) and the forces diminish and are neutralised. The material in the
depolarised membrane potentials shown in (b) and (c) is centre of the bone consequently does not need to contribute
determined by the number of Na’ ions entering the axon to its strength. Reduction in weight of the bone due to the
from the extracellular fluid. The solutions in which (a), (b) absence of bone along its central axis is advantageous to the
and (c) were recorded contained progressively fewer Na animal as it lightens the weight of the femur without reducing
ions. its mechanical efficiency. The perimeter of the femur is
16.4 Sodium ions are pumped out of the axon and potassium ions composed of compact bone which resists the tension and
are pumped in. The movements of the ions are linked compressional forces. Spongy bone at the head of the femur
(coupled) and mediated by the cation pump. is a meshwork of interconnecting bony struts. They maintain
16.5 The longitudinal resistance of axoplasm decreases with the rigidity of the bone but with the minimum of weight.
increasing diameter of the axon. As the resistance decreases, Cartilage acts as a cushion between two articulating bones.
the length of the membrane influenced by the local circuit Its matrix can be deformed by compression but will return to
increases and this lengthens the distance between adjacent its original shape because it possesses good powers of
depolarisations and leads to an increase in conduction extensibility. The cartilage also reduces friction between the
velocity. smooth, moving articular surfaces.
The tendons consist of inextensible, white fibrous tissue
16.6 The frog is a cold-blooded (poikilothermic) organism, active
and attach the muscles to the femur. The pull of the muscle is
within the temperature range 4—25 °C, whereas the cat, being
concentrated over a small area. Tendon organs operate to
warm-blooded (homeothermic), maintains a constant
temperature of 35 °C. This increase in temperature increases prevent muscle rupture if the muscle is suddenly subjected to
the speed of conduction of the nerve impulses by a factor of a heavy load.
three. Ligaments are also composed of inextensible, white fibrous
tissue and connect the femur to bones articulating with it at
16.7 Graph (a) indicates that the frequency of impulses in the joints. They confine the movement of the components of
sensory neurone is directly related to the stimulus applied to
each joint to a specific direction and therefore promote the
the muscle spindle. It is known that the stimulus produces a
depolarisation known as the generator potential and, as efficiency of its operation. Ligaments also strengthen the
shown in graph (b), the magnitude of this potential is directly joint.
related to the frequency of sensory impulses. It may be 17.2 (a) It allows free movement of the rib cage of the mammal.
concluded from these data that increasing stimuli produce (b) The flexible suspension enables the animal to withstand
increasing generator potentials whose amplitudes are direct- the shock sustained by the forelimbs when it lands at the end
ly related to the frequency of impulses in the sensory of a jump.
neurone. (c) The forelimbs possess a wide range of movement, which is
16.8 The path taken by light as it passes through the eye is as useful for such activities as climbing, cleaning the face,
follows: conjunctiva+cornea—aqueous humour— lens— manipulating food and digging.
vitreous humour—retina. 17.3 A band remains the same length; H band becomes shorter; I
16.9 Light from an object falling onto several rods which are band becomes shorter.
linked to the brain by separate neurones may not have 17.4 The myosin filaments (constituting the A band) remain the
sufficient energy to produce a propagated action potential in same length whilst actin filaments slide over them towards
each neurone and therefore the light may not be detected. If, each other. This effectively shortens the H and I bands.
however, the same light falls on three rods which are linked 17.5 (a) The central non-contractile portion of the muscle spindle
to the same neurone supplying the brain, the separate is stretched. Nerve impulses are fired from the annulo-spiral
generator potentials produced by the rods would summate nerve endings and pass to the central nervous system.
and produce a propagated action potential which would be Impulses are propagated from the central nervous system via
registered in the brain as light. efferent neurones to the extrafusal fibres causing them to
16.10 When looking directly at an object, light reflected from it contract. This is the stretch reflex.
passes along the optical axis of the eye and strikes the retina (b) The muscle spindle is no longer stretched and the number
at the fovea which contains cones only. During daylight this of impulses fired from the central non-contractile region of
will produce a detailed image in the brain due to the high light the spindle is diminished.
910
(c) Impulses are fired from the tendon organs which
reach are low and the plant cannot lose latent heat and reduce its
inhibitory neurones in the spinal cord. When these
are temperature.
activated this causes a reflex inhibition of contraction of the
muscle that is being stretched. Therefore active resistance 18.3 19.6%
to
stretching is decreased. This mechanism offers a form 18.4 During this period the subject was allowed to equilibrate
of
protection to the muscles, preventing possible rupture when with his surroundings.
they are suddenly overloaded. 18.5 There is a direct relationship between these two variables
(d) The central region of the spindle is stretched and fires which suggests that the rate of sweating is controlled by
impulses to the spinal cord. Motor neurones are stimulated activity of the hypothalamus.
to
carry impulses to the extrafusal fibres of the muscle causin 18.6 The direct relationship between skin temperature and
g
contraction. When contraction in the intrafusal fibres evaporation during the first 20 min established that an
is
matched by an equal contraction of the extrafusal fibres, the equilibrium exists between the two. As the evaporation rate
central region of the spindle is stretched no further and astate falls, due to the action of the hypothalamus in response to
of equilibrium is reached. This mechanism is very important the ingestion of iced water, latent heat of evaporation is not
as it sets the tone of muscles. It ensures that the muscle is not being lost from the skin and this accounts for the observed
completely slack, and by increasing the stretching of the rise in skin temperature.
spindle so increases the muscle’s response. If the muscle was 18.7 ‘Fever’ is due to the resetting of the hypothalamic
slack when a heavy load was applied, the muscle could suffer
‘thermostat’ at a higher temperature. Until the core
considerable damage. temperature rises to that temperature there is a discrepancy
17.6 It allows greater movement of Ca’* ions needed for muscle between ‘normal’ body temperature and the cold conditions.
contraction. In these conditions the body responds by shivering and the
17.7 Synchronous. This type of muscle has much more sarcoplas- body continues to feel cold until the core temperature
mic reticulum because it requires more nérvous impulses to reaches the temperature of the hypothalamic thermostat.
operate it, and each nerve impulse depends on the release of
Ca’* ions by the sarcoplasmic reticulum.
17.8 (a) Streamlined shape
(6) Smooth surface — scales overlap each other in an
Chapter 20
appropriate direction 20.1 (a) The true statement is (ii), ‘Gametes are always haploid’.
— mucus/oily covering thus reducing Even this is not always true since polyploid parent cells would
friction give rise to gamete cells with more than the haploid number
(c) Various types of fin to promote forward propulsion and of chromosomes; for example the gametes of hexaploid
stability during swimming wheat plants would be triploid.
(d) Highly muscular body (b) Exceptions to statements (i), (iii), (iv) and (v):
(e) Lateral line (table 4.12) (i) Asexual reproduction in plants with alternation of
(f) Swim bladder in bony fishes generations involves meiosis, so the gametophytes
(g) Highly coordinated neuromuscular activity produced show variation.
17.9 (a) Large, powerful flight muscles (iii) Gametes are produced by mitosis in life cycles A and
(b) Streamlined body shape D (fig 20.1).
(c) Sharp narrow wings, designed for low drag (iv) Meiosis does always produce haploid cells (except in
(d) Large wings possess mostly primary feathers (for forward polyploid organisms). Mitosis can produce haploid cells,
propulsion) as in the growth of multicellular haploid organisms (such
(e) Fast wing beat (10 per second) as Spirogyra, life cycle A, and growth of gametophyte,
(f) Forward thrust achieved by both the downbeat and upbeat life cycle D, fig 20.1). It can also produce diploid cells as
of the wings. in the growth of multicellular diploid organisms.
17.10 This position increases the effective length of their limbs. (v) Mitosis can occur in haploid cells, as in the examples
Consequently each stride taken is longer and so propels the given in (iv) above.
body forward over a greater distance. Assuming that the 20.2 (a) If the parent plants that produced the pollen grains can
speed of movement of the limbs remains the same, the be identified, then certain deductions can be made about the
sprinter will therefore move forwards at a faster pace. climate that such plants would have grown in.
(b) Any interference by Man with the natural vegetation
would be reflected in the pollen record. For example, pollen
Chapter 18 of weed species and agricultural plants, such as wheat, would
indicate clearance of natural vegetation for agriculture.
18.1 Hyperventilation increases the tidal movements of air, Similarly, absence of pollen from trees in some areas would
carbon dioxide is expelled from the lungs and the carbon indicate forest clearance.
dioxide tension of the alveolar air decreases. The carbon 20.3 If a plant species is dioecious, half of its individuals do not
dioxide level of the blood comes to a new equilibrium with produce seeds. Also, there is a large wastage of pollen which
the alveolar air as it passes through the pulmonary is a disadvantage in terms of material and energy resources.
capillaries. This reduces the carbon dioxide tension of the 20.4 Separate sexes is more economic in animals than in plants
blood and increases the blood pH. The abnormal alkalinity of because there is less risk in transferring male gametes as a
the blood produces symptoms of dizziness and fainting and result of locomotion and behaviour patterns.
the inhibitory effect of lack of stimulation of the chemorecep- 20.5 3 (50%). Remembering that the pollen grain is haploid:
tors supplying the respiratory centres decreases the ventila- Parent plant genotype Possible pollen genotypes
tion rate. '
S,S, = in equal numbers
18.2 The rates of transpiration and evaporation are inversely 2
proportional to the level of atmospheric humidity. When S, pollen grains would be compatible with S,S, style tissue
humidity is high the rates of transpiration and evaporation S, pollen grains would be incompatible with §,S, style tissue
911
(b) Red light stimulates lettuce seed germination, but far-red
Note that neither S, nor S, pollen grains would be compatible
with the style of the parent plant (S,S,), so that self-
light inhibits it (section 15 .4.2). Seeds under a leaf canopy,
fertilisation is impossible. where the light will be enriched in far-red, might therefore be
inhibited from germinating until a break in the canopy
20.6 (a) The part of the bee’s body receiving most pollen will be ensures that they will not be too shaded for efficient
that which brushes against the anthers while the bee is taking photosynthesis and growth.
nectar. Thus pollination will generally occur between anthers
and stigmas at the same height within the flower, that is 21.6 At the onset of germination, food reserves in the barley
between pin-eyed and thrum-eyed flowers.
grain, principally starch, with some protein, are mobilised.
(b) It encourages outbreeding. Starch is converted to sugars, and proteins to amino acids,
and these are translocated to the embryo for use in growth.
20.7 The functions of the cell organelles suggest that the cells Therefore, endosperm dry mass decreases while embryo dry
manufacture materials for use within the cell. The raw mass increases.
materials for these processes come from the breakdown of At the same time there is an overall loss in dry mass during
materials entering the cell, using enzymes stored in the the first week. This is due to aerobic respiration, which
lysosomes. The synthesised products are packaged by the consumes sugar, in both endosperm and embryo (though toa
Golgi apparatus and stored for subsequent usage. greater extent in the latter). At about day 7 the first leaf
20.8 (a) Adding the numbers of corpora lutea and corpora emerges and starts to photosynthesise. The resulting increase
albicanti together shows the number of ovulations which in dry mass more than compensates for respiration losses so
have occurred, that is 67 in all. Assuming one oocyte was that a net increase in dry mass is observed. At the same time
released from each follicle per month this gives the the rate of growth of the embryo, now a seedling, increases.
reproductive age at 5 years 7 months. Hence the 22-year-old
21.7 (a) There is a gain in dry mass of 8.6 g, calculated as follows.
woman began ovulating at approximately the age of 163 ;
Mass of seeds = 51.2 g
years. = 51.2/2 = 25.6 g
Mass of fatty acid
(b) Of the 42000 follicles only 219 were primary oocytes,
M, fatty acid = 256
having a diameter over 100 um. Again, assuming one oocyte
Therefore 1 mole = 256 g, so 25.6 g = 0.1 mole
would be released per month, the potential number of years
From the equation,
for ovulation would be 18 years, 3 months.
0.1 mole fatty acid > 0.1 mole sugar + 0.5 mole water
(c) Five follicles contained two oocytes each, therefore + 0.4 mole carbon dioxide
potentially five pairs of twins might have been produced M. sugar = 342
which would not be identical twins.
Therefore 25.6 g fatty acid — 34.2 g sugar + water
(d) Non-disjunction occurring during anaphase I of meiosis. + carbon dioxide
20.9 Blood would flow in the reverse direction along the ductus Water is not included in the dry mass and carbon dioxide is
arteriosus. lost as a gas, therefore the gain in dry mass = (34.2 — 25.6) g
= 8.6g.
Chapter 21 (b) Respiration would result in a decrease in dry mass. In
21.1 The following conclusions may be drawn from these curves: reality there would still be an increase in dry mass.
(a) relative growth is greatest during embryological develop- (c) Volume of carbon dioxide evolved from the seeds =
ment, 8.96 dm* at STP, calculated as follows:
(b) rate of growth is greatest during infancy and adolescence, from the equation, 0.1 mole fatty acid — 0.4 mole carbon
(c) maximum growth, in terms of a particular parameter, such dioxide. 0.4 mole carbon dioxide occupies 0.4 x 22.4 dm’ at
as height, is greatest in the adult. STP = 8.96 dm’.
21.2 (a) There is loss of mass due to respiration of food reserves in (d) By hydrolysis, catalysed by a lipase. The other component
the seed. of the lipid is glycerol.
(b) Green leaves have grown and opened above the ground. (e) 51 carbon atoms (the lipid would be tripalmitin; the fatty
(c) Photosynthesis. Its rate must now be greater than acid is palmitic acid). Each lipid molecule comprises three
fatty acid molecules, each with 16 carbon atoms, plus one
respiration.
(d) This is due to dispersal of fruits and seeds. glycerol molecule with three carbon atoms.
(f) Sucrose or maltose.
21.3 Body mass averages about 33.5 kg from November to March
(g) Oxygen reaches the storage tissue by diffusion through the
at a time when the average daily intake is 320 kJ. Throughout
testa and micropyle.
this period the mean monthly temperature does not rise
above —10 °C. In order to maintain a constant blood 21.8 (a) The dominant food store is lipid, which comprises about
temperature of approximately 35 °C, despite a temperature 70% of the dry mass of the seeds before germination. By
difference of 45 °C, the husky, a homeotherm, must have an day 4 the mass of lipid is starting to decrease and the mass of
extremely high metabolic rate. This places a great demand on sugar to rise. Lipid is therefore being converted to sugar and
an adequate supply of food to provide the energy sources for translocated to the embyro. Note that no sugars can be
respiratory activity. Since the data show that the average formed by photosynthesis since germination occurs in
body mass throughout the winter period is below that of the darkness. At day 5 the RQ of the embryo = 1, indicating that
summer, this provides extra evidence of the metabolic the embryo is respiring the sugar derived from the lipids. At
demands imposed on maintaining a constant body tempera- the same time, the cotyledons (RQ = 0.4—0.5) are gaining
ture. energy from the conversion of lipid to sugar, and possibly
21.4 Small seeds have relatively small food reserves; it is
from oxidation of sugar and fatty acids.
therefore important that the growing shoot reaches light
quickly so that photosynthesis can start before the reserves C,,H,,0, + 130, > C,H,,0,, + 6CO, + 6H,O + energy
are exhausted. ricinoleic sucrose
acid (fatty
21.5 (a) Chlorophyll strongly absorbs red and blue light, but not acid derived
green and far-red light (see chlorophyll absorption spectrum from a lipid)
fig 9.9).
oi
RQ = 6/13 = 0.46 cells from A grown in '4N
Conversion of lipid to sugar takes place with an increase in cells grown ———.-S*’]?—7".
dry mass, so total dry mass of the seedlings increases up to 6 in SN DNA first generation second generation
or 7 days. Beyond this point, the lipid reserves are running
low, so rate of use of sugar starts to exceed the rate of
production. Net mass of sugar, and total mass of seedlings,
then starts to decrease. Sugar is used in respiration and in
anaerobic reactions. =e — '4N - containing DNA
(b) At day 11, the RQ of the whole seedlings would probably
be slightly less than 1.0. It is a combination of two reactions:
the main one is the oxidation of sugar in respiration, RQ = 1,
but there would probably still be a small contribution from ae. —— —5N - containing DNA
the conversion of lipid to sugar, RO 0.4—0.5.
21.9 Normally insufficient oxygen is able to penetrate the testa to
-
allow exclusively aerobic respiration; the RQ is a combina-
tion of the RQ for aerobic respiration (probably about 1.0) A B C
and that for anaerobic respiration, which is infinity (0), conservative replication
Removal of the testa allows more rapid penetration of
oxygen by diffusion, with a consequent increase in aerobic
respiration and decrease in RQ. Ethanol is a product of
anaerobic respiration so less accumulates when the testas are
removed.
21.10 During exposure of the pupa to a constant low temperature
of 3 °C for 6 weeks thoracicotropin is produced and stored by
neurosecretory cells in the brain. Transferring the pupa to a = | —'4N/©N
aaa - containing
constant temperature of 25 °C causes the release of thoraci-
cotropin into the blood which activates the thoracic glands to — N - containing DNA
produce moulting hormone, diapause is broken and pupal-
adult metamorphosis occurs.
C
Chapter 22 dispersive replication
Fig 22.2(ans) Diagrams explaining two further theories of

22.1 (a) Meiosis DNA replication. The appearance of DNA in a caesium
(b) W -—interphase chloride density gradient according to the theories presented
X — telophase I in Fig 22.14
Y — telophase II
(c) Gamete cells
22.2 See fig 22.2(ans).
22.3 Bases G
original sequence GTA! GTA| GTA; GTA, GTA;GTA;GTA| GTA
22.4 4 bases used once = 4X 1 = 4 = 4 adding three bases GTA, CGT AGT!ACG TAG| TAC GTA' GTA
4 bases used twice = 4 x 4 = 4 = 16
4 bases used three times = 4 x 4 x 4 = 4 = 64 { | sequence
4+C aG +C| restored
therefore the mathematical expression is x” where
x = number of bases and y = number of bases used. deleting three bases |GTA!GTGiTAG " AGT |GTA_ GTA'GTA
22.5 See fig 22.5(ans).
22.6 UAC AAG CUC AUG GUA CAU UGC -A -A -A
22.7 This suggests that the process of differentiation does not
sequence restored
involve the loss of, or damage to, the genetic material (DNA)
but the switching off of selected genes. The DNA was still Fig 22.5(ans) Answer to question expressed
able to function given the new environment of an undiffe- diagrammatically. The general principle behind restoring the
rentiated cell. Further experiments involving transplanting normal triplet sequence by the addition or deletion of 3 bases
donor nuclei from cells at a later stage of development did not is to add or delete three bases at any position along the
produce viable embryos. This was taken as evidence that length of the polynucleotide code
certain irreversible changes occur in the genetic material
during development which influence differentiation by
causing genes to be switched off irreversibly. However, later
experiments carried out by Gurdon at Oxford in the 1960s plays the dominant role in the process of development, the
involved transplanting nuclei from endothelial cells of cytoplasm (in this case the egg cytoplasm) plays a major role
tadpole intestine into enucleate egg cells from a mature frog. in determining the extent of the genetic expression of the
In many cases the eggs developed into normal frogs with the nuclear material, that is differentiation. In this case it
characteristics of the nucleus-donor tadpoles. This technique induced the nucleus to fulfil its complete genetic potential
is known as cloning and suggested that whilst the nucleus and produce a complete organism.
913
23.3 (a) If short black hair appeared in the F, phenotypes, then
Chapter 23 short hair must be dominant to long hair and black hair must
be dominant to white.
23.1 See fig 23.1(ans) below. (b) See fig 23.3(ans) below.
(a) Let: bet:
B represent brown fur (dominant) B represent black hair

b represent grey fur (recessive) b represent white hair
S represent short hair
Parental phenotypes brown fur x grey fur s represent long hair
Parental genotypes (2n) BB x bb
F, phenotypes short black hair X short black hair
Meiosis SsBb x SsBb
F, genotypes (2n)
a
Gametes (n) x (0) (o)
F, genotypes (2n) Bb Bb Bb Bb
Meiosis @ | @|@|@
F, phenotypes all brown fur Sb sB sb
Gametes (n)
(as shown by(# and @ ) SB SB SB
F, phenotypes brown fur x brown fur a a a
F, genotypes (2n) Bb x Bb
Meiosis Random fertilisation Sb sB sb
Gametes (n) (b) x (») Sb Sb Sb
(as shown by Punnett square)
Random fertilisation |_| =) B CL)
F genotypes (2n) BB Bb Bb bb
F, phenotypes 3 brown fur 1 grey fur F, genotypes (2n) SB Sb sB sb.
(as shown in each square) sB sB a | il
(b) =| go fi
Experimental phenotypes brown fur x grey fur
Experimental genotypes (2n) Bb x bb SB Sb sB sb
Meiosis sb sb sb dl sb |
Gametes (n) x (») (») B O
Offspring genotypes (2n) Bb Bb bb bb F, phenotypes 9 short 3 short 3 long 1 long
black hair white hair black hair white hair
Offspring phenotypes 1 brown fur 1 grey fur
In the case of monohybrid inheritance, the offspring from a heterozygous

genotype crossed with a homozygous recessive genotype produce equal
(Symbols) a >, silly avame
numbers of offspring showing each phenotype: in this case 50% brown fur
and 50% grey fur.
23.4 See fig 23.4(ans) below.
Let:
R,r and S,s represent two allelomorphic pairs of
23.2 If an organism having an unknown genotype is testcrossed genes controlling flower colour.
with a homozygous dominant organism, all the offspring will
show the dominant characteristic in the phenotype, as shown Parental phenotypes purple X purple
in fig 23.2(ans) below. Parental genotypes (2n) RrSs_ X RrSs
Meiosis
wets
T represent a dominant allele

9 ol
t represent a recessive allele Gametes (n) RS

(as shown by(% and 8 ) RS
Testcross homozygous X homozygous | heterozygous X homozygous eo .
phenotypes
Testcross genotypes ties UP Tt x TT Random fertilisation RS
(2n) (as shown by Punnett square)
Meiosis
Gametes (n) OO x @®a) OO x O® Offspring genotypes (2n)
Offspring genotypes TY TY TT TT TY 0) cette. ale (as shown in each square)
(2n)
Offspring phenotypes all tall (homozygous) all tall (4 homozygous,
’, heterozygous)
Offspring phenotypic ratio 9 purple : 7 white

(Symbols) @ O
914
23.5 The two alleles segregate during metaphase I and anaphase
I
23.6 The number of different combinations of chromosomes (a) Let:
in
the pollen gamete cells is calculated using the formula 2”
N represent normal wing (dominant)
where n is the haploid number of chromosomes.
’
n represent miniature wing (recessive)

In crocus, since 2n = 6, n = 3. R represent red eye (dominant)
Therefore, combinations =2 = 8. r represent white eye (recessive)
23.7 The F, phenotypes show that purple flower and short stem XX represent female fly ( 2)
XY represent male fly (@’)
are dominant and red flower and long stem are recessive. The
approximate ratio of 1:1:1:1 in a dihybrid cross suggests that (i) Parental phenotypes miniature wing, red eye X normal wing, white eye 2
the two genes controlling the characteristics of flower colour Parental genotypes (2n) XE ee
and stem length are not linked and the four alleles are
Meiosis
situated on different pairs of chromosomes (fig 23.7(ans)
below). Gametes (n) (x) x (x*) Ce
et
F, genotypes (2n) XX Ne Xne Ae KY x BEY:
P represent purple flower —————————
p represent red flower F, phenotypes normal wing, red eye? normal wing, white eye
S represent short stem
s represent long stem (ii) Assuming no crossing-over between the genes for wing length and eye
Since the parental stocks were both homozygous for both characters the colour in the female, the following results are likely to appear:
F, genotypes must be PpSs. F, phenotypes normal wing, whiteeyeo? X normal wing, red eye 2
Testcross phenotypes purple flower, short stem x red flower, long stem F, genotypes (2n) KNEE exe Xon
Testcross genotypes (2n) PpSs x ppss Meiosis
Meiosis
Gametes (n)
S| ®|©|@ Gametes (n) (x*)(*) x Ce
(as shown byo% and ©) PS Ps pS ps
F, genotypes (2n) xx XE AE xy OY,
Random fertilisation ps ps ps ps
(as shown in Punnett square) F, phenotypes normal wing, normal wing, miniature wing, normal wing,
red eye 9 whiteeyeQ redeyeo white eye
(listed in each square)
(b) The lack ofa 1:1:1:1 ratio of phenotypes resulting from this cross
Offspring phenotypes 1 purple flower, short stem: | purple flower, long stem: indicates crossing-over between the genes for wing length and eye colour
1 red flower, short stem: 1 red flower, long stem in the female.
Testcross phenotypes normal wing, red eye? X miniature wing, white eye <j
Testcross genotypes (2n) xerXN x x™y
23.8 (a) Homologous chromosomes Meiosis
(6) Body colour and wing length
COC Coey
(c) This cannot be concluded from the data since the position Gametes (n)
of the antenna shape gene is not shown. (as shown by 9 and)
23.9 Out of the 800 seeds produced, only 24 show the results of
Random fertilisation (x*) xr xo XNr yar x™ xo XNR Or
crossing-over between the genes for seed colour and seed
shape. In the other 776, the alleles for seed colour and seed
(as shown in Punnett square)
: ¢ : 2
shape have remained linked as shown by their approximate Offspring genotypes (2n)
1:1 ratio. (*) KEY exe ey. ENE ORY:
(as listed in squares)
Hence the crossover value is (24/800) x 100 = 3%. Cf cS co o
Therefore the distance between the genes for seed colour and
Offspring phenotypes wing: miniature normal miniature normal
seed shape is 3 units. eye: ied white white red
Experimental results 36 35 18 17
The alleles for wing length and eye colour are shown on the two F, female
(X) chromosomes in the explanation above. Crossing-over between the
alleles will give the recombinant genotypes shown above. Out of 106 flies,
35 show recombination of alleles (18 + 17), therefore the crossover value
is 35/106 = approximately 30%.
915
23.11 See fig 23.11(ans) below. 23.12 See fig 23.12(ans) below.
et:
Magpie moth
Let: B represent black coat colour
G represent ginger coat colour
N represent normal colour (dominant) XX represent female cat
n represent pale colour (recessive) XY represent male cat
Parental phenotypes pale colour male X normal colour female

Parental phenotypes ginger-coat male x black-coat female
Parental genotypes (2n) EXGCENA x ee
EITHER OR Meiosis
n@ n N Gametes (n) S) (y) x
n N@ N
Meiosis K EXE UXexe XEYuan XG
F,, genotypes (2n)
Gametes (n) F, phenotypes tortoiseshell- black-coat colour ed
x x coat colour 2
xa y: ee xX X Xe (The parental female must be homozygous for black coat colour since this is
Random fertilisation ot 9 of 9 the only condition to produce a black-coat colour phenotype).
Offspring genotypes (2n) x tortoiseshell-coat female

F, phenotypes black-coat male
EITHER OR F, genotypes (2n) BY. x XxeKe
niNin[N| Ni N n|Ni n n| Ni n
Meiosis
Gametes (n) @) x (x)
F, genotypes (2n) XBKG KBXB xsy xKBy
F, phenotypes tortoiseshell- black- ginger- black-
coat colour 2 coat colour Y coat colour
o% coat colour o
\ jee C7 See See
Offspring phenotypes normal normal Bee ay pale /
colour Q colour of colour colour?
From the results for the offspring phenotypes it is seen that the heterogametic 23.13 (a) See fig 23.13(ans) below.
sex in the magpie moth is the female. (b) There is a probability of4 (25%) that each child will have
blood group A. So the probability that both will have blood
Cat
group A is4 X 4 = 1/16 (6.25%).
et: Let:
B represent black colour (dominant)
I represent the gene for blood group
b represent yellow colour (recessive)
A represent the allele A (equally dominant)
B represent the allele B
Parental phenotypes black colour male ~X_ yellow colour female O represent the allele O (recessive)
EITHER OR Parental phenotypes blood group A x blood group B
B be b Be B b Parental genotypes (2n) ee x rene
Meiosis Meiosis
Gametes (n) Gametes (n) (*) (1°) x ae (1°)

x x
Random fertilisation w © w 0
Offspring genotypes (2n) Tale ere LOTS 11g
OLY, Xe ox XX xy. Offspring phenotypes blood groups AB A B O
Random fertilisation oo Q oS 2
EITHER OR
B| b| Bi bl b b B| bl B BL bLB
Ly RY Sh yaaa vo
Offspring phenotypes black —_-yellow black >< black !
colour? colour ao colour ow colour
From the results for the offspring phenotypes it is seen that the heterogametic
sex in the cat is the male.
916
Let:
P represent pea comb

R represent rose comb
a single P allele and a single R allele occurring together produce walnut comb
a double homozygous recessive genotype produces single comb
W represent white feathers (dominant)
w represent black feathers (recessive)
If eight different phenotypes are produced from the cross, each parent must
possess as many heterozygous alleles as possible. Hence the genotypes
are as shown below:
Parental phenotypes black, rose-comb cock x white, walnut-comb hen

Parental genotypes (2n) wwRrpp WwRrPp
Meiosis
Gametes (n)
(as shown by 9 and)
3 "
| | Pa
WRP WRp WwrP Wrp wRP wRp wrP wrp

Random fertilisation wRp wRp wRp wRp wRp wRp wRp wRp
(as shown in Punnet square)
O A © A @ A @ A
Offspring genotypes (2n) WRP WRp WrP Wrp wRP wRp wrP wrp
(as shown in squares) wrp wrp wrp wrp wrp wrp wrp wrp
O A oC @ A a &
Offspring phenotypes: 3 white, walnut-comb: 3 black, walnut-comb: 3 white, rose-comb: 3 black, rose-comb: 1 white, pea-comb:
(Symbols) O ® A A
1 black, pea-comb: | white, single-comb: | black, single-comb
G
23.15 Since both dominant alleles, W, white, and B, black, are
present in the heterozygous F, genotype, and the phenotype
is white, it may be concluded that the alleles show an epistatic Chapter 24
interaction where the white allele represents the epistatic
gene and the black allele represents the hypostatic gene. 24.1 A control experiment in which each variable was systematic-
The F, generation is shown in fig 23.15(ans) below. ally eliminated.
24.2 Redi’s basic assumption was that the presence of ‘worms’
Using the symbols given in the question, was due to the entry of the flies through the open flasks.
24.3 Sealing the broths would prevent the entry of organisms to
F, phenotypes
the vessels. Lack of air within the vessels may have deprived
white cock X white hen
organisms of oxygen for respiration.
F, genotypes (2n) WwBb X WwBh
24.4 Pasteur’s basic assumptions were that each generation of
Meiosis organisms develops from the previous generation and not
(as shown by ,fand 2)
ane | spontaneously.
24.5 Micro-organisms develop in the nutrient broth due to
Gametes (n) contamination of the broth by organisms in the atmosphere.
(as listed in Punnett square)
we| we | we| We 24.6 Sealed tubes, boiling, autoclaving, direct access to atmos-
phere, indirect access to the atmosphere.
O @ O| ©
24.7 Access to atmosphere.
F, genotypes (2n)
(as listed in the squares) weO| weO| weO| wbO 24.8 Pair A are not heated, pair B are boiled for 10 min, pair C
are autoclaved at 15 lb pressure for 20 min, pair D have
differential access to the atmosphere.
24.9 Spallanzani — tubes 3 and 4
Tod) eaaee ie Pasteur - tubes 7 and 8
24.10 Tubes 1, 3, 5 and 7
O O @| ®
24.11 The experimental design takes full account of scientific
bg mekgt de method. There is a hypothesis and the experimental design

includes appropriate controls and systematically eliminated
© © ® @ experimental variables. If identical experiments yield consis-
tent results these data may be regarded as valid. The degree
F, phenotypes 12 white colour: 3 black colour: 1 brown colour of validity attached to conclusions based on these, or any
(Symbols) © @ data, depends upon how accurately they interpret the data.
O47
Chapter 25
25.1 The carrier genotype is the heterozygous genotype. The
Hardy-Weinberg equation is used to calculate genotype
frequencies. The equation may be represented as
Dp SD Gita q =a
where
p= frequency of homozygous dominant genotype,
2pq = frequency of heterozygous genotype,
gq = frequency of homozygous recessive genotype.
The incidence of cystic fibrosis in the population appears in
individuals with the homozygous recessive genotype, hence
qg is 1 in 2000 or 1/2000 = 0.0005.
Thereforeg = V0.0005
= 0.0224.
Since p + q
Pp
The frequency of the heterozygous genotype (2pq) is

therefore
2 x (0.9776) x (0.0224)
= 0.044
= 1in 23
~ 5%
Approximately 5% of the population are carriers of the
recessive gene for cystic fibrosis.
25.2 Fasciola hepatica, the liver fluke, is a parasite which infests
sheep. It has an intermediate host, the snail Limnaea
truncatula, which lives in fresh water and damp pastures.
Draining ponds and wet areas would bring about a change in
environmental conditions which would exert a selection
pressure tending to eliminate Limnaea. As the numbers of
the snail fall this would reduce the numbers of available hosts
which would lead to a decrease in the numbers of the
parasite, Fasciola.
25.3 Reduced selection pressure at the extremes of each new
population would favour increased variability. New pheno-
types may show adaptations to the areas previously occupied
by the eliminated subspecies and spread inwards to occupy
the vacated ecological niche. The initial geographical
separation of the cline may have initiated allopatric specia-
tion. If the ring was reformed, gene flow may be impossible
due to genetic isolation and each subpopulation would
diverge genetically even further to form distinct species, as is
the present case in the British Isles where the species exist
sympatrically. If the genetic isolation between the two
subpopulations was not too great, hybrids may form when
the subpopulations were reunited. This zone of hybridisation
may act as a reproductive barrier as is the case with the
carrion and hooded crows.
918
a
Index
abscisic acid, 543, 549, 554, 555-6 allometric growth, 772 Arthropoda, 508
commercial application, 556 allopatric speciation, 900-1 excretion, 703-4
discovery of, 555 alternation of generations, 731 exoskeleton, 626
effects of, 556 amines, 603 eye, 597-8
geotropism, 549, 556 ammonia, 695 growth, 773
growth inhibitor, 776 amnion, 759, 767 locomotion, 650-1
low temperature adaptation, 672 amniote egg, 767 nervous system, 585
photoperiodism, 563 amniotic cavity, 767 osmoregulation, 703-4
structure of, 556 amniotic fluid, 759 articular facets, 631
summary of role in growth and Amoeba asexual reproduction, 723-8
development, 554 irritability, 585 budding, 725
synthesis and distribution, 556 locomotion, 645 fission, 723
abscission, 558-9, 563 osmoregulation, 700 fragmentation, 725
accommodation, 572, 594 amoeboid movement, 645 spore formation, 723-4
acetylcholine (ACh), 570, 573, 641-2 Amphibia vegetative propagation, 725-8
acetylcholinesterase (AChE), 570 circulation, 510-11 association areas, 582-3, see also cerebrum
acinus, 685 metamorphosis, 789-90 Astacus osmoregulation, 704
acromion, 632 osmoregulation, 708 aster, 798
acrosome, 751 secretion, 708 asynchronous muscle, 651
reaction, 757 skeleton, 628 atherosclerosis, 671, 689
actin, 636, 637-8 amplification, 572 atlas, 631
action potential, 567, 568 ampulla, 600 atrio-ventricular node, 524
amplitude, 568 anaemia, sickle cell, 856 atrium, 509
involuntary, 574 anaphase, 798-9, 802-5 atropine, 573
saltatory conduction, 569 anatomy, comparative, 875 autograft, see transplantation, types of
threshold stimulus intensity, 568 Andalusian fowls, 847 autosomes, 843
active transport, 697 androecium, 734 autotomy, 792
in nephron, 712, 718 androgen-receptor, protein molecule, 761 auxins, 543-50, 554
adaptation, 572, 588, 672 aneuploidy, 853 apical dominance, 557
to extreme climate, animals, 682, 683 angiotensin, 609, 720 cell division, 557
to temperature changes, plants, 672 anguilliform locomotion, see fish, locomotion commercial applications, 549
adaptive radiation, 873, 877-9 anisogamy, 732 effect on abscission, 558-9
of marsupials, 878 Annelida, 507 fruit set and fruit development, 559
adenine, 818 excretion, 701, 703 geotropism, 547-8
adenosine triphosphate (ATP), 640-1 locomotion, 648 mode of action, 549
adenylate cyclase, 604, 830 nervous system, 585 phototropism, 543-6
adipose tissue, 676 osmoregulation, 701, 703 pollen tube growth, 559
adrenal glands, 608-9 annual rings, 783-4 summary of roles in growth and
cortex, 609 antagonism, 557 development, 554
medulla, 609-10 antagonistic muscles, 632, 635, 648 synthesis, 546
adrenaline, 602-4, 609 antennal glands, 704, 705 Avery, McCarty and McCleod, 812
adrenocorticotrophic hormone (ACTH), 609 anther and development of pollen grains, Aves see birds
adsorption, 776 736-7 axial filament, 751
adventitious buds, 782 anthropocentrism, 611 axial skeleton, 629
adventitious roots, 782 anti-diuretic hormone (ADH), 602, 605, axis, 631
aerotaxis, 542 719-20 axon, giant, 565
aerotropism, 541 antibody, 534 axon, squid, 566, 569
aestivation, 794 clonal selection theory, 535 axoneme, 646
after-ripening, 776 antigen, 534
agglutination-inhibition test, 756 antrum, 677 B cells, 534-6
agglutinogens, 538 apical dominance, 554, 557 production of, 535
aggression (agonistic behaviour), 619 apocrine, 677 backcross, 834
agranulocytes, 514 apomixis, 730 bacteriophage, 813
albinism, 816, 885 apoplast pathway bark, 785
aldosterone, 720 leaves, 482-3 balance, 599-600
alkaptonuria, 816 roots, 495 barbs, 657
all or nothing response, absolute refractory appendicular skeleton, 629, 632 barbules, 657
period, 637, see also muscle limb girdles, 632 baroreceptor, 526
arachnoid membrane, 578 Barr body, 844
allantoin, 696
allantochorion, 759 _archenteron, 791 basal body, 800, see also centriole
allantois, 709, 759, 767 Aristotle, 858 basal ganglia, 580, 584
alleles, (allelomorph), 836, 837, 847, 885 arterio—venous anastomoses, 679 basal metabolic rate, 678
Allen’s rule, 682 arteriole, 507 basement membrane, 713
allograft, see transplantation, types of artery, 507, 517 basophils, 514
O19
bombykol, 618 cavum
Bateson, 832 aorticum, 510
bats, flight, 661-2 bone, 627
deposition, 628 pulmocutaneous, 510
Beadle and Tatum, 817
development of skeleton, 628 cell
bees, role of parthenogenesis in life cycle, 730
botulinum, 573 amacrine, 595
waggle dance, 621
bouton terminale, 570 ‘C’, 608
behaviour, innate, 611-22 chloride, secretory, 707
aggression, 619-20 Bowman’s capsule, 710, 713
Boysen-Jensen, 544 cycle, 796
biological rhythms, 614-16
brachiation, 628 differentiation, 769, 774
courtship and mating, 618
bradycardia, 670 diffuse bipolar, 595
individual—characteristics, 613
brain, 578-85 division, 769, 796, 800
instinct, 613
areas, 582 division and plant growth substances, 557
motivation, 613
association, 582, 583 effector, 565
patterns, selected, 611
convolutions, 581 excitable, 565
releasing mechanisms, 615
diencephalon, 584 expansion, 769
simple reflex, 613
function, 580-2 horizontal, 595
social, 620, 621
species characteristics, 613 human, 582-5 Leydig, 749, 751
mesencephalon, 584 monosynaptic bipolar, 595
territorial, 616-17
territorial, intraspecific competition, 617 metencephalon, 584 neurosecretory, 605
behaviour, learned, 611, 622-3 motor, 582 nonexcitable, 565
behaviour, study of, 611 myelencephalon, 584 nurse, 750
ethological approach, 611 nuclei, 580 parallel sensory, 588
mechanistic approach, 611 premotor, 583 secondary sense, 588
vitalistic approach, 611 sensory, 582 solenocyte, 701
Bergman’s rule, 682 structure, 580 vitrellar, 597
Bernard, Claude, 665 telencephalon, 582 centre
Berry, Prof R J, 866 ventricle, 578 cardiovascular, 669
bile, 685, 689 wave frequencies, 580 respiratory, 668
bilirubin, 689 branchial arches, 876 centriole, 800, see also basal body
biliverdin, 689 breeding centromere, 798-811
binary fission, 723 artificial selection, 893 centrum, 626
binocular vision, 597 inbreeding, 893-4 cephalisation, 585
biochemistry, comparative, 881-3 non-random, 888-9 cerebellum, 584
biogenesis, 858 outbreeding, 894-5 cerebral cortex, 582
biological control, 618 plant and animal, 874-5 cerebrospinal fluid, 578
biological rhythms, 614-16 pure, 831 cerebrum, 582-4
bipedal gait, 663 bromocriptine, 765 association areas, 582, 583
birds brown fat, 794 motor areas, 582, 583
circulation, 512 budding, 725 sensory areas, 582
excretion, 709-10 buds silent areas, 583
feathers, 656-8 lateral, 780-1 cervix, 752
flight, 657-61 see also apical dominance chaetae, 648
osmoregulation, 709-10 bulb, vegetative propagation, 725 chemoreceptors, 591-3
salt gland, 710 chemotaxis, 542
birth, 761-6 calcitonin, 602, 606, 608 chiasmata, 803, 809, 840
changes in foetal circulation, 765 calorigenesis, 607 chimpanzees, 621
foetus, 761 Calvin cycle, 489 chitin, 626
gestation period, 761 cambium, 490, 782 chloride shift, 531
blastocoel, 758, 791 cork, 784, 785 choice chamber, 612
blastocyst, 756, 758 vascular, 782, 783, 784, 785, 786 cholecystokinin, 602
blastomeres, 758, 791 capacitation, 757 cholesterol, 609, 689
blastopore, 758 capillarity, 492 cholinergic, 570
blastula, 791 capillary, 507, 517 chorion, 759, 767
bleaching, 586 inflammation, 533 chorionic gonadotrophin (HCG), 602, 756
blood capitulum, 631 chorionic villi, 760
control of metabolites, 670 carangiform locomotion, see fish, propulsion in choroid plexus, 578
control of respiratory gases, 668-9 carbohydrate metabolism in liver, 686-7 anterior, 578, 584
defensive functions of, 531-3 carbon dioxide carriage, 530 posterior, 585, 578
blood, mammalian, 512-23, see also heart, carbon monoxide, 530 chromatid, 796, 803
mammalian carbon chromatin, 811
carbon dioxide, 530-1 circulation in plant, 497 chromosome, 796
cells, 512-15 in pea plant, 498-500 and linkage group, 842
clotting, 531-3 carboxyhaemoglobin, 530 autosomal, 843
functions, 527 cardiac cycle, 522, 523 giant, 843
groups, 538-9, 847 cardiac muscle, 635 homologous, 837
inflammation, 533 cardiac output, 525 independent assortment, 810
oxygen carriage, 528 carpel, at fertilisation, 738 polyploidy, 895
pH control, 721 carpometacarpus, 658 puffs, 830
phagocytosis, 533 carrier, 845 sex, 843
plasma, 511-12 cartilage, 627 structural changes, 855
pressure, 526-7 types of, 627 theory of heredity, 836
pressure, control of, 670 cascade effect, 603 X and Y, 761, 843
pressure, diastolic, 671 Casparian strip, 495, 496 cilia, 646
pressure, systolic, 671 caste system, 621 circadian rhythm, 616
wound healing, 534 catecholamines, 609 circulatory system, Annelida, 507
blood group inheritance, 847 cation pump, 566 Arthropoda, 508
Bohr effect, 529 cavitation, 492 vertebrates, 509
920
circumoesophageal connectives, 585 crossing-over, 840-1
circumpharyngeal connectives, 585 production of cytokinin, 553
cryptorchidism, 749 replication, 813
cistron, 817 crystalline cone, 597
civetone, 618 role in inheritance, 812-13
cupula, 600 structure of chromosomes, 811
classification curare, 573 transcription, 822
immunological evidence, 883 curve, sigmoid, 769 dog, gait, 662, 663
Linneaus, 876 cuticle, 487 dogfish
phenetic, 874 transpiration loss, 482 circulatory system, 509-10
phylogenetic, 874 cuttings, 727 locomotion, 653—4
clavicle, 632 cybernetics, 665 dominance
cleavage, 791 cytochrome, 881-2 apical, 557
climacteric, 559 cytokinesis, 749, 800 incomplete, 846-7
cline, 900 cytokinins, 553, 554, 555 dopamine, 573
clitoris, 752 cell division, 557 dormancy, 792-3
clone, 723, 728, 729, 775 commercial application, 555 bud, 554
clotting, 531-3 discovery, 553 seed, 554, 776
co-ordination mode of action, 555 dormancy and photoperiodism, 563
endocrine, 610 summary of roles in growth and dormin, 672
nervous, 610 development, 554 dorsal light reaction, 612
coacervation, 861 synthesis and distribution, 555 dorsal vessel, 507
cochlea, 599 cytoplasm, 828 dorso-ventral muscles, 650
cockroach, vascular system, 508 cytoplasmic proteins, 774 down, 657
codon, 819 cytosine, 818 Down’s syndrome, 853
Coelenterata, excretory mechanisms, 700 drag, 658
nervous system, 585 dark, adaptation to, 596 driving force, 661
osmoregulatory mechanisms, 700 dartos muscle, 749 drought, adaptation to
cohesion-tension theory, 481, 492 Darwin, Charles, 543-4, 860, 862, 864, 865, endurers, 698
colchicine, 854 866, 868, 872 evaders, 698
collecting dust, 719 de Vries, Hugo, 853 duct of Bellini, 719
colostrum, 765 deamination, 687 bile, 685
colour blindness, 596 Decidua, 760 common hepatic, 685
colour vision, 596 demi-facets, 631 ductus arteriosus, 765
competition, 892 dendrites, 569 ductus venosus, 765
interspecific, 865, 892 dendrochronology, 783 dura mater, 578
intraspecific, 617, 865, 892 dendroclimatology, 783 dynein, 646
compound eye, 589 depolarisation, 567, 570
concentration gradient, 712 deuterostome, 870 ear, mammalian, 598-600
conditioning, 623 development balance, 599-600
conductance, 567 embryonic induction, 829 hearing, 599
conduction, 569 flowering plants, 776-86 ossicles, 876
decremental, 585 genetic control, 827 structure and function, 598
saltatory, 569 in plants and animals, 774-5 vestibular apparatus, 599
continental drift, 871 in vertebrates, 790-2 saccule, 599
contour, 657 role of environment, 829 semicircular canals, 599
control systems, 665-7 role of genes, 829 utricle, 599
regulators, 668 diabetes earthworm, 507-8
conus arteriosus, 509 insipidus, 720 dorsal vessel, 507
convergence, 588-9 mellitus, 610 vascular system, 507-8
copulation, 756 diapause, 793-4 eccrine, 677
coracoid, 632 diapedesis, 533 ecdysis, 626
Cori cycle, 687 diaphysis, 627 hormonal control, 787-9
cork, 784-5 diastole, 522, 523 ecdysone, 787
corm, vegetative propagation, 725-6 differential mortality, 890 Echinodermata, excretion, 706
corolla, 734 differential permeability, 712 ectotype, 892
corpora cardiaca, 788 differential reproductive potential, 890 ectoderm, 759
corpora quadrigemina, 584 differentiation, 769, 774 ectoplasm, 645
corpus albicantus, 755 diffusion, passive, 655, 712 ectotherms, 673, 674-5
corpus allatum, 788 diffusion pressure deficit, 481 temperature regulation in, 675
corpus callosum, 582 diffusion shells in transpiration, 484 effectors, definition of, 601, see also cells
corpus luteum, 753, 755 dioecious plants, 728, 738 egestion, definition of, 693
Correns and Tschermak, 836 discontinuous growth, 773 egg, cleidoic, 709, 767
cortical reaction, 757 discrimination, 573, 588 ejaculation, 756
costal cartilages, 632 dispersal of, fruits and seeds, 745-8 elasmobranchs, 707
cotyledons, 743 dispersal electro-osmosis, 505
countercurrent, animal, 745-6 electroencephalograms, 580
exchanger, kidney, 718 chance, 748 embryo sac, 737
heat exchanger, 683 self-, 748 embryology, 790
coupling water-, 748 comparative, 879-81
excitation-secretion, 570 wind-, 747 embryonic development, 759-61
cranium, 629 displacement activity, 620 embryonic development of human, 759-61
Crassulaceae, 485 diurnal rhythm, 616 in seed, 744
creatinine, 690 DNA, 796 sexual, 761
cretinism, 608, 816 comparative biochemistry, 881 embryonic disk, see yolk sac
crocodile, circulation, 511 double helix, 814 embryonic induction, 829
cross genetic code, 818-20 end-plate potential, 572
back, 834 ligase, 814 endocrine system, 565, 601-11
reciprocal, 832 polymerase, 814 hormone action, 601-4
O21
s
ultrafiltration, 698 tertiary, 754

endoderm, 759
exocytosis, 570 foramen ovale, 765
endolymph, 598, 599
exoskeleton, 625, 626 fossils, 867—70
endometrium, 752, 758
exteroceptors, see sensory receptors, types of founder principle, 889
endomysium, 572
endoplasm, 645 extraembryonic membranes, 759 fountain zone, theory of, 645
endorphin, 574 extrafusal fibres, 643 fovea, 590
extrapyramidal tracts, 583 fowl, comb shape, 848, 849
endoskeleton, 625, 626
eye, mammalian, 593-7 fragmentation, 725
endosperm, 743, 744
accommodation, 594 freezing point depression, 697
endotherms, 673, 675-84
adaptation to extreme climates, 682 binocular vision, 597 frequency code, 569
adaptation to low temperatures, 682-3 colour vision, 596 Frisch, Karl von, 621
loss of heat, 678 depth of focus, 594 frog
skin structure, 676 photoreception, 594, 595-7 circulatory system, 510-11
sources of heat, 678 retina, 594—5 cloning, 729
engram, 622 rods and cones, 595 locomotion, 729
enkephalins, 574 structure and function, 593-4 see also Amphibia
entropy, 671 fruit
environment, effects of changes in, shown by facilitation, 573, 585 development of, 743
invertebrates, 665 falciform ligament, 684 dispersal, 745, 746
environment, internal fallopian tube, 752 FSH (follicle stimulating hormone), 602, 751,
control, 667-8 fimbrial, 752 (SSS
temperature, 671 fat metabolism in liver, 689 fungicide, systemic, 473
environmental influences on phenotype, 852 fate maps, 791 fusiform initials, 782
enzyme feathers, 657-8
induction, 862 feedback, negative, 666 Galapagos Islands, 872-3, 902
repression, 826-7 modular, components of, 666-8 gametes, 728, 737
eosinophils, 514 feedback, positive, 567 in flowering plants, 737
epidermis, 676-7 feedback control, 589 gametogenesis, 749, see also spermatogenesis
epididymis, 749 femur, 634 and oogenesis
epilepsy, 581 Fensom and Spanner, 505 gametophyte, 731
epiphysis, 606, 627 ferritin, 689 gamma aminobutyric acid, 573
epistasis, 849-50 fertilisation ganglia
erector pili, 677 flowering plants, 743, 744 basal, 584
erythropoiesis, 689 humans, 756, 757, 758 cerebral, 585
erythrocytes, 513, 514 membrane, 758 ganglion, 574
formation in liver, 689 fibrin, 531, 532 dorsal root, 580
ethene, 543, 549, 554, 556 fibrinogen, 532 sympathetic, 577
commercial application, 556 fibula, 634 gas gland, 654
discovery of role, 556 filoplume, 657 gastrin, 602
effects of, 556 filtration pressure, 714 gastrula, 791
structure, 556 fish gastrulation, 791
summary of role in growth and anadromic, 707 gene, 815, 831
development, 554 catadromic, 707-8 complex, 849
synthesis and distribution, 556 euryhaline, 707-8 enzyme hypothesis, 816
ethology, 611 excretion, 706 flow, 889-90
etiolation, 559 locomotion, 652-5 frequency, 885
eucalyptus, 500 osmoregulation, 706-7 interactions, 846
eugenics, 895 fission, 723 lethal, 849
euglenoid motion, 648 flagella, 646 linkage, 838-40
euploidy, 854, 895 flight mapping, 841
euryhaline fish, 707-8 bats, 661-2 mutation, 815, 852, 856
evaporation, heat loss, regulated, 674 bird, 657-61 natural selection of, 866
latent heat of, 481 flapping, 659-60 poly, 851
evolution, theory of, 862-4 gliding, 661 pool, 885
biochemical, 857, 860 hovering, 661 promoter,.827
convergent, 879 insect, 650-1 recessive, 816
evidence for, 866-74 soaring, 661 recombination unit, 815
geographical distribution, 869, 871-4 florigen, 562 regulator, 825
in action, 866 flower reshuffling, 852
natural selection, 864-6 embryo sac, 737 role in development, 829
parallel, 879 fertilisation, 743 sex-linked, 845
past, 866 half-flowers, 735 testicular feminisation, 761
saltatory, 868 insect pollinated, 734-6 Y-linked testis-determining, 761
synthetic, 885 parts, 733-4 generation
see also palaeontology phytochrome control of flowering, 560-2 first filial (F2), 832
excretion pollen grains, 736 second filial (F1), 832
definition, 693 representative examples, 734-6 generator potential, 586
effect of environment, 699, 704-10 structure and function, 733-7 Genesis, 857
functions, 693 symmetry, 734 genetic control, 825
in animals, 694, 700-10 foetal circulation, 765-6 Jacob-Monod hypothesis, 825-6
in fish, 706 foetal membrane, 759 genetic counselling, 895
in plants, 694 foetus, 761 genetic drift, 889
in vertebrates, 706 follicle stimulating hormone (FSH), 602, 751, genetic load, 889
nitrogenous products, 695-7 53a genital ridges, 761
products, 694 primary, 753 genotype frequency, 886
secretion, 699 primordial, 752 geotaxis, 542
selective reabsorption, 699 secondary, 754 geotropism, 547-9
O72
germ layers, 791 guanine, 818 hyaline, see cartilage, types of
primary, 791 guard cells, 489
germinal epithelium cell, 750 hyaluronidase, 757
gustation, 591 hybridisation, interspecific, 900, 903
germination, 776-9 guttation, 493 hybrids, 894
conditions needed, 776 gynaecium, 734 Hydra, budding, 724
embryo growth, 778 gyplure, 618 nerve net, 585
epigeal, 778
hydrophytes, 697
hypogeal, 778 H-Y antigen, 761 hydrostatic skeleton, 625
physiology, 776-7 Haeckel, 879 hydrotactic, positively, 613
respiration in seeds, 778 haem, 689 hydrotropism, 541
gestation, 761 haemocoel, 507, 508 hyperglycaemia, 670
gibberellic acid see gibberellins haemoglobin, 528
gibberellins, 543, 549, 550-553, 554
hyperparathyroidism, 606
and carbon monoxide, 530 hyperplasia, 769
acid, 550 breakdown in liver, 689 hyperpolarise, 570
cell division, 557
comparative biochemistry, 882 hypertension, 671
commercial applications, 553 haemophilia, 533, 845 hypertonic, 697
discovery of, 550 haemopoiesis, 513 hypertrophic, 769
effects, 551-3 haemopoietic tissue, 513 hyperventilation, 669
effects on flowering, 562 hair follicles, 677 hypophysis, 605
mode of action, 553 halophytes, 697-8 hypostatic, 849
parthenocary, 559 halteres, 876 hypothalamo-hypophysial portal system, 605
photoperiodism, 563 Hammerling, 828 primary plexus of, 605
structure of, 550 haptonastic movement, 542, 543 secondary plexus of, 605
summary of role in growth and haptotropism, 541 hypothalamus, 584, 604-5
development, 554 Hardy-Weinberg equation, 886-8 FSH and LH release, 753
synthesis and distribution, 550-1 equilibrium, 886-8 heat balance, 680-2
girdle see skeletal system hearing, 599 hypophysial portal system, 605
gland, definition of, 601 heart rate, control of, 670 median eminence, 604
glenoid cavity, 632 heart, mammalian, 520-7, see also blood, nuclei, 604
Glisson’s capsule, 684 mammalian hypothyroidism, 608
globin, 689 cardiac cycle, 522 hypotonic, 697
glomerular filtration rate, 713 contraction, 524-5
glomerulus, 713 excitation, 524-5 IAA (indoleacetic acid), 546
glucagon, 602, 610 regulation of rate, 525 ICSH (interstitial cell stimulation hormone),
glucocorticoids, 602 structure, 520-2 751
gluconeogenesis, 687 heartwood, 784 imbibition, 776
glycogenesis, 610, 687 heat imipramine, 573
glycogenolysis, 687 exchanger, countercurrent, 683 immovable joints, 629
goitre, 608 heliotherms, basking, 675 immune system, 534-40
Graafian follicle, 753, 754 load, 683 B cells, 534-6
grafting, 727 production, 691 interferon system, 540
granulocytes, 514 source for animals, 673 T cells, 534-5
grebe, courtship, 618 helicotrema, 599 transplantation, 539
green glands, 704, 705 helix, 814 types of immunity, 537-8, 760
grey matter, 580 Henle, loop of, 710-12, 717 immunisation, 538
Griffith, Frederick, 812 hepatocytes, 684 immunity
ground meristem, 779 hermaphrodistism, 728, 730 acquired active, 583
ground tissue, 779-80 hermaphroditism, floral structure, 740 acquired passive, 537
growth, 769-74 heroin, 573 natural active, 537
adventitious, 782 herring, locomotion, 654-5 natural passive, 537
allometric, 772 heterozygous, 833 immunogen, 534
and development, 773 Hertwig, 836 immunoglobulin, 534
control of, 774 heterosis, 894 classes, 536-7
curves, 797, 798 heterostyly, 740 immunological memory, 537
definition of, 769 heterothermism, 672 implantation, 758-9
discontinuous, 773 HGC (human chorionic gondotrophin), 756 imprinting, 623
factors influencing, 774 hibernation, 794 inbreeding, 893-4
hormone releasing factor. 792 diurnal, 794 independent assortment, 810
in arthropods, 773 His, bundle of, 525 indoleacetic acid (IAA), 546
in flowering plants, 776-86 histones, 811 industrial melanism, 896—7
isometric, 772 homeostasis, definition of, 665 infection, control of, 671
mammalian hormones, 605, 792 role of kidney in, 712 inflammation, 533
measurement of, 796-8 homeotherm, 673, 681 inflorescence, 733
negative, 769 homocercal tail, 654 infundibulum, 605
of lateral roots, 781 homograft, 761 inheritance
patterns, 772-3 homology, 875-7 chromosomal basis, 836
plant growth substances, 542-59 homozygous, 833 dihybrid, 834
positive, 769 hormones, 601-11, 690, see also growth monohybrid, 832
primary, of root, 781 substances, plant of blood groups, 847
primary, of shoot, 779 horn, lateral, 580 polygenic, 851
primary plant body, 779 ventral, 580 sex-limited, 844
repair and regeneration, 792 Hoyle, Fred, 861 inhibition, 573
secondary, 782-5 human chorionic gonadotrophin (HGC), 756 initials, 779, 781, 782, 783
sigmoid, 769 human placental lactogen, 761 inner cell mass, 758
see also development humidity, effect on transpiration, 486 insects
growth substances, survey of role in plant Huxley, Aldous, 895 adaptive radiation of mouthparts, 877
growth and development, 554 Huxley, T. Henry, 866 diapause, 793
923
larval stages, 786-7 Meissner’s corpuscle, 591
insects—cont. melatonin, 606
ecdysis, 787-8, 793 latent period, 638
lateral membrane
flight, 650-1 basilar, 599
moulting, 787-8, 793 drag, 653
meristems, 782 differentially permeable, 475
pollination, 734-6, 739-42 effect of heat and alcohol, 481
taste and smell, 592-3 roots, 781
leaf primordia, 780 plasma, 475, 577
walking, 650 tectorial, 599
insensible water loss, 674 learning, 622-3
membrane granulosa, 752
insertion, 635 Leeuwenhoek, Anton von, 858
lenticels, 784-5 membrane potential, 565
insight learning, 623 depolarisation, 567
instincts, 613 transpiration loss, 482
leucocytes, 513-14 positive feedback, 567
insulin, 602, 610
Leydig cells, see cells memory, 622
integration, 573 memory cell, lymphatic, 835-6
intercellular fluid, 519 LH (luteinising hormone), 751, 753
life cycles, variety, 731 Mendel, Gregor, 815, 831
interlobular blood vessels, 685 inheritance experiments, 832-7
interoceptors, see sensory receptors, types of lifting force, 658
light, effects on transpiration, 485 traits, 831
interstitial cell stimulating hormone (ICSH),
limbs, 633-4 meninges, 578
751
pentadactyl, 875 menstrual cycle, 753
intervertebral discs, 629
limiting factors, 852 meristem, 779
intrafusal fibres, 643
linkage, 838-40 apical, 779, 781
intralobular vein, 685
groups, 842 initials, 781
intromittent organ, 756
Linnaeus, 876 intercalary, 779
ions
liver, mammalian, 684-91 lateral, 782
active uptake, 496
blood storage, 689 mescaline, 573
passive uptake, 496
carbohydrate metabolism, 686-7 Meselsohn and Stahl, 815
irritability, 565
islets of Langerhans, 610 detoxification, 690 mesencephalon, 584
excretion, 695 mesoderm, 759, 791
isogamy, 732
isograft, see transplantation, types of fat metabolism, 689 mesophytes, 485, 698
isolating mechanisms, 901 function, 686-91 metabolites, control of, 670
isometric contraction, 638 heat production, 691 metachronal rhythm, 647
isometric growth, 772 hormones, 690, 792 metacromion, 632
isotonic contraction, 697 lobules, 685 metagenesis, 732
mineral storage, 689 metamorphosis, 786-90
protein metabolism, 687-9 Amphibia, 789
Jacob—Monod hypothesis, 825
structure, 684-6 Insecta, 787-9
jaundice, 689 metanephridium, 701
Johannsen, 831 vitamin storage, 689
loading tension, 528 metaphase, 798-9, 802-5
joints, see skeletal system
locomotion, 625 metenkephalin, 574
junctional folds, 572
locus, 837 micelles, 689
juvenile hormone, 787
long-day plants, 561-2 Miller, Stanley, 860
long-day response see response mineral salts
kangaroo rat, adaptation to drought, 699 longitudinal muscle, 650 active transport, 496
keratin, 676 Lorenz, Konrad, 614 passive uptake, 496
Kettlewell, Dr H B D, 896 lungs, excretion, 695 recirculation, 497
kidney, 706, 710-21 luteinising hormone (LH), 602, 751, 753 remobilisation, 497
bladder, 706 luteolysin, 756 translocation through plants, 496
function, 712 lymph, 519, 520 uptake by roots, 495-6
hormonal control, 712 lymphocytes, 514 mineralocorticoids, 602
mesonephric, 706 thymocytes, 535 mitosis, 796, 797-9, 800-1, 810
metanephric, 706 lymphokines, 535 comparison with meiosis, 811
nephron, 706, 710-12, 713-15 lysergic acid diethylamide (LSD), 573 monoamine oxidase inhibitor, 573
position and structure, 710 monocytes, 514
see also Henle, loop of macrophage, 533 monoecious plants, 728
tubules, 715 maculae, 599 Morgan, Thomas H, 838, 845
kinesis, 542, 612 Malpighian layer, 676 morphine, 573
kinetin, 553, 555, 557 Malpighian tubules, 703 morphogenesis, 769, 774, 775
klinostat, 547 Malthus, Rev Thomas, 864 morula, 758
klinotactic response, 612 Mammalia mosaic eggs, 828
Kohler, 623 circulation, 512, 517-19 mosaic image, 598
Kornberg, 813 composition of blood, 512 motivation, 613
Krause, bulb of, 591 heart, 520-7 motor areas, see cerebrum
Kuppfer cells, 686, 690 mammary glands, 764 motor end-plate, 572
kymograph, 638 marram grass, 486 motor unit, 641
marsupials, adaptive radiation, 878 motorium, 648
labia, 752 Mason and Maskell, 502 moulting hormone, 787, 788
majora, 752 mass flow, 473 moulting see insects
minora, 752 mastigonemes, 646 multiple alleles, 847
labour, 762-4 mechanoreceptors, 590-1 Minch’s hypothesis, 504
Lack, David, 616 medulla oblongata, 585 muscarine, 573
lactation, 764-5 medullary rays, see rays muscle, 635-45
lactic acid, 641 megakaryocytes, 515 ‘all or nothing response’, 638
lactogenesis, 765 meiosis, 728, 732, 796, 801-10 cardiac, 635
lactose synthetase, 765 comparison with mitosis, 810, 811 contractile response, 638
lacunae, 759 movement of chromosomes, 837 direct, 650
Lamarckism, 864 sexual reproduction, 810 energy supply, 640-1
Lang, A., 505 variation, 852 excitation-contraction-coupling, 640
924
indirect, 650 chemical influences, 573 secretion, 699
innervation, 641-2 function, 572-3 selective reabsorption, 699
mode of action, 638 neurone, 565 ultrafiltration, 698
skeletal, 635-43 adrenergic, 570 osmosis, 474-81, 712
sliding filament theory, 639 afferent, 574 osmotic potential, 474, 697
smooth, 635, 643 auditory, 599 osmotic pressure, determination of, 474, 477-9
Striated, 635-8 cholinergic, 570 ossification, 628
tone, 644 efferent, 574 osteoblast, 628
muscle spindle, 591, 643-4 excitory, 576, 577 osteoclasts, 628
stretch reflex, 643-4 inhibitory motor, 577, 583 ostia, 508
muscle twitch, 638 internuncial, 575, 580 ostraciform locomotion, see fish, locomotion
muscone, 618 motor, 574 otoconia, 599-600
mutagenic radiations, 853 postganglionic, 577 outbreeding, 894-5
mutation, 852-3, 856 preganglionic, 577 ovarian cycle, 753
chromosome, 853 sensory, 574 ovary
effects of, 856 sudomotor, 679 cortex, 752
frequency, 853 visceral, 577 human, 752, 753
point, 852 neuronemes, 646 medulla, 752
random, 896 neurophysins, 605 ovule, development of, 737
somatic, 852 neurotransmitter substance, 570 oxygen dissociation curve, 528, 529
myelin, 568 neutrophil, 514 oxyhaemoglobin, 528
myofibril, 636 phagocytosis, 533 oxytocin, 602, 605, 762
myoglobin, 530 nicotine, 573
myogram, 638 Nirenberg, M, 819 Pacinian corpuscle, 586, 591
myometrium, 752, 762 node of Ranvier, 569 palaeontology, 867-70
myonemes, 648 noradrenaline, 570, 602, 609 horse, 868-9, 870
myosin, 636, 637 nucleoli, 798 pancreas, 610
myotomes, 652 nucleosomes, 812 panspermia theory, 860
myxoedema, 608 nucleotide base sequence, 818 Paramecium
nucleus, 827-8 locomotion, 646, 647, 648
nasal gland, 710 osmoregulation, 700
nasty (nastic movements), 542 obturator foramen, 633 parameter, definition of, 666
natural selection, 864, 872, 896 occipital condyles, 629 parasympathetic, 643
evidence for, 865-6 ocellus, 597 parathormone, 606
resistance, 896 odontoid process, 631 parathyroid gland, 602, 606
nature and nuture, 852 oedema, 533 parthenocarpy, 559
neo-Darwinism, 866, 885 oestradiol, 753, 755 parthenogenesis, 730
neoteny, 790 oestrogen, 602, 753, 761, 762 Pasteur, Louis, 858
neotonin, 787, 788 oestrous cycle, 753, 755, 756 pea plant, carbon in, 498-500
nephridium, 701 ommatidia, 589, 597 Mendel’s experiments, 831-7
nephridiopore, 701 On the Origin of Species, 862, 864, 866, 883 pecking order, 620
nephron, 706, 710-12 oocytes, 752 pectoral girdle, 632
cortical, 711 primary, 752 pelvic girdle, 632
juxta-medullary, 711 secondary, 752 penguins, territorial behaviour, 617
structure and function, 713-15 oogamy, 732 pentadactyl limb, 633
nephrostome, 701 oogenesis, 752-3 perennating organs, 725-7
nerve cord, 574 oogonia, 749, 752 pericarp, 743, 876
ventral, 585 Oparin, Alexander, 860 perilymph, 598
nerve impulse, 565-9 operon, 826 periosteum, 628
refractory period, 568 opsonin, 533 peristalsis, 643
speed of conduction, 568 optic chiasma, 597 personality, 583
transmission, 568 optic vesicle, 594 pectoralis
nerve net, 585 organ of Corti, 599 major muscles, 659
nerve tracts, 576, 580 organiser, 829 minor muscles, 659
ascending nerve fibre, 576, 580 primary, 829 pH, control of blood level, 721
descending nerve fibre, 576, 580 organogeny, 791 phagocytes, 533
nerves orgasm, 756 phagocytosis, 533
cranial, 574 Orgel, 860 phellem, 784-5, see also cork
mixed, 574 origin of life, theories of, 857-62 phelloderm, 784-5
spinal, 580 biochemical, 857, 860-1 phellogen, 784-5
vagus, 574 cosmozoan, 857, 860 phenoxyacetic acid, 550
nervous system, 565-74 special creation, 857 phenylketonuria, 816
autonomic, 577-8 spontaneous generation, 857, 858-9 pheromones, 592-3, 618
central, 578-85 steady state, 857, 859-60 biological control by use of, 618
parasympathetic, 577 see also evolution, theory of phloem, 473
peripheral, 574 origin of muscle, 635 development of, 779, 780, 781
phylogenetic development, 585, 587 ornithine cycle, 688, 695 mechanism of translocation, 502—6
sympathetic, 577 orthokinesis, 613 mineral transport, 496-7
synapse, 569-74 osmoconformer, 697 movement in, 502
vertebrate, 574-85 osmoregulation, 693, 696, 697, 706 protein, 502, 505, 506
neural spine, 629 and nitrogenous excretion, 696-7 secondary, 782, 783, 784
neuroendocrine environmental effects, 699, 704-10 structure, 497, 500, 501
reflex, 606 functions, 693 translocation, 500, 502
response, 606 in animals, 700-10 translocation of organic solutes, 497-506
transducer, 606 in plants, 697-8 photomorphogenesis, 559, 561, 779
neurohaemal organs, 605 mechanisms, 697 photons, 593
neuromuscular junction, 570, 572 phylogenetic, review of, 699 photoperiodism, 560-2, 563, 615
925
s
photoreception, 595-7 prezygapophyses, 629 polysynaptic spinal, 575

colour vision, 596 primary growth of plants, 779-82 polysynaptic spinal/brain, 575-6
colour vision, purple, 595 primrose, pin-eyed and thrum-eyed flowers, spinal, 575, 613
photosynthetic slump, 672 740 stretch, 643-4, 645
phototaxis, 542 principle of independent assortment, 834-5 reflexes, simple, in vertebrates, 613
phototropism, 542 principle of segregation, 833 flexion, 613
phyllotaxis, 780 procambium, 779, 780, 781 stretch, 613
phylogeny, 814 progesterone, 602, 773, 775, 761, 762, 765 refraction of light, 594
phytochrome, 559-60 prolactin, 602, 753, 883 refractory period, 525, 568
discovery of, 559 action of in vertebrates, 883 absolute, 568
pia mater, 578 promotor gene, see gene relative, 568, 638
pineal body, 584, 606 pronephros, 706 regeneration, tissue, 792
pituitary gland, 605 pronuclei, 758 regulator gene, see gene
adenohypophysis, 605 prophase, 798-9, 802-5 Reissner’s membrane, 599
gonadotrophins, 775 proprioceptors, 586, see sensory receptors, relay hypothesis of phloem translocation,
neurohypophysis, 605 types of 505-6
posterior, 604 protease, 757 releaser, 614
somatotrophin, 792 protein renal
stalk, 605 kinases, 609 cortex, 710
placenta, 759, 760 metabolism in liver, 687-8 medulla, 710
chorionic villi, 760 synthesis, 820-1 papillae, 710
plasma see blood, mammalian prothoracic gland, 787, 788 pelvis, 710
plasma prothoracicotrophic hormone (PPTTH), 788 pyramids, 710
clearance, 713 protoderm, 779 threshold, 610
protein production, 683 protonephridium, 701 remiges, 657
plasmagel, 645 protostome, 870 renin, 602, 609, 720
plasmasol, 645 Protozoa, osmoregulation and excretion, repair, tissue, 792
plasmodesmata, 483 700-01 replication
plasmolysis, 476, 477 Pseudopodia, 645 conservative, 814
plate tectonics, 871 pterylae, 657 dispersive, 814
platelets, 515 punctuated equilibria, 868 semi-conservative, 814
Platyhelminthes, 700-1 Punnett square, 835 repolarisation, 567
pleiotrophy, 849 Purkinje cells, 584 repressor molecule, 825
podocytes, 713 Purkinje tissue, 525 reproduction, 723-67
poikilotherm, 673 pyramidal tracts, 583 sexual, see sexual reproduction
polar body, 752 pyrogens, 682 Reptilia
first, 752 pyrrole rings, 689 circulation, 511
second, 752 pyruvic acid, 641 cleidoic egg, 709
pollen, development of grains, 736, 742 excretion, 709
pollination, 732, 737-43 osmoregulation, 709
quanta, 593
cross-, 738 skeleton, 628
quill, 657
features favouring, 738 reserpine, 573
insect-, 743-6, 739-42 respiration
self-, 738 rabbit, skeleton, 629-35 control of gases, 668-9
tube-growth, 559, 743 race, 900 in germinating seeds, 778
wind-, 740 geographical, 900 respiratory quotient (RQ), 778
polyembryony, 732 rachis, 657 response
cyclic, 732 radiation, adaptive, 877-9 long-day, 793
polymorphism, 621, 732, 892, 897-900 radioimmunoassay, 601 short-day, 793
balanced, 899 ramus communicans resting potential, 565
genetic, 899 (grey), 577-8 reticular activating system, 583
transient, 899 (white), 577-8 reticular formation, 583
polyploidy, 854, 895 ratio reticulo-endothelial system, 686
allopolyploidy, 854 dihybrid, 835 reticulospinal tract, 583
autopolyploidy, 854 monohybrid, 833 retina, 594-5
euploidy, 854 ray initials, 782 retinene, 595
polyspermy, 758 rays, 782, 783, 784, 785, 786 retrices, 657
pons, 584 receptor potential, 586 reverberating circuits, 622
population, 889 receptors rhabdome, 597
donor, 889 a adrenergic, 610 rheotactic response, 618
pioneer, 889 6 adrenergic, 610 rhesus factor, 538
recipient, 889 feedback control, 589 rhizome, vegetative propagation, 726
population genetics, 885-90 lateral inhibition, 589 rhodopsin, 595, 789
porphyropsin, 789 properties of, 588, 589 rib, 632
portal tract, 685 rapidly addapting (phasic), 588 ring, annual, see annual rings, 783-4
portal vein, 509 slowly adapting (tonic), 588 ringing experiments, 502, 503
postsynaptic membrane, 570 spontaneous activity, 589 rings of Balbiani, 830
postsynaptic potential recombination, genetic, 840 RNA, 819
excitatory, 571 Redi, Francesco, 858 comparative biochemistry, 881
inhibitory, 571 reduction division, 796 messenger, 821, 822
postzygapophyses, 629 reflex ribosomal, 821
potometer, 484 action, 574-7 role, 821
prefrontal lobes, 583 arc, 574-7 transcription, 822
pregnancy testing, 756 conditioned, 576 transfer, 821
pressure flow hypothesis see Miinch’s inhibitory, 644 translation, 822-3
hypothesis milk ejection, 765 robin, territorial behaviour, 616
presynaptic membrane, 570 monosynaptic, 575 rods and cones, 585
926
root
sex hormones spores, 723-5
adventitious, 782 female, 753, 755, 756, 761, 762, 765
dorsal, 580 sporophyte, 731
male, 751, 752 starch-statolith hypothesis, 548
hair, 481 sex linkage, 844, 845
lateral, 781 Starling’s law, 526
sexual reproduction, 728-67 stem
pressure, 493
Angiospermae, 732-48 primary anatomy, 490
primary anatomy, 493-4 comparison with asexual reproduction, 731
primary growth, 781-2 primary growth, 779-80
female reproductive system, 752-6 secondary growth, 782-5
secondary growth, 785-6 life cycles, 731-2 stereoscopic vision, 597
uptake of water, 493-4 male reproductive system, 749-52 sternum, 623
ventral, 580
origins, 730 steroids, 603
root cap, 549, 781
phylogenetic review, 766 stimuli, 565
royal jelly, 621 short-day plants, 561-2 behavioural, 614
RQ see respiratory quotient short-day response see response motivational, 614
Ruffini, organ of, 591
sickle cell anaemia, 856 releasers, 614
runner, vegetative propagation, 726 sieve plate, 500-1 sign, 614
sieve tubes, 500, 501, 503, 506 terminating, 614
sacroplasmic reticulum, 636 loading, 506 stolon, 726
salt glands, 709, 710 ultrastructure, 501-2 stomata
sapwood, 784 silent areas, see cerebrum distribution, 487
movement of water in, 493 sinking force, 661 effect of wind, 487
sarcolemma, 572, 636 sinks, 496-7 exit of water, 483
sarcomere, 637 sino-atrial node, 524 mechanisms of opening and closing, 489
sarcoplasm, 636, 637 sinus venosus, 509 structure, 489
scarification, 776 sinuses, 508, 509 sunken, 486
scent posts, 617 sinusoids, 685 transpiration loss, 482
schizogeny, 723 skeletal muscle, 625, 635 stratification, 563, 776
Schmidt-Nielsen, Knut, 699 skeletal system, 625 stratum corneum, 676
Schwann cell, 568 functions, 625 stratum granulosum, 676
scotopsin, 595 girdle, 623-4 stretch reflex, 644, 651
scrotal sac, 749 hydrostatic, 625-6 stridulation, 618
sebaceous glands, 677 joints, 634 subarachnoid space, 578
sebum, 677 limbs, 633-4 suberin, 485, 785
secondary growth of plants, 782-6 rabbit, 629-35 subliminal stimulus, 636
secondary growth see growth secondary vertebrate, 627-35 sucrose, Osmotic pressure of solutions, 478
secondary phloem, see phloem skin suction pressure see water potential
secretion, by kidney, 712 excretion, 694 summation, 588, 589
secretion, definition of, 693, 699 human, 676-794 spatial, 571
seed sweat production, 679 temporal, 571
advantages and disadvantages, vertebrate, 677-8 super-gene, 899
744 skull, 629 sweat glands, 677
development to dormancy, 743 smell, sense of, 592 sweat production and evaporation, see skin
dispersal, 745, 748 smooth muscle, 635 swim bladder, 654
germination, 776-9 social swimming, fish, 652-5
selection, 890-900 behaviour, 620 sympathetic system 643
artificial, 893, 895 ‘caste’ system, 621 symplast pathway, 482-3
in Man, 895-6 hierarchy, 620 leaves, 482-3
directional, 891-2 organisation, 621 movement of ions, 496
disruptive, 892 sodium roots, 494
inbreeding, 893-4 control of blood level, 720 synapse, 569-74
outbreeding, 894-5 inactivation, 567 axodendritic, 569
pressure, 890, 892 somatomedin pump, 566, 607, 690, 792 axosomatic, 569
stabilising, 891 somatostatin, 792 chemical, 570
selective advantage, 865 somatotrophin, 690, 792 chemical influences, 573
disadvantages, 865 sound, 598 electrical, 570, 572
predation, 896 Spallanzani, Lazzaro, 858 excitatory, 571
readsorption, 699, 712, 715 special creation, 857 fatigued, 573
semen, 756, 757 speciation, 900-4 function, 572-3
seminiferous tubules, 749 allopatric, 900, 901 mechanism of transmission, 570-1
sensitivity, 565, 588 interspecific, 900, 903 synaptic
sensory areas, see cerebrum intraspecific, 900-1 cleft, 570
sensory receptors, 565, 586-90 sympatric, 900, 902 convergence, 573
adaptation, 588 species knob, 570
convergence, 588-9 concept of, 900 vesicle, 570
feedback control, 589 polytypic, 900 synchronous muscle, 651
function, 590-600 ring, 902 synergism, 556-7
lateral inhibition, 589 Spemann and Mangold, 828 syngergists, 635
properties, 588 spermatid, 750, 751 systole, 522, 523
structure, 590-600 spermatocyte
summation, 571, 589 primary, 750, 751 T cells, 534-5, see also transplantation
transducers, 586 secondary, 750 T system, 572
types of, 586 spermatogenesis, 750-2 tachycardia, 670
serotonin, 573 spermatogonia, 749, 750, 751 tap root, swollen, 727
Sertoli cells, 750, 751 spermatozoa, 749, 751, 757, 758 taste, 591
serum, 533 spermiogenesis, 750 hairs, 592
Sewall-Wright effect, 889 sphygmomanometer, 670-1 taxis, 541-2, 612
sex determination, 843 spinal cord, 578, 580 tectum, 584
oe
\
transplantation, 539 vascular system
teleology, 611 closed, 507
teleost, marine, 707 graft rejection, 539
lcellss539 open, 507
telophase, 789-9, 802-5
tissue matching, 539 vascular, cambium, 782, 783, 784, 785
temperature regulation, 671 vasoconstriction, 526
adaptation to high, 683-4 types of, 539
transport vasodilation, 526
adaptation to low, 682-3
in animals, 507 vasomotor centre, 526-7
adaptation to, in plants, 672 vasopressin, 605
body, 673-4 in plants, 473-506
transverse processes, 629 vegetative propagation, 725-8
core, 673
transverse tubule system, 572 vein, 507, 517
effects on transpiration, 485
triad, 636 ventricle, 509, 578
heat balance, 680-2
trimethylamine oxide, 696 ventricular
influence on animals, 673
triplet code, 818 diastole, 522
influence on plants, 671
codon, 819 systole, 522
loss of heat, 678
degenerate, 819 venule, 507, 518
sources of heat, 678
frame shifts, 818 vernalisation, 562
terrestrial ectotherms, in 675
non-overlapping, 818 vertebrae, 630-2
tendon, 644
translation, 820, 824 caudal, 632
territorial behaviour, 616
universal, 820 cervical, 630
testcross, 834
trophoblast, 758 lumbar, 632
testis, human, 749
trophoblast cells, 756 sacral, 632
endocrine function, 751
trophoblastic villi, 759 thoracic, 631
testosterone, 602, 749, 752
trophollaxes, 621 vertebral column, 629-32
tetanus, 525
tropism, 541 vertebrarterial canal, 630
thalamus, 584
tropomyosin, 637-8 vertebrates, circulatory system, 509
theca, 753
troponin, 637-8 vestibular glands, 752
externa, 753
tuber, 726-7 visible spectrum, 593
interna, 753
tubercle, 632 visual
thermal gradient, 674
thermoreceptors, 591 tubercular facets, 631 acuity, 590
Thorpe, 623 tuberculum, 631 cortex, 597
thrombin, 531 tubulin, 800 field, 593
thrombosis, 533, 690 tunica pathway, 597
thrombus, 533 externa, 517 purple, see photoreception
through conduction tracts, 585 intima, 517 vitamin storage in liver, 689
thymine, 818 media, 517 Von Baer, 879
thymocytes, 535 turgor pressure, 476, 697 vulva, 752
thymosin, 535 twitch fibres, 643
thymus in immune system, 535 wall pressure see turgor pressure
thyroid gland, 606-8 ultrafiltration, 698, 712, 713, 715 Wallace, Alfred Russel, 864, 872
calcitonin, 606 umbilical cord, 760 water balance in terrestrial organisms, 708-9
structure of, 606 unidirectionality, 572 water movement
thyroglobulin, 606 universal across leaf, 481-8
thyroxine, 606, 789 donors, 538 across root, 493-5
triiodothyronine (T;), 606 recipients, 538 up stem, 492-3
thyroid stimulating hormone (TSH), 607, 789 urea, 688, 695 water potential, 474, 476-7
thyrotoxicosis, 608 uric acid, 695-6, 703 determination of, 479
thyrotrophin releasing hormone (TRH), 607, urine, 706 diffusion pressure deficit, 481
789 formation, 716 of air, 481
Tinbergen, Niko, 614, 619-20 hypertonic, 716, 719 suction pressure, 481
tinsel flagellum, see flagellum hypotonic, 719 water ascent in xylem, 492
tonic fibres, 643 uterine cycle, 753, 756 Watson and Crick, 813
tonicity, 697 uterus, 752 Went, 544-5
tonoplast, 475 white fibrous cartilage, see cartilage, types of
torpor, 794 vaccination, 538 white matter, 580
toxoids, 538 vaccine, 538 Wilberforce, Bishop Samuel, 866
transanimation, 688 vacuolar pathway, 482-3 wilting, 672
transcription, 822 leaves, 482-3 wind pollination, 740
transduction, 586 roots, 494 wind, effect on transpiration, 487
transfer cells, 496, 506 vacuole wood
transforming principle, 812 contractile, 700 diffuse porous, 783
translocation, 473 plant cell, 475 ring porous, 783
in organic solutes, 497 vacuum activity, 620 see also xylem secondary
in phloem, 500, 502 vagina, 752
of mineral salts through plants, 496-7 valves, 517-18 yellow elastic cartilage, see cartilage, types of
of organic solutes in phloem, 497-506 vane, 657 yolk sac, 759, 767
of water through plants, 481-95 vapour pressure, 486
transpiration, 481 variation Zea mais
cohesion-tension theory, 481, 492 continuous, 851 cytokinin, 553
cooling effect, 488 discontinuous, 851 mineral recirculation, 497
environmental factors, 484, 485-7 intraspecific, 851 root, primary anatomy, 493
functions, 488 sources of, 852 zeatin, 553
mineral salt distribution, 488 vas deferens, 749 zona pellucida, 753
plant and internal factors, 487-8 vasa efferentia, 749 zone of tolerance, 674
rate of, 484 vasa recta, 718-19 zygote, 728, 758
stream, 481 vascular bundles, 492
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BIOLOGICAL
SCIENCE &—
These two books provide a comprehensive account of biological
information required by students following General Certificate of Education
advanced level and Higher School Certificate courses in the biological
sciences. In addition, all the topic areas recommended by the Interboard
Working Party on the advanced level Common Core in Biology are
incorporated. First year students in higher education will also find the books
useful.
Biological Science 1: Organisms, Energy and Environment considers

examples of biological organisation at all levels from viruses and
prokaryotes to fungi, plants and animals. Detailed information on
biochemistry, cell structure, histology, nutrition and mechanisms of energy
utilisation in plants and animals is presented. In addition, a
thorough account of qualitative and quantitative aspects of ecology is
provided and guidance is given in the techniques required for
synecological and autecological investigations.
Biological Science 2: Systems, Maintenance and Change contains

information on transport, co-ordination, movement, homeostasis, excretion,
reproduction and development, genetics, evolution and speciation.
Accounts and theories on the origin of life, evolution and mechanisms of
speciation have been prepared in the light of recent developments and an
objective approach to these topics has been adopted throughout.
Comprehensive appendices in Book | give information on fundamental
ay
physical and chemical concepts, scientific method, biological techniques,
biomathematics and classification.
Numerous experiments and investigations representing essential practical

work at advanced level are included. These have all been fully tested.
Questions are interspersed throughout both books. These are designed to

test comprehension and application of knowledge. Answers to these
questions are provided in a special chapter at the end of each book.
_ The two books are fully illustrated with more than 700 line drawings and 250
black and white photographs. —
—~ ee SN
aN
\ S

N.P.O. Green, G.W. Stout, D.J. Taylor - Editor - Roland Soper - Biological Science 2. 2-Cambridge University Press (1985)

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BIOLOGICAL “=

N.P.O. GREEN B.Sc., M.I.Biol

G.W. STOUT B.Sc., M.Ed., M.LBiol

D.j. TAYLOR B.Sc., Ph.D., M.I.Biol

The right of the

CAMBRIDGE UNIVERSITY PRESS

©Cambridge University Press 1984

First published 1984

Printed in Great Britain by Ebenezer Baylis & Son Ltd

British Library cataloguing in publication data

Cover design by Andrew Weall and Associates

Table 14.1 Some mass flow systems of animals and plants.

Mass flow system Material(s) moved Driving force Location

respiratory system air or water respiratory muscles Mollusca, Chordata

Uptake Transport Elimination

* Movement of gases is considered in further detail in chapter 11.

O solute molecules or ions Method

a ioWhich solutionhas the higher water potential(ys2 Results

cell wall - freely permeable to small molecules

14.1.6 Plasmolysis and turgor pressure (TP)

__ the cell wall and the shrunken protoplast in plasmo--

used instead, referring to the pressure of the wall against

sucrose solution water/cm sucrose solution/cm* Concentration of

Molarity of sucrose Length of beetroot Length of beetroot

0.00 (distilled water) 4.8 5.0 5.3 eooe =kG

cut portions of scape

plane of cut epidermis

ee A, B, C Strips after they have been allowed to come to

Fig 14.6 Experiment on dandelion scapes.

(1) Stomata: by evaporation of water from cells and

(a) lignin cellulose xylem vessel

SYMPLAST cell sap inside vacuole

evaporation of water from cell

cells. Theoretically each stoma has a diffusion gradient, or Method

syringe containing water for pushing air

seas fee [hans

14.3.8 Functions of transpiration

protoxylem endarch, i.e. nearer centre lysigenous canal formed from

eye sclerenchyma fibres

(a) root hair root hair b)

(b) Primary anatomy of the root of

(a) cell wall piliferous

radial wall transverse wall

vacuolar pathways is not known.

14.6 Uptake of mineral salts and their SS

the root is the release of ions into the xylem. To achieve

Leaf removed from bag, and plant left

Whole plant Quantitative determination

Autoradiography of '*C-labelled plants plant is cut up into its component parts

radioactive sample + scintillation fluid. As

Fig 14.21 (opposite and above) Stages in an experiment to

14.8.1 Features of phloem translocation

plastid hts oes

phloem protein may or may not be

14.8.2 Ultrastructure of sieve tubes

(a) (b) (c)

of an aphid inserted into

==—) mass flowof solution a manometer attached to B would

semi-permeable membrane semi-permeable membrane