Concepts of Biology
Concepts of Biology
Concepts of Biology
SECOND EDITION
Sylvia S. Mader
TM
Mader, Sylvia S.
Concepts of biology / Sylvia S. Mader. — 2nd ed.
p. cm.
Includes index.
ISBN 978-0-07-340348-9 — ISBN 0-07-340348-2 (hard copy : alk. paper)
1. Biology—Textbooks. I. Title.
QH308.2.M234 2011
570—dc22
2010017189
www.mhhe.com
PART I PART IV
Organisms Are Composed of Cells 22 Plants Are Homeostatic 429
2 Basic Chemistry and Cells 24 21 Plant Organization and Homeostasis 430
3 Organic Molecules and Cells 42 22 Transport and Nutrition in Plants 450
4 Structure and Function of Cells 62 23 Control of Growth and Responses in Plants 468
5 Dynamic Activities of Cells 84 24 Reproduction in Plants 488
6 Pathways of Photosynthesis 104
7 Pathways of Cellular Respiration 122 PART V
Animals Are Homeostatic 505
PART II
25 Animal Organization and Homeostasis 506
Genes Control the Traits 26 Coordination by Neural Signaling 524
of Organisms 140 27 Sense Organs 548
8 Cell Division and Reproduction 142 28 Locomotion and Support Systems 568
9 Patterns of Genetic Inheritance 168 29 Circulation and Cardiovascular Systems 588
10 Molecular Biology of Inheritance 190 30 Lymph Transport and Immunity 608
11 Regulation of Gene Activity 210 31 Digestive Systems and Nutrition 626
12 Biotechnology and Genomics 228 32 Gas Exchange and Transport in Animals 648
33 Osmoregulation and Excretion 664
PART III 34 Coordination by Hormone Signaling 680
Organisms Are Related and Adapted 35 Reproduction and Development 698
to Their Environment 246
13 Darwin and Evolution 248 PART VI
14 Speciation and Evolution 270 Organisms Live in Ecosystems 724
15 The Evolutionary History of Life on Earth 290 36 Population Ecology 726
16 Evolution of Microbial Life 308 37 Behavioral Ecology 742
17 Evolution of Protists 332 38 Community and Ecosystem Ecology 758
18 Evolution of Plants and Fungi 350 39 Major Ecosystems of the Biosphere 780
19 Evolution of Animals 378 40 Conservation Biology 798
20 Evolution of Humans 410
iii
“This book uses everyday language to immerse the student into the world
of science.” —Michael P. Mahan, Armstrong Atlantic State University
iv
Imagine trying to redesign a vital mechanical part of an airplane, natural selection. Particularly in plants, many examples of gene
Applications are used throughout Concepts of Biology to while still keeping that plane in flight. Sounds nearly impossible,
doesn’t it? This was one of the puzzles facing early evolutionary
duplication have been found—for example, the wild mustard
plant has undergone at least two duplications of all its chromo-
show how biological concepts relate to students’ lives. biologists. After all, mutations are the main way in which new
traits and features arise during evolution, and yet most mutations
somes in the past, as well as duplication of several individual
genes at various times in history.
cause damage. If a feature is important, how can it be altered An intriguing example of gene duplication involves the
while still allowing an organism and its offspring to survive? sweet-tasting proteins. Of the thousands of proteins studied so
• NEW How Life Changes applications emphasize Geneticists have shown one possible way mutations can
accumulate without impairing present function: gene duplication
far, most have no noticeable flavor—but about half a dozen have
an intensely sweet taste. These rare, sweet-tasting proteins are
(Fig. 13B.1). An extra (and possibly unused) copy of a gene may found in plants and plant products from several different conti-
evolution as the unifying theme of biology and result from errors during cell division, efforts to repair breakage
to DNA, or other mechanisms. The surprising idea here is that
nents: The protein “curculin” is found in the fruit of a Malaysian
herb (Fig. 13B.2); “mabinlin” can be extracted from a traditional
these seeming accidents actually can provide raw material for Chinese herb; “thaumatin” is found in the fruit of a West African
how it pertains to students’ lives. rain forest shrub; and “brazzein” comes from a fruit that grows
wild in Gabon, Cameron, and Zaire. Each of these proteins tastes
sweet only to humans and certain monkeys. From the plant’s
• How Biology Impacts Our Lives single
g copypy of gene
g
point of view, the proteins likely provided an advantage: Sweeter
fruits would be eateneate more mo often and their seeds distributed
more widely, ensuring ng the growth of more plants with genes for
n
applications examine issues that making sweet proteins.
unusual proteins come
ns. A question still remains: How did these
n
me about?
m ab
H O W B I O L O G Y I M PAC T S O U R L I V E S duplicate copies of gene
Application
affect our health and environment. 8B Tissues Can Be Grown in
FIGURE 13B.1
the Lab
Duplication of a
No one yet knows ws exa
w
gene duplication is a like
exactly how these proteins originated, but
likely answer. The proteins look nothing
alike, are found in unrelated
unrela
u plants, and clearly did not come
gene followed by
Most people are now aware that stem cells can undergoathe cell cycle
mutation in one them a human embryo must be destroyed. Embryos fromaresome
some-ancient sshared plant gene. Each protein, however,
• How Science Progresses and generate tissues for the cure of devastating human
such as diabetes, cancer, brain disorders, and heart
diseases,
of the genes is a
way ailments
mutated copy of gene
times “left over” at fertility clinics, but even so many
for complexity reject the use of ES cells because it means the destruction
does resemble
people other proteins
Brazzeinofand a mabinlin,
prote
liin, for
normally found in healthy plants.
fo example, closely resemble “protein-
(Fig. 8B). For many years, scientists have known abouttotwo types
Theofnew potential human life. Adult stem cells are difficult to ase inhibitors,”
glean from proteins
eeins thatt can help prevent further damage
applications discuss scientific stem cells: embryonic stem (ES) cells and adult stem cells.
Embryonic stem cells are simply the cells of an early embry-
arise:
protein might
function diff erently
the human body, and they do not multiply readily in when a plant is injured.
the labora-
protein
ured. Interestingly, however, neither sweet
u
has that function.
ttion. Similar stories are true of most sweet-
tory. Also, their potential to become all different types of tissues
research and advances that have onic stage. These cells can stay alive longer and are
producing different tissues than adult stem cells, but
thanbetter
one.
the original
to acquire
at is not as great as that of ES cells. One drawback to both
proteinstem
and adult product
cells is the dangerproteinofproduct
rejection by the
tastingESproteins:
ordinary
cells Theyy closely
functions, b
recipient. but
clos resemble other plant proteins with
the sequences necessary for those other
ut th
Remember the many different types of proteins thatfunctions occur in the seem to bee missing
mis or mutated. It’s as though pre-
helped us gain valuable biological plasma membrane? Some of those mark the cell as existing
us, and if a transplanted tissue or organ carries different
belonging
Presumably
genes
to were recycled
eecycle to become genes for sweet proteins.
mark-a gene duplication
uplica
u in the distant past resulted in an
ers, our body works against them until they die. This “extra” gene that could
is called lld mutate
mu freely, while still leaving a “good”
knowledge. rejection of the transplant. copy of the gene to ssuppo
extra copy of the genee acq
support the plant’s functions. In time, the
acquired mutations that happened to pro-
Breakthrough vide a sweet taste, and nd pla
n plants with that mutation gained a spe-
human embryo human skin
cialtoappeal
becomefor local diners.
diners
d
All applications end with several By now, scientists are experienced at coaxing stem cells
specialized cells, but research would really benefit from an unlim-
ited source of stem cells in order to achieve the goal of replacing
F O R M YO U R O P I N I O N
Form Your Opinion questions that can diseased or damaged tissues in the human body. The scientific
community is now hopeful that such a source has been found, 1. Humans
H and perhaps apes and mon-
Stem cells thanks to a little-known Japanese scientist who worked alone for keys
k like sweet foods. How does
serve as a basis for class discussion. embryonic induced pluripotent
ten years in a tiny laboratory. Through patient research, Shinya
Yamanaka was able to discover why ES cells are pluripotent—able
this
t benefit plants containing sweet
proteins?
p
stem cells (ES) stem cells (iPS)
to become any type of tissue in the body. He hypothesized that 2. Are
A humans influencing the evolution
pluripotent cells produce certain proteins that specialized cells do of
o plants when they propagate them?
See page xxvii for a complete listing of not produce. Yamanaka worked with mouse skin cells until he
knew that only four particular genes do the trick of making cells
When
W they genetically modify them
Different and
a then propagate them?
In the second edition, the “Connecting the Concepts” Energy from the sun flows through all liv-
ing things with the participation of chlo-
Photosynthesis Cellular respiration
roplasts and mitochondria. Through the process of
feature appears at the end of each chapter. This feature photosynthesis, chloroplasts in plants and algae
capture solar energy and use it to produce carbo- sun carbohydrate
includes narrative and several questions to help students hydrates, which are broken down to carbon dioxide
and water in the mitochondria of nearly all organisms.
The energy released when carbohydrates (and other organic
O2
understand how the concepts in the present chapter are molecules) are oxidized is used to produce ATP molecules.
When the cell uses ATP to do cellular work, all the captured
energy dissipates as heat.
related to one another and to those in other chapters. During cellular respiration, oxidation by removal of
hydrogen atoms (e– + H+) from glucose or glucose prod-
chloroplast
heat
mitochondrion
ucts occurs during glycolysis, the prep reaction, and the Krebs
cycle. The prep reaction and Krebs cycle release CO2. The elec-
heat
trons are carried by NADH and FADH2 to the electron transport
chain (ETC) in the cristae of mitochondria. Oxygen serves as the CO2+H2O
final acceptor of electrons, and H2O is produced. The pumping of
hydrogen ions by the ETC into the intermembrane space leads to ATP
ATP production.
PUT THE PIECES TOGETHER
2. Explain the statement, “if chloroplasts and mitochondria
1. Tell how the pre-eukaryotic cell must have produced ATP. What are descended from a free-living common ancestor it would
event in the history of life would have allowed cellular respira- explain their structural similarities.” What are some structural
tion to evolve? Explain. similarities?
PREFACE v
Multilevel Perspective
Illustrations depicting complex structures
show macroscopic and microscopic views
to help you see the relationships between
increasingly detailed drawings.
Combination Art
Drawings of structures are paired with micrographs to
give you the best of both perspectives: the realism of
photos and the explanatory clarity of line drawings.
vi PREFACE
Sporangium
Process Figures
Complex processes are broken down into a series of
Mitosis
smaller steps that are easy to follow. Numbers guide
stalk
Sporophyte you through the process.
zygote
In process figures, numbered steps are coordinated with
diploid (2n)
FERTILIZATION
haploid (n)
MEIOSIS the narrative for an integrated approach to learning.
2 Fertilization:
Flagellated sperm egg Spores
sperm 5 Spore dispersal:
produced in
Mitosis Spores are released
antheridia swim in
external water to when they are most
archegonia, each likely to be dispersed
onia
Archegonia
bearing a single egg. by air currents.
archegonium buds
Antheridia
10.6 During translation, polypeptide synthesis occurs one amino acid at a time
Although we often speak of protein synthesis, some proteins
have more than one polypeptide, so it is more accurate to recog- amino acid methionine large ribosomal subunit
nize that polypeptide synthesis occurs at a ribosome. Polypep-
tide synthesis involves three events: initiation, elongation, and met
E site P site A site
termination. Enzymes are needed so that each of the three events
will occur, and both initiation and elongation also require an
input of energy.
U A
5ʹ A U C mRNA met
Initiation During initiation all translation components come G
TRANSCRIPTION ing site. The start codon AUG is at the P site. The fi
TRANSLATION rst, or initia-
first,
1 DNA in nucleus serves as
a template for mRNA. tor, tRNA pairs with this codon because its anticodon is UAC. An mRNA binds to a small The large ribosomal subunit
As you can see by examining Figure 10.4B, AUG is the codon ribosomal subunit. An initiator completes the ribosome.
for methionine. Methionine is always the3fi
DNA mRNA moves intoof a tRNA with the anticodon UAC Initiator tRNA occupies the
rst
first amino acid
2 mRNA is processed large and small cytoplasm and pairs with the mRNA start P site. The A site is ready
polypeptide. After the small ribosomal
ribosomal subunits 5′ unit has attached,
becomes associated a codon AUG. for the next tRNA.
before leaving the nucleus.
mRNA large ribosomal subunit joins to the small with subunit. Although
ribosomes.
introns
primary similar in many ways, initiation in eukaryotes is much more
mRNA 3′ complex. FIGURE 10.6A Participants in the initiation event assemble as
mature amino shown. The first amino acid is typically methionine.
4 tRNAs with
mRNA acids
nuclear pore anticodons carry
amino acids
to mRNA.
ribosome tRNA
U A
U A C C
polypeptide
5′ A U G 3′
anticodon Color Consistency
codon
5 During initiation, anticodon-codon
Consistent use of color organizes
8 During termination, a
CC
C
complementary base pairing
begins as the ribosomal subunits ribosome reaches a stop information and clarifies concepts.
come together at a start codon. codon; mRNA and
ribosomal subunits
anticodon
5′
C C C UG G U U U disband.
G G G A C C A A A G U A
3′
PREFACE vii
“The organization of the text around the Chromosome Anomalies Can Be Inherited
8.13 Nondisjunction causes chromosome number
anomalies 162
major theories of Biology is a wise path 8.14 Chromosome number anomalies can be
observed 163
8.15 Chromosome structure anomalies can also be
to follow; it integrates the chapters into observed 164
APPLI C AT I O N S
themes and points out the development HOW LIFE CHANGES
Evolution of the Spindle Apparatus 147
HOW BIOLOGY IMPACTS OUR LIVES
of a theory. . . . Tissues Can Be Grown in the Lab 151
HOW BIOLOGY IMPACTS OUR LIVES
Protective Behaviors and Diet Help Prevent Cancer 154 Colon cancer cell
they are to know. occurs in the life cycle of organisms determines the adult chromosome number. Even though meiosis is different from mitosis, the two
processes bear certain similarities.
Section Introductions orient students to concepts in a many chromosomes as the parent cell.
In Figure 8.7, the diploid (2n) number of chromosomes is four,
and there are two pairs of chromosomes. The short chromosomes
centrioles
homologous 2n = 4
chromosomes
short, easy-to-understand manner. are one pair, and the long chromosomes are another. They are
homologous chromosomes (also called homologues) because they
look alike and carry genes for the same traits, such as finger length.
However, one homologue could call for short fingers and the other CHROMOSOME DUPLICATION
for long fingers.
Prior to the first division, called meiosis I, DNA replication has synapsis
occurred, and the chromosomes are duplicated. During meiosis I,
the homologous chromosomes come together and line up side by
side. This so-called synapsis results in an association of four chro-
matids that stay in close proximity during the first two phases of 2n = 4
meiosis I. Also, because of synapsis, there are pairs of homologous sister
chromosomes at the equator during meiosis I. (Keep in mind that chromatids
only during meiosis I is it possible to observe paired chromosomes
Figure Legends have been expanded in this edition at the equator.) Synapsis leads to a reduction in the chromosome
number because it permits orderly separation of homologous chro-
MEIOSIS I
Duplicated
homologous pairs
mosomes. The daughter nuclei are haploid because they receive
to reinforce the discussion and to improve student only one member of each pair. The haploid (n) nature of each
daughter cell can be verified by counting its centromeres. Each
synapse and
then separate.
learning. DNA occurs between meiosis I and meiosis II. The period of time
between meiosis I and meiosis II is called interkinesis.
During meiosis II, the sister chromatids of each chromo-
some separate, becoming daughter chromosomes that are dis- MEIOSIS II
tributed to daughter nuclei. In the end, each of four daughter Sister chromatids
cells has the n, or haploid, number of chromosomes, and each separate, becoming
daughter chromosomes.
chromosome consists of one chromatid.
In humans, the daughter cells mature into gametes (sex
cells—sperm and egg) that fuse during fertilization. Fertiliza-
tion restores the diploid number of chromosomes in the zygote,
the first cell of the new individual. If the gametes carried the
diploid instead of the haploid number of chromosomes, the
Check Your Progress questions at the end of chromosome number would double with each fertilization. After
several generations, the zygote would be nothing but
n=2 n=2
chromosomes. FIGURE 8.7 Meiosis produces daughter cells that are genetically
each section help students assess and/or apply 8.7 Check Your Progress At the completion of meiosis I, are the
different from the parent cell. Four daughter cells result because
meiosis includes two divisions: During meiosis I, the homologous
chromosomes separate, and during meiosis II the chromatids
cells diploid (2n) or haploid (n)? Explain.
separate, becoming daughter chromosomes.
their understanding of a concept.
CHAPTER 8 Cell Division and Reproduction 155
viii PREFACE
aid to learning these terms in a nonthreatening way. • Humans have 22 pairs of autosomes and one pair of sex
chromosomes. A Y chromosome is shorter than an X
chromosome. Males are XY and females are XX.
(telophase).
• In plants and animals, mitosis allows growth and repair. In
humans, adult stem cells undergo mitosis to replace worn-out
• Following DNA replication, each chromosome has two sister cells.
chromatids held together at a centromere. Kinetochores 8.5 Cytokinesis divides the cytoplasm
which develop at centromeres function during cell division.
• In animal cells, cytokinesis involves a cleavage furrow.
• Homologus chromosomes have genes for the same trait—e.g.,
• In plant cells, cytokinesis involves the formation of a new
type of hairline. Sister chromatids have exact genes—e.g.,
plasma membrane and cell wall at a cell plate.
widow’s peak.
8.2 The eukaryotic cell cycle has a set series of stages Cancer Is Uncontrolled Cell Division
• In the cell cycle, interphase (G1, S, G2 stages) precedes the M 8.6 Cell cycle control is lacking in cancer cells
stage, which includes mitosis and cytokinesis. • Checkpoint G1 involves cell cycle control; checkpoint G2
ensures that DNA replicated properly; checkpoint M ensures
G0 that chromosomes are distributed accurately to daughter cells.
• In general, if the cell cycle is unable to continue, apoptosis
occurs. Apoptosis initiated by p53 is programmed cell death
Int orchestrated by unleashed enzymes.
er
G1 p h • Due to mutations, carcinogenesis occurs and cancer is
Cyto (growth)
as
kine present when cells divide uncontrollably and a tumor
e
sis
S
develops. Cell cycle control and apoptosis are lacking.
tosis
M (DNA synthesis) • Cancer cells have abnormal characteristics: lack
differentiation, have abnormal nuclei, form tumors,
Mi
undergo metastasis (formation of tumors distant from
G2
primary tumor), and promote angiogenesis (formation of
(growth) new blood vessels).
(n), depending
lloid (n
a tetrad) and crossing-over (exchange of genetic material)
between nonsister chromatids occurs.
2. Which is a correct contrast between autosomes and sex 8.9 Sexual reproduction increases genetic variation
apply their understanding of a concept. meiosis II. No DNA replication occurs.
• Meiosis II: During stages designated by the Roman numeral
II, the chromatids of duplicated chromosomes from meiosis I
chromosomes in humans?
a. 22 pairs—one pair
on the species.
• The number of centromeres equals the number
chromosomes a cell has.
mber of
m • Crossing-over recombines genetic information and increases
the variability of genetic inheritance on the chromosomes.
separate, producing a total of four daughter cells for meiosis. b. control gender—control enzymes • The gametes contain all possible combinations of
• Centrosomes form the spindle apparatus, which w helps ensure
c. are always duplicated—are always single chromosomes because of independent assortment.
8.11 Life cycles are varied orderly separation of chromatids.
d. are always visible—are never visible
• Asexual reproduction results in offspring that are genetically 3. In the cell cycle,
identical to the single parent. a. mitosis cannot occur without interphase.
• In the haploid life cycle, asexual reproduction occurs when b. the single event during interphase is chromosome CHAPTER 8 Cell Division and Reproduction 165
a haploid parent produces offspring by mitosis that are also duplication.
haploid. In sexual reproduction, only the zygote is diploid and c. cells are metabolically inactive during interphase.
undergoes meiosis to produce haploid offspring. Algae and d. a DNA double helix divides in two.
fungi often have the haploid life cycle.
• In the alternation of generations life cycle, which usually Mitosis Maintains the Chromosome Number
occurs in plants, the diploid sporophyte produces spores by 4. The two identical halves of a duplicated chromosome
meiosis. A spore undergoes mitosis to become a gametophyte, a. always stay together. c. become daughter chromosomes.
which produces gametes. When the gametes fuse, the diploid b. are different sizes. d. are called homologues.
zygote becomes a sporophyte.
• In the diploid life cycle, which usually takes place in animals, For questions 5–8, match each description to a phase of mitosis in
the diploid adult produces gametes by meiosis, which are the key.
the only haploid part of the life cycle. Mitosis is involved in
“It would be fun to teach and growth.
8.12 Meiosis can be compared to mitosis
KEY:
a. prophase
b. metaphase
c. anaphase
d. telophase
learn using this book.” • See Figure 8.12 and note that homologous chromosomes
only pair during metaphase I of meiosis and that four haploid
5. The nucleolus disappears, and the nuclear envelope
breaks down.
6. The spindle disappears, and the nuclear envelopes form.
daughter cells result from meiosis but not mitosis.
PREFACE ix
The second edition of Concepts of Biology continues to pre- Part III: Organisms Are Related and Adapted to Their
sent concepts clearly and make biology relevant through the Environment The evolution chapters include extensive revi-
use of excellent writing, instructional art, and effective ped- sions. Chapter 14, Speciation and Evolution, now includes real-
agogical tools. This new edition also includes life examples of various processes that can cause speciation.
• Significant content changes, as outlined below. The influence of regulatory genes during development helps
explain how species can share the same genes but have differ-
• Enhanced evolutionary coverage, including extensive
ent phenotypes. Chapter 15, The Evolutionary History of Life on
updates to the evolution chapters and the addition of
Earth, will help instructors introduce their students to cladis-
How Life Changes applications.
tics. It explains the rationale behind the replacement of Lin-
• Media integration, including a robust set of teaching and nean classification with that of cladistics in a way that allows
learning tools through McGraw-Hill’s Connect™ Biology. instructors to be up-to-date, while not overburdening begin-
ning students. Chapter 17, Evolution of Protists, introduces a
Content Changes new evolutionary tree of protists based on molecular data. The
Chapter 1, Biology, the Study of Life, was rewritten. It begins chapter still emphasizes the biological and ecological relevance
with a discussion of the scientific process and proceeds to of each type of protist. Chapter 18, Evolution of Plants and
an overview of the major concepts of biology (cell theory, Fungi, employs a new evolutionary tree based in part on molec-
gene theory, theory of evolution, theory of homeostasis, and ular data. Land plants and stoneworts, which are charophytes,
theory of ecosystems). Basic evolutionary principles are pre- share a common green algal ancestor. All land plants protect the
sented, and a depiction of the Tree of Life introduces the embryo, and thereafter each of four innovations can be associ-
three domains of life and the various types of eukaryotes. ated with a particular group of land plants. Chapter 19, Evolu-
Part I: Organisms Are Composed of Cells Chapter 4, tion of Animals, introduces the new evolutionary tree of
Structure and Function of Cells, presents an improved dis- animals based on molecular and developmental data. The biol-
cussion on cell structure and two new tables. Table 4.4 sum- ogy of each group is discussed as before. Chapter 20, Evolution
marizes the differences between plant and animal cells, and of Humans, was rewritten to include a description and impor-
Table 4.16 summarizes the eukaryotic cell structures and tance of the newly studied fossil Ardipithicus ramidus.
their functions. A new figure (Fig. 4.13) stresses that plant Part IV: Plants Are Homeostatic Chapter 21, Plant Orga-
cells have both mitochondria and chloroplasts. Chapter 5, nization and Homeostasis, includes a rewrite of Section 21.1
Dynamic Activities of Cells, includes sharpened energy to better explain the overall organization of a plant and the
transformation analogies with references to everyday occur- functions of roots, stems, and leaves. This supports a rewrite
rences (see Figs. 5.1, 5.3B, and 5.5A). Cell communication of Section 21.8 which discusses more authoritatively how
was stressed with the addition of a new section (Section plants maintain homeostasis. Chapter 22, Transport and
5.13). Chapter 7, Pathways of Photosynthesis, provides an Nutrition in Plants, includes a reorganization of Sections 22.4
improved discussion of mitochondrial structure. Changes to and 22.6 to increase student understanding of phloem struc-
Figures 7.4 and 7.7B offer views from the whole cell to par- ture and function, and root structure and function. Chap-
ticles on the mitochondrial inner membrane. ter 24, Reproduction in Plants, was reorganized and rewritten
Part II: Genes Control the Traits of Organisms All to better present an overview of the flowering plant life cycle
chapters in Part II were rewritten to present concepts at a and place it in an evolutionary context.
student-friendly pace, and thereby increase student interest Part V: Animals Are Homeostatic Chapter 26, Coordina-
and learning. Modern genetics has been updated. Chapter 11, tion by Neural Signaling, was reorganized and begins with
Regulation of Gene Activity, is now at an appropriate level an overview of the structure and function of the human ner-
and explains how humans can make do with far fewer pro- vous system, before comparing this system to that of other ani-
tein-coding genes than expected. Chapter 12, Biotechnology mals. Chapter 30, Lymph Transport and Immunity, includes
and Genomics, offers a short, but complete, discussion of the an updated and rewritten discussion of immunity to be con-
human genome, including the several types of DNA sequences sistent with current immunity literature. Chapter 32, Gas
that are not protein-coding genes. Exchange and Transport in Animals, provides a discussion
My Lectures—Tegrity
Tegrity Campus™ records and distributes your class lecture,
with just a click of a button. Students can view anytime/
anywhere via computer, iPod, or mobile device. It indexes
as it records your PowerPoint presentations and anything
shown on your computer so students can use keywords to
fi nd exactly what they want to study. Tegrity is available as
an integrated feature of McGraw-Hill Connect™ Biology or
as standalone.
xi
xiii
xiv ACKNOWLEDGMENTS
ACKNOWLEDGMENTS xv
xvi
CONTENTS xvii
xviii CONTENTS
CONTENTS xix
xx CONTENTS
CONTENTS xxi
xxii CONTENTS
Viewpoint The Eyes Have It 548 Skeletal Muscle Cells Contract Individually 582
28.9 A muscle cell contains many myofibrils 582
Sensory Receptors Respond to Stimuli 550
28.10 Sarcomeres shorten when muscle cells contract 582
27.1 Sensory receptors can be divided into five
28.11 Axon terminals bring about muscle cell contraction 583
categories 550
28.12 A muscle cell has three sources of ATP
27.2 Sensory receptors communicate with the CNS 551
for contraction 584
Chemoreceptors Are Sensitive to Chemicals 552 28.13 Some muscle cells are fast-twitch and some are slow-
27.3 Chemoreceptors are widespread in the animal twitch 584
kingdom 552
27.4 Mammalian taste receptors are located in the
mouth 552
29 Circulation and Cardiovascular
27.5 Mammalian olfactory receptors are located in
Systems 588
the nose 553 Viewpoint Not All Animals Have Red Blood 588
Photoreceptors Are Sensitive to Light 554 A Circulatory System Helps Maintain Homeostasis 590
27.6 The vertebrate eye is a camera-type eye 554 29.1 A circulatory system serves the needs of cells 590
27.7 The lens helps bring an object into focus 555 The Mammalian Heart Has Four Chambers 592
27.8 The retina sends information to the visual cortex 556 29.2 The mammalian heart is a double pump 592
Mechanoreceptors Are Involved in 29.3 The heartbeat is rhythmic 594
Proprioception and Sense of Touch 558 Blood Vessels Transport Blood 595
27.9 Proprioceptors are located in skeletal muscles, and
29.4 The structure of blood vessels is suited to their
touch receptors are located in skin 558
function 595
Mechanoreceptors Are Involved in 29.5 Blood vessels form two circuits in mammals 598
Hearing and Balance 559 29.6 Blood pressure is essential to the flow of blood in each
27.10 The mammalian ear has three well-developed circuit 599
regions 559
Blood Has Vital Functions 600
27.11 Hearing receptors are in the inner ear 560
29.7 Blood is a liquid tissue 600
27.12 Balance receptors are also in the inner ear 562
29.8 Exchanges between blood and tissue fluid occur at
capillaries 602
28 Locomotion and Support 29.9 Blood clotting involves platelets 603
Systems 568 29.10 Blood types must be matched for transfusions 604
CONTENTS xxiii
xxiv CONTENTS
CONTENTS xxv
Viewpoint Life Under Glass 780 The Causes of Today’s Extinctions Are Known 805
40.5 Habitat loss is a major cause of wildlife extinctions 805
Climate Is Dictated by Temperature and Rainfall 782
40.6 Introduction of alien species contributes to
39.1 Solar radiation and winds determine climate 782 extinctions 806
39.2 Topography and other effects also influence 40.7 Pollution contributes to extinctions 807
climate 783
40.8 Overexploitation contributes to extinctions 809
39.3 Ocean currents affect climate 784
40.9 Disease contributes to extinctions 809
On Land, the Biosphere Is Organized
Habitat Preservation and Restoration Require
into Terrestrial Ecosystems 785
Much Effort and Expertise 810
39.4 Major terrestrial ecosystems are characterized
40.10 Habitat preservation is of primary importance 810
by particular climates 785
40.11 Habitat restoration is sometimes necessary 811
39.5 The tundra is cold and dark much of the year 786
39.6 Coniferous forests are dominated by A Sustainable Society Is Achievable 813
gymnosperms 786 40.12 A sustainable society will preserve resources 813
39.7 Temperate deciduous forests have abundant life 788 40.13 Energy sources should be renewable 814
39.8 Temperate grasslands have extreme seasons 788 40.14 Water sources should be conserved 815
39.9 Savannas have wet-dry seasons 789 40.15 Agriculture can be more diverse 816
39.10 Deserts have very low annual rainfall 789 Appendix
39.11 Tropical rain forests are warm with abundant Answer Key A-1
rainfall 790
Fresh Water and Salt Water Are Organized Glossary G-1
into Aquatic Ecosystems 791
Credits C-1
39.12 Fresh water flows into salt water 791
Index I-1
39.13 Marine ecosystems include those of the coast and
the ocean 792
xxvi CONTENTS
APPLI C AT I O N S
HOW BIOLOGY IMPACTS OUR LIVES
Organic Farming 6
HOW LIFE CHANGES
Evolution’s Many Applications 17
queen
larvae
workers
eggs
H O W B I O L O G Y I M PAC T S O U R L I V E S Application
1A Organic Farming
Besides being health con- oxygen in the blood. In adults, nitrates are implicated in causing
scious, people who buy digestive tract cancers. Certainly they can cause an algal bloom,
organic may also be socially recognized as a green scum on the water’s surface.
conscious. Organic farming is In response to these problems, organic farmers severely limit
part of a movement to make the use of nitrogen fertilizers and instead rely on crop rotation,
agriculture sustainable by alternately planting a nitrogen-providing
using farming methods that legume and a nitrogen-requiring crop such
protect the health of people as wheat. Organic farmers also cut way
and ecosystems and preserve back on the use of herbicides and pesti-
the land so that it can be pro- cides, and this may be the primary reason
ductive for our generation and all future generations. you and others buy organic. The long-term
Modern agricultural methods have been dramatically suc- consumption of these chemicals has been
cessful at increasing yield, but at what price? We now know that associated with such health problems as
modern farming practices lead to topsoil depletion and ground- birth defects, nerve damage, and cancer.
water contamination. Without topsoil, the nutrient-rich layer Children may be especially sensitive to
that nourishes plants, agriculture is impossible, and yet modern health risks posed by pesticides; this is the FIGURE 1A
farming practices such as tilling the land and allowing it to lie chief reason lawns sprayed with pesticides Legume plants have
fallow (bare) allow topsoil to erode and disappear. One solution carry warning signs. We should all be nodules.
is to use a legume as a ground cover because it both protects and aware that we too can contribute to an
nourishes the soil (Fig. 1A). The researchers who did the study organic lifestyle by limiting the use of synthetic chemicals on our
described in Section 1.2 used pigeon peas as a way to enrich the lawns and gardens. In doing so, we improve our health and help
soil between winter wheat plantings. preserve the environment for ourselves and future generations.
Instead of growing legumes, farmers in recent years are
F O R M YO U R O P I N I O N
accustomed to making plants bountiful by applying more and
more synthetic nitrogen fertilizer. Unfortunately, nitrogen fertil- 1. The United States exports its current farming technology,
izers pollute wells used for drinking water and also huge bodies with all its long-range problems, to other countries. Should
of water, such as the Chesapeake Bay, the Gulf of Mexico, and this be continued?
the Great Lakes. Nitrates in the drinking water of infants leads 2. What circumstances might discourage a farmer from growing
to the “blue-baby” syndrome and possible death due to lack of food organically, and how might these obstacles be overcome?
20
Control Pots
= no treatment
Test Pots
15 = fertilization treatment
10
0
year 1 year 2 year 3
The results
pots preplanted with pigeon peas did not produce as high a yield Analysis of Results After two years, the yield from pots
as the control pots. treated with nitrogen fertilizer was less than it had been the first
year. Indeed, wheat yield in pots following a summer planting of
Conclusion The hypothesis was not supported. Wheat
pigeon peas was the highest of all the treatments. After three
yield following the growth of pigeon peas was not as great
years, wheat yield in pots treated with nitrogen fertilizer was
as that obtained with nitrogen fertilizer treatments.
greater than in the control pots but not nearly as great as the
yield in pots following summer planting of pigeon peas. Com-
Follow-Up Experiment and Results The researchers
pared to the first year, wheat yield increased almost fourfold in
decided to continue the experiment, using the same design and
pots having a pigeon pea/winter wheat rotation.
the same pots as before, to see whether the buildup of residual
soil nitrogen from pigeon peas would eventually increase wheat Conclusion The hypothesis was supported. At the end
yield to a greater extent than the use of nitrogen fertilizer. This of three years, the yield of winter wheat following a
was their new hypothesis: pigeon pea/winter wheat rotation was much better than
for the other types of test pots.
Hypothesis A sustained pigeon pea/winter wheat rota-
tion will eventually cause an increase in winter wheat To explain their results, the researchers suggested that the soil
production. was improved by the buildup of the organic matter in the pots
as well as by the addition of nitrogen from the pigeon peas.
They predicted that wheat yield following three years of pigeon
They published their results in a scientific journal,1 where their
pea/winter wheat rotation would surpass wheat yield following
experimental method and results would be available to the
nitrogen fertilizer treatment.
scientific community.
1.2 Check Your Progress What would your control group and test
groups be composed of if you were testing whether a parasite could
1
Bidlack, J. E., Rao, S. C., and Demezas, D. H. 2001. Nodulation, nitrogenase
activity, and dry weight of chickpea and pigeon pea cultivars using different
reduce the size of a fire ant colony?
Bradyrhizobium strains. Journal of Plant Nutrition 24:549–60.
Biosphere
Regions of the Earth’s crust,
waters, and atmosphere inhabited
by living things
Ecosystem
A community plus the physical environment
Community
Interacting populations in a particular area
Population
Organisms of the same species
in a particular area
Organism
An individual; complex individuals
contain organ systems
Organ System
Composed of several organs
working together
Organ
Composed of tissues functioning
together for a specific task
Tissue
A group of cells with a common
structure and function
Cell
The structural and functional
unit of all living things
Molecule
Union of two or more atoms
of the same or different elements
Atom
Smallest unit of an element composed of
electrons, protons, and neutrons
cell
a. b. c.
FIGURE 1.3B Only micrographs (pictures taken microscopically), such as the one in (c), can reveal that organisms are composed of cells.
CHAPTER 1 Biology, the Study of Life 9
ing) components of the environment. One example of an ecosys- Solar Chemical cycling
tem is a North American grassland, which is inhabited by energy
populations of rabbits, hawks, and many other animals, as well
as various types of grasses. These populations interact by form-
heat
ing food chains in which one population feeds on another. For
example, rabbits feed on grasses, while hawks feed on rabbits
and other organisms. producers
As Figure 1.6 shows, ecosystems are characterized by chem-
ical cycling and energy flow, both of which begin when plants,
such as grasses, take in solar energy and inorganic nutrients to heat
produce food (organic nutrients) by photosynthesis. Chemical
cycling (gray arrows) occurs as chemicals move from one popu-
consumers
lation to another in a food chain, until death and decomposition
allow inorganic nutrients to be returned to the photosynthesizers
once again. Energy (red arrows), on the other hand, flows from
the sun through plants and the other members of the food chain
as they feed on one another. The energy gradually dissipates and
returns to the atmosphere as heat. Because energy does not
cycle, ecosystems could not stay in existence without solar
energy and the ability of photosynthesizers to absorb it.
The Biosphere Climate largely determines where different
ecosystems are found in the biosphere. For example, deserts
exist in areas of minimal rain, while forests require much rain.
The two most biologically diverse ecosystems—tropical rain for-
ests and coral reefs—occur where solar energy is most abun- decomposers heat
dant. The human population tends to modify these and all
ecosystems for its own purposes. Humans clear forests or grass-
lands in order to grow crops; later, they build houses on what FIGURE 1.6 A grassland is a major ecosystem. Chemicals cycle
was once farmland; and finally, they convert small towns into because decomposers return inorganic nutrients to producers which
cities. As coasts are developed, humans send sediments, sewage, provide organic nutrients to consumers including decomposers. With
and other pollutants into the sea. each transfer of nutrients, energy is lost as heat.
Tropical rain forests and coral reefs are home to many organ-
isms. The canopy of the tropical rain forest alone supports a It has long been clear that human beings depend on healthy
variety of organisms, including orchids, insects, and monkeys. ecosystems for food, medicines, and various raw materials. We
Coral reefs, which are found just offshore in the Southern Hemi- are only now beginning to realize that we depend on them even
sphere, provide a habitat for many animals, including jellyfish, more for the services they provide. The workings of ecosystems
sponges, snails, crabs, lobsters, sea turtles, moray eels, and some ensure that environmental conditions are suitable for the contin-
of the world’s most colorful fishes. Like tropical rain forests, ued existence of humans.
coral reefs are severely threatened as the human population
increases in size. Aside from pollutants, overfishing and collec- 1.6 Check Your Progress Give a specific example to illustrate that
tion of coral for sale to tourists destroy the reefs. we depend on natural ecosystems.
Crocodilians
reptile characteristics
bird characteristics
feathers Dinosaurs
common
ancestor
(reptilian)
Dinosaurs
teeth
tail with vertebrae
Archaeopteryx
common ancestor
claws
Past Present
Time
Deer prefer a diet of smooth leaves over hairy leaves. Plants with
hairy leaves reproduce more than other plants in the population.
FIGURE 1.9A Predatory deer act as a selective agent to bring FIGURE 1.9B Rockhopper penguins are adapted to swimming in
about change in a plant population. the icy waters of Antarctica.
KEY:
a. is organized d. is homeostatic
Experiments and/or
b. uses materials e. responds to stimuli observations c.
and energy f. forms ecosystems
c. reproduces g. evolves
28. organisms exhibit behavior
29. populations interact Conclusion d.
30. giraffes produce only giraffes
31. common descent with modification
The scientific method consists of making observa- composed of cells and that cells are the fundamental units of life.
tions, formulating a hypothesis, testing the hypoth- The theory of homeostasis tells us that all organisms have mecha-
esis, and coming to a conclusion on the basis of the results (data). nisms that allow them to keep their internal environment rela-
The conclusions of many studies have allowed scientists to develop tively constant. The gene theory tells us all organisms have genes,
the five theories (cell theory, gene theory, theory of homeosta- hereditary units that undergo mutations leading to the variety of
sis, theory of ecosystems, and theory of evolution) on which this life. Even so, all life-forms share similar characteristics because
book is based. Theories are conceptual schemes that tell us how they can trace their ancestry to a common source as stated by the
the world works. All theories of biology are related. For example, theory of evolution. All life-forms live in ecosystems where inter-
the gene theory is connected to the theory of evolution because actions allow them to acquire the materials and energy they need
mutations create differences between the members of a popula- to continue their existence. Human beings are also dependent on
tion. Better-adapted members have the opportunity through natural ecosystems, and when they preserve the biosphere, they are pre-
selection to reproduce more, and in that way a species becomes serving their own existence as well.
adapted to its environment.
Any two theories are related. For example, evolution is also con-
PUT THE PIECES TOGETHER
nected to the theory of ecosystems because, as natural selection occurs,
species become adapted to living in a particular ecosystem. We can 1. Give your own example (not taken from this reading) to show
connect this observation to the cell theory because, if a gazelle’s nerve that two theories are related.
cells can conduct nerve impulses faster to its muscle cells than a lion’s 2. Explain in your own words how bacteria become resistant to an
nerve cells, the gazelle is more likely to escape capture. antibiotic.
In exploring the theories, we have also discussed the char-
acteristics of life. The cell theory taught us that all organisms are
C H AP T E R O U T LI N E
Cells Are the Basic Units of Life
4.1 All organisms are composed of cells 64
4.2 Metabolically active cells are small in size 65
4.3 Prokaryotic cells evolved first 67
4.4 Eukaryotic cells contain specialized organelles:
An overview 68
Protein Synthesis Is a Major Function of Cells
4.5 The nucleus contains the cell’s genetic
information 70
4.6 The ribosomes carry out protein synthesis 71
4.7 The endoplasmic reticulum synthesizes and
transports proteins and lipids 72
4.8 The Golgi apparatus modifies and repackages
proteins for distribution 73
Vacuoles and Vesicles Have Varied Functions
4.9 Lysosomes digest biomolecules and cell parts 74
4.10 Peroxisomes break down long-chain fatty acids 74
4.11 Vacuoles are common to plant cells 74
Hooke saw almost nothing except for outlines, which we
4.12 Vesicles allow the organelles of the
endomembrane system to work together 75 know today are the cell walls of plant cells. Similarly, you can
make out the demarcations between onion root cells in the
A Cell Carries Out Energy Transformations micrograph on page 63. After comparing these to the nerve
4.13 Chloroplasts and mitochondria have opposite cells below, you might conclude that a cell is an entity, a unit
functions 76 of a larger whole.
The Cytoskeleton Is Dynamic
4.14 The cytoskeleton maintains cell shape and assists
movement 78
4.15 Cilia and flagella permit movement 79
Cell Structures Work Together
Table 4.16 Eukaryotic Cell Structures 80
APPLI C AT I O N S
HOW SCIENCE PROGRESSES
Microscopes Allow Us to See Cells 66
HOW SCIENCE PROGRESSES
Pulse-labeling Allows Observation of the Secretory
Pathway 73
HOW LIFE CHANGES
How the Eukaryotic Cell Evolved 77
Animal nerve cells
62
63
Lilac, a plant
Rabbit, an animal
50 mm
Micrograph of leaf reveals cells.
FIGURE 4.1A Lilac leaf, with a photomicrograph below. Micrograph of intestine reveals cells. 140 mm
protein
chloroplast
plant and mouse rose
frog egg
animal
amino cells
acid
virus
ostrich
most bacteria human egg ant egg
atom
blue whale
electron microscope human
light microscope
human eye
FIGURE 4.2A The sizes of living things and their components. In the metric
system (see back endsheet), the units are based on the power of 10.
(1 meter = 102 cm = 103 mm = 106 μm = 109 nm.)
protein flagellum
molecules
4.3 Check Your Progress Why can’t you define or recognize a cell
FIGURE 4.3B A prokaryotic cell is structurally simple but
by the presence of a nucleus?
metabolically complex.
Plasma membrane:
outer surface that
regulates entrance and FIGURE 4.4A Animal cell anatomy.
exit of molecules
protein
phospholipid
NUCLEUS: command center of cell
Nuclear envelope: double
membrane that encloses nucleus
Chromatin: diffuse threads
Cytoskeleton: protein fibers that containing DNA
maintain cell shape and assist Nucleolus: region that produces
movement of cell parts ribosomal subunits
Mitochondrion: organelle
that carries out cellular
Vesicle: membrane-bounded respiration, producing
sac that stores and transports ATP molecules
substances
Polyribosome: string of
*Lysosome: vesicle that ribosomes
digests biomolecules
and even cell parts
Golgi apparatus: processes, packages,
*Not found in plant cells and secretes modified cell products
Ribosomes: carry
out protein synthesis *Chloroplast: carries
out photosynthesis,
Centrosome:
producing sugars
microtubule organizing
center (lacks centrioles)
nuclear
envelope
nucleolus
DNA
Cytoplasm
mRNA
2 mRNA attaches to ribosome and
polypeptide synthesis begins.
receptor
ER membrane
protein
Endoplasmic
reticulum (ER) 71
rough
endoplasmic
reticulum
(RER)
interior
of ER
smooth
endoplasmic
reticulum
(SER)
0.08 µm
1. Why would Palade have labeled sulfur and not carbon in FIGURE 4B The secretory pathway.
the amino acids? (See Fig. 3.9A.)
2. Where else might Palade have found the labeled amino
acids in the cell? (See Fig. 4.6.)
4.12 Vesicles allow the organelles of the endomembrane system to work together
The endomembrane system includes various membranous saliva or when the pancreas produces digestive enzymes. Simi-
organelles that work together and communicate by means of larly, lipids move from the smooth ER to the Golgi apparatus and
transport vesicles. It includes the endoplasmic reticulum (ER), can eventually be secreted.
the Golgi apparatus, lysosomes, and the transport vesicles. 5 In animal cells, lysosomes produced by the Golgi appa-
Figure 4.12 shows how the components of the endomem- ratus 6 fuse with incoming vesicles from the plasma mem-
brane system work together: 1 Proteins, produced in the rough brane and digest biomolecules and debris. White blood cells
ER, are carried in 2 transport vesicles to 3 the Golgi appara- are well-known for engulfing pathogens (e.g., disease-causing
tus, which sorts the proteins and packages them into vesicles viruses and bacteria) that are then broken down in
that transport them to various cellular destinations. lysosomes.
4 Secretory vesicles take the proteins to the plasma mem-
brane, where they exit the cell when the vesicles fuse with the 4.12 Check Your Progress What parts of the cell are responsible
membrane. This is called secretion by exocytosis. For example, for producing and exporting the proteins found in the endomem-
brane system?
secretion into ducts occurs when the salivary glands produce
secretion
FIGURE 4.12 The organelles plasma
membrane
of the endomembrane system.
incoming vesicle
incoming vesicle
4 secretory vesicle
5
3 Golgi apparatus
lysosome
protein
2 transport vesicle
transport vesicle
lipid
1 rough endoplasmic
reticulum
smooth endoplasmic
reticulum
ribosome Nucleus
mitochondrion
200 nm
cristae
chloroplast
thylakoids
1.5 µm
FIGURE 4.13 Plant cells carry on photosynthesis in green leaves where chloroplasts absorb solar energy because they contain the green
pigment chlorophyll in thylakoid membranes. Mitochondria in plant and animal cells carry on cellular respiration, a process that produces ATP on
the membranous invaginations called cristae.
76 PART 1 Organisms Are Composed of Cells
FIGURE 4.14B The motor molecule kinesin is moving a vesicle 4.14 Check Your Progress A cell is dynamic. In general, what
along a microtubule track. accounts for the ability of cell contents to move?
Ribosomes Small particles, each with two subunits Are sites of protein synthesis
Storage, Transport, Vesicle Tiny membranous sac Stores and transports proteins and lipids
and Digestion
Cell Shape and Plasma Phospholipid bilayer with embedded Regulates entrance and exit of molecules
Movement membrane proteins into and out of cell
Cell wall In plant cells, outer layer of cellulose Helps maintain shape of cell, protects and
supports
Cytoskeleton Network of protein fibers Supports organelles, assists movement of
cell and its parts
Flagella and cilia Microtubule-containing cellular extensions Move the cell or move substances along
its surface
Our knowledge of cell anatomy has been gathered by Table 4.16 shows you other ways to group the organelles.
studying micrographs of cells. This has allowed cytol- Lysosomes and peroxisomes are vesicles with digestive functions:
ogists (biologists who study cells) to arrive at a generalized picture Lysosomes digest various biomolecules while peroxisomes break
of cells, such as those depicted for an animal and a plant cell in Sec- down lipids.
tion 4.4. Eukaryotic cells, taken as a whole, contain several types of The origin of the eukaryotic cell links together what you know
organelles, and the learning outcomes for the chapter suggest that about the structure of prokaryotic and eukaryotic cells because the
you should know the structure and function of each one. A concept endosymbiotic theory says that mitochondria and chloroplasts were
to keep in mind is that “structure suits function.” For example, ribo- once free-living prokaryotes.
somal subunits move from the nucleus to the cytoplasm; therefore, In Chapter 5, we continue our general study of the cell by con-
it seems reasonable that the nuclear envelope has pores. Finding sidering some of the functions common to all cells. For example, all
relationships between structure and function will give you a deeper cells exchange substances across the plasma membrane, and they
understanding of the cell and boost your memory capabilities. also carry out enzymatic metabolic reactions, which either release
Also, realizing that the organelles work together is helpful. If you or require energy.
wanted to describe the involvement of cell parts to make a protein,
you would start with the nucleus because chromosomes contain DNA,
which specifies the order of amino acids in a particular protein. From
PUT THE PIECES TOGETHER
there, you would mention the ribosomes at the rough endoplasmic 1. Use the structure of the prokaryotic cell to support the endo-
reticulum (RER), transport vesicles, the Golgi apparatus, and a possi- symbiotic theory.
ble final destination for the protein. Analogies can help. For example, 2. Explain how the structure of the endoplasmic reticulum suits
the endomembrane system can be compared to a post office: Proteins its function.
(the letters) are deposited into the RER (the local post office), which 3. Microtubules are a part of the cytoskeleton and are found in
sends them to a Golgi (the regional sorting center) from which they cilia and flagella. What function of the cytoskeleton is consis-
are sent to their correct destinations. The pulse-labeling technique, tent with the presence of microtubules in these structures?
described in Section 4B, provides evidence to support this analogy.
Enhanc
Enhance
a c your understanding of evolution and
diversit
diversity through media and applications!
Media
Chapter 13 Darwin and Evolution
Animations Evolution of Homologous Genes
Genetic Drift
Video Finches—Natural Selection
Virtual Lab Dinosaur Dig
Chapter 14 Speciation and Evolution
Videos Finches—Adaptive Radiation
Cichlid Specialization
Chapter 15 The Evolutionary History of Life on Earth
Animations Geological History of Earth
Breakup of Pangaea
Three Domains
Phylogenetic Tree
Video Prehistoric Hurricanes
Virtual Lab Classifying Using Biotechnology
Chapter 16 Evolution of Microbial Life
Animations How the HIV Infection Cycle Works
Prion Diseases
Binary Fission
Bacterial Spore Formation
Bacterial Transformation
Bacterial Conjugation: Transfer of a Plasmid
Videos Virus Lytic Cycle
Virus Crisis
How Viruses Attack
Cranberries vs. Bacteria
Chapter 17 Evolution of Protists
Animations Endosymbiosis
Malaria: Life Cycle of Plasmodium
Video Plankton Diversity
Chapter 18 Evolution of Plants and Fungi
Videos Carnivorous Plants
Spore Dispersal
Christmas Tree Threat
Chapter 19 Evolution of Animals
Animation Early Vertebrates
Videos Portuguese Man-of-War
Clam Locomotion
Sea Urchin Reproduction
Frog Reproduction
Walking Catfish
Trials of a Tadpole
Leaf-Tailed Gecko
Bird Radar
Mom Grizzly Teaches Her Cubs
Virtual Labs Earthworm Dissection
Classifying Arthropods
Virtual Frog Dissection
Chapter 20 Evolution of Humans
Video Tool Use by Chimps
246
247
248
Butterfly-
pollinated
flower
petal resembles
a female bumblebee
Bumblebee-pollinated flower,
Ophrys elegans
long thin beak
hummingbirds
hover
the long, thin butterfly proboscis to reach the nectar. Hum-
mingbirds fl ap their wings rapidly—called hovering—in
order to remain in one spot while they feed during the day
from odorless, red flowers that curve backward. A humming-
bird’s long, thin beak can access the nectar through a slender
floral tube.
Moth-pollinated flowers are white, pale yellow, or pink—
colors that are visible at night, when moths are active. The flow- floral tube with
ers give off a strong, sweet perfume that attracts moths, which curved back margins
hover as they extend a long, thin proboscis to gather the nectar
at the base of a floral tube. The Madagascar star orchid (Angrae- Hummingbird-pollinated flower
cum sesquipedale) has a very long floral tube that holds its nec-
tar much like a long, thin goblet would hold a drink. When
Darwin fi rst saw a picture of this orchid, he exclaimed, “What
insect could suck it?” Later, he said in his book on orchids, “In
Madagascar there must be moths with proboscises capable of
an extension to ten and eleven inches [25.4 cm–27.7 cm]!” Many
were skeptical, but Darwin was vindicated when in 1903, the
zoologists Lionel Walter Rothschild and Karl Jordan discovered proboscis
a large hawk moth living in Madagascar that has a proboscis
25–30 cm in length. As the hawk moth approaches the flower, it
unrolls its proboscis and inserts it into the floral tube in order
to feed.
We begin our study of evolution in this chapter by exam-
ining the work of Charles Darwin, who provided evidence
that evolution consists of descent from a common ancestor hawk moth hovers
and adaptation to the environment. Further, Darwin offered
a mechanism for evolution he called natural selection. He
called it natural selection because the environment, in a Hawk moth
sense, chooses which members of a population reproduce, pollinated flower
and in that way, adaptation to the environment is eventually
achieved.
249
Great
Britain
Europe
North
America
ATLANTIC
OCEAN
PACIFIC Africa
OCEAN
Galápagos INDIAN
Islands South OCEAN
America
Australia
FIGURE 13.1C Middle: Charles Darwin and the route of the HMS Beagle. Circles pinpoint highlights of Darwin’s trip.
probe for insects. Why were these animals found only in the often contain fossils, are uplifted from below sea level to
Galápagos Islands? Had they evolved there? form land. Lyell’s book went on to support a uniformitarian-
When Darwin explored the region that is now Argentina, ism hypothesis, which states that geologic changes occur at
he saw raised beaches for great distances along the coast. He a uniform rate. This idea of slow geologic change is still
thought it would have taken a long time for such massive accepted today, although modern geologists have concluded
movements of the Earth’s crust to occur. While Darwin was that rates of change have not always been uniform. Darwin
making geologic observations, he also collected fossils that was convinced that the Earth’s massive geologic changes are
showed today’s plants and animals resemble, but are not the result of slow processes and that, therefore, in contrast to
exactly like, their forebears. Darwin had brought Charles thought at that time, the Earth was old enough to have
Lyell’s Principles of Geology on the Beagle voyage. This book allowed evolution to occur.
said that weathering causes erosion and that, thereafter, dirt
and rock debris are washed into the rivers and transported to 13.1 Check Your Progress Look again at Figure 1.1B, a diagram
that illustrates the scientific method. Which part of the diagram
oceans. When these loose sediments are deposited, layers of
applies to Darwin’s approach so far?
soil called strata (sing., stratum) result. The strata, which
b.
FIGURE 13.5A a. Paleontologists carefully remove and study fossils. b. The deeper
the stratum, the older the fossils found there.
a.
the fossil remains. Many more prebird fossils have been discov-
tail head ered recently in China. These fossils are progressively younger
wing than Archaeopteryx: The skeletal remains of Sinornis suggest it
had wings that could fold against its body like those of modern
birds, and its grasping feet had an opposable toe, but it still had
a tail. Another fossil, Confuciusornis, had the first toothless
feet
beak. A third fossil, called Iberomesornis, had a breastbone to
which powerful flight muscles could attach. Such fossils show
how the bird of today evolved.
Scientists had always thought whales had terrestrial ancestors.
wing Now, fossils have been discovered that support this hypothesis (see
Fig. 14.1A). Ambulocetus natans (meaning “the walking whale that
swims”) was the size of a large sea lion, with broad, webbed feet
on its forelimbs and hindlimbs that enabled it to both walk and
Archaeopteryx fossil swim. It also had tiny hoofs on its toes and the primitive skull and
teeth of early whales. Figure 13.6B is an artist’s re-creation, based
on fossil remains of Ambulocetus, which lived in freshwater
streams. An older genus, Pakicetus, was primarily terrestrial, and
reptile characteristics
yet also had the dentition of an early toothed whale. A younger
bird characteristics
genus, Rodhocetus, had reduced hindlimbs that would have
been no help for either walking or swimming, but may have
been used for stabilization during mating.
feathers
The origin of mammals is also well documented. The
synapsids are proto-mammals whose descendants diversi-
fied into different types of premammals. Slowly, mammal-like
fossils acquired skeletal features that adapted them to live more
efficiently on land. For example, the legs projected down-
ward rather than to the side as in reptiles. The earliest true
mammals were shrew-sized creatures that have been
unearthed in fossil beds about 200 million years old.
teeth Section 13.7 discusses the evidence for common descent
based on comparative anatomy.
tail with vertebrae
13.6 Check Your Progress Suppose fossils of hummingbirds indi-
claws cated they had shorter, thicker beaks than at present. What would you
expect to find about the flowers they pollinated?
13.9 Check Your Progress Explain the observation that all organ-
FIGURE 13.9 Biochemical differences indicate degrees of
isms use DNA as their genetic material.
relatedness among organisms.
FIGURE 13.10 The HapMap project compares DNA sequences among African, Asian, and European populations to discover unique base
sequence differences.
sperm
0.04 DD 0.16 Dd
homozygous recessive (dd). From these genotype frequencies,
you can calculate the allele frequencies, in the population:
0.80
d
genotypes DD Dd dd
frequency of genotypes 0.04 0.32 0.64 0.16 Dd 0.64 dd
in the population Offspring
The frequency of the gametes (sperm and egg) produced by this 2pq = frequency of Dd genotype (dark-colored) = 2(0.20)(0.80) = 0.32
population will necessarily be the same as the allele frequencies.
q2 = frequency of dd genotype (light-colored) = (0.80)2 = 0.64
Assuming random mating (all possible gametes have an equal
chance to combine with any other), we can use these gamete 1.00
frequencies to calculate the ratio of genotypes in the next genera- FIGURE 13.11 Calculating gene pool frequencies for F1 and F2.
tion by using a Punnett square (Fig. 13.11).
There is an important difference between a Punnett square succeeding generation of a sexually reproducing population as
that represents a cross between individuals and the one shown long as five conditions are met:
in Figure 13.11. In Figure 13.11, we are using the gamete frequen-
cies in the population to determine the genotype frequencies in 1. No mutations: Allele changes do not occur, or changes in one
the next generation. As you can see, the results show that the direction are balanced by changes in the opposite direction.
genotype frequencies (and therefore the allele frequencies) in 2. No gene flow: Migration of alleles into or out of the popula-
the next generation are the same as they were in the previous tion does not occur.
generation. In other words, the homozygous dominant moths 3. Random mating: Individuals pair by chance, not according
are still 0.04; the heterozygous moths are still 0.32; and the to their genotypes or phenotypes.
homozygous recessive moths are still 0.64 of the population. 4. No genetic drift: The population is very large, and changes
This remarkable finding tells us that sexual reproduction alone in allele frequencies due to chance alone are insignificant.
cannot bring about a change in genotype and allele frequencies. 5. No natural selection: No selective agent favors one geno-
Also, the dominant allele need not increase from one generation type over another.
to the next. Dominance does not cause an allele to become a In real life, these conditions are rarely met, if ever, and allele
common allele. frequencies in the gene pool of a population do change from one
The potential constancy, or equilibrium state, of gene pool generation to the next. For example, when the trees darken due
frequencies was independently recognized in 1908 by G. H. Hardy, to industrial pollution, gene pool frequencies change because
an English mathematician, and W. Weinberg, a German physi- predatory birds (the selective agent) can find and therefore eat
cian. They used the binomial equation (p2 + 2pq + q2 = 1) to light-colored moths. The dark melanic color becomes common
calculate the genotype and allele frequencies of a population, as among moth populations. This is called industrial melanism.
illustrated in Figure 13.11. From their findings, they formulated
the Hardy-Weinberg principle, which states that an equilibrium 13.11 Check Your Progress How do you know when microevolu-
of allele frequencies in a gene pool will remain in effect in each tion has occurred?
gene flow
self-
pollination
Pisum arvense
stamen
stigma
FIGURE 13.13B Occasional
FIGURE 13.13A The anatomy cross-pollination between a
of the garden pea (Pisum sativum) population of Pisum arvense and a
ensures self-pollination and population of Pisum sativum is an
Pisum sativum nonrandom mating. example of gene flow. Pisum sativum
F O R M YO U R O P I N I O N
1. Humans and perhaps apes and mon-
keys like sweet foods. How does
this benefit plants containing sweet
proteins?
2. Are humans influencing the evolution
of plants when they propagate them?
When they genetically modify them
and then propagate them?
3. In what way is artificial selection
A Curculigo plant The fruits develop at base of leaves. harmful to the plants and animals
FIGURE 13B.2 The sweet protein curculin is present in the fruit of a Curculigo plant. selected to reproduce?
FIGURE 13.14A Chance events can cause allele frequency 13.14 Check Your Progress Could genetic drift have set back the
coevolution of flowers and their pollinators? Explain.
changes and genetic drift.
FIGURE 13.15A Phenotype ranges before and after three types of selection. Blue represents favored phenotype(s).
Amount of Color
predation
above waterfall
High
predation
0 4 8 12
below waterfall Months
much less or much greater than usual). When a baby is small, In disruptive selection, two or more extreme phenotypes
its systems may not be fully functional, and when a baby is are favored over any intermediate phenotype (Fig. 13.15Ac,
large, it may have experienced a difficult delivery. Stabilizing right). For example, British land snails (Cepaea nemoralis) have
selection reduces the variability in birth weight in human popu- a wide habitat range that includes low-vegetation areas (grass
lations (Fig. 13.15B). fields and hedgerows) and forests. In forested areas, thrushes
Directional selection occurs when an extreme phenotype is feed mainly on light-banded snails, and the snails with dark
favored, and the distribution curve shifts in that direction. Such shells become more prevalent. In low-vegetation areas, thrushes
a shift can occur when a population is adapting to a changing feed mainly on snails with dark shells, and light-banded snails
environment (Fig. 13.15Ab). become more prevalent. Therefore, these two distinctly different
Two investigators, John Endler and David Reznick, both at phenotypes are found in the population (Fig. 13.15D).
the University of California, conducted a study of guppies, which Stabilizing selection, discussed in Section 13.16, maintains
are known for their bright colors and reproductive potential. the heterozygote, especially if it has an advantage over the
These investigators noted that on the island of Trinidad, when homozygote, as seen in sickle-cell disease.
male guppies are subjected to high predation by other fish, they
tend to be drab in color and to mature early and at a smaller size. 13.15 Check Your Progress If the flowers of a species are pres-
ently only one color and the pollinator prefers this color, is stabilizing
The drab color and small size are most likely protective against
selection occurring? Explain.
being found and eaten. On the other hand, when male guppies
are exposed to minimal or no predation, they tend to be colorful,
to mature later, and to attain a larger size.
Endler and Reznick performed many FIGURE 13.15D Disruptive
experiments, and one set is of particular selection in snails.
interest. They took a supply of guppies
from a high-predation area (below a Forested Low-lying
waterfall) and placed them in a low- areas vegetation
predation area (above a waterfall)
(Fig. 13.15Ca). The waterfall pre-
vented the predator fish (pike) from
entering the low-predation area.
They monitored the guppy popula-
tion for 12 months, and during that
year, the guppy population above the
waterfall underwent directional selection
(Fig. 13.15Cb). The male members of the
population became colorful and large in
size. The members of the guppy population
below the waterfall (the control popula-
tion) remained drab and small.
Darwin took a trip around the world as the natural- Hardy-Weinberg equilibrium (allele frequencies stay the same) if
ist aboard the HMS Beagle. During his trip, he col- mutation, gene flow, nonrandom mating, genetic drift, and natural
lected fossils and made several observations that made him think selection did not occur. However, these events do occur, and they
evolution occurs. Darwin was aware of artificial selection, and he are the agents of evolutionary change that lead to microevolution,
had read an essay by Malthus suggesting that the members of a recognizable by allele frequency changes. Mutations provide the
population compete with one another for resources. Darwin began raw material for evolution. Genetic drift results in allele frequency
to see that a competitive edge would allow certain members of a changes due to a chance event, as when only a few members of a
population to survive and reproduce more than other members of population are able to reproduce because of a natural disaster or
the population. Assuming that advantageous traits are inheritable, because they have founded a colony. Natural selection is the only
future generations would eventually acquire adaptations to the local agent of evolution that results in adaptation to the environment.
environment. Darwin called this process, by which a population Chapter 14 concerns macroevolution, the manner in which
adapts to its environment, natural selection because nature selects new species arise. The origin of new species is essential to the his-
which members of a population will reproduce to a greater extent. tory of life on Earth, which we consider in Chapter 15.
Natural selection is like artificial selection except the environment
PUT THE PIECES TOGETHER
instead of a breeder selects which plants or animals will reproduce.
Evolution explains the unity and diversity of life. Life is uni- 1. We now know that evolution by natural selection can be
fied because of common descent, and it is diverse because of observed over a short period of time (years, months). Give
adaptations to particular environments. Darwin used the expres- examples.
sion “descent with modification” to explain evolution. Support for 2. Why would you expect evolution to have a genetic basis? Use
common descent includes transitional fossils, anatomic features industrial melanism to support the genetic basis of evolution.
(homologous structures, vestigial structures, and embryologic simi- 3. Why would it be incorrect to say that bacteria became resistant
larities), biogeographic data, and molecular evidence. in order to escape being killed by antibiotics?
In the 1930s, biologists developed a way to apply the
principles of genetics to evolution. Populations would be in a
APPLI C AT I O N S
HOW BIOLOGY IMPACTS OUR LIVES
The Many Uses of Corn, an Allotetraploid 280
HOW SCIENCE PROGRESSES
The Burgess Shale Hosts a Diversity of Life 282
Liger
270
Mules Zorse
271
Acadian flycatcher, Empidonax virescens Willow flycatcher, Empidonax trailli Least flycatcher, Empidonax minimus
FIGURE 14.1B Three species of flycatchers. The call of each bird is given on the photograph.
distinguish species, the most important criterion, according to reproduce with one another; therefore, they are separate spe-
the biological species concept, is reproductive isolation—the cies. They live in different habitats. The Acadian flycatcher
members of a species have a single gene pool. While useful, inhabits deciduous forests and wooded swamps, especially
the biological species concept cannot be applied to asexually beeches; the willow flycatcher inhabits thickets, bushy pas-
reproducing organisms, to organisms known only by the fossil tures, old orchards, and willows; and the least fl ycatcher
record, or to species that interbred when they lived near one inhabits open woods, orchards, and farms. They also have dif-
another. The benefit of the concept is that it can designate spe- ferent calls. Conversely, when anatomic differences are appar-
cies even when trait differences may be difficult to find. The ent, but reproduction is not deterred, only one species is
flycatchers in Figure 14.1B are very similar, but they do not present. Despite noticeable variations, humans from all over
the world can reproduce with one another and
belong to one species. The Massai of East
Africa and the Eskimos of Alaska are kept
apart by geography, but we know that, should
they meet, reproduction between male and
female would be possible (Fig. 14.1C).
The biological species concept gives us a
way to know when speciation has occurred,
without regard to anatomic differences. As soon
as descendants of a group of organisms are able
to reproduce only among themselves, specia-
tion has occurred.
In recent years, the biological species con-
cept has been supplemented by our knowledge
of molecular genetics. DNA base sequence data
and differences in proteins can indicate the
relatedness of groups of organisms but it can’t
indicate when speciation has occurred. Macro-
evolution starts with speciation but anatomic
data such as the differences in the whales in
Figure 14.1A play a prominent role in decipher-
ing the history of life.
Pre
Premating Mating Fertilization
Habitat isolation
Hab Zygote mortality
Species at same locale
Spec Fertilization occurs, but
occupy different habitats.
occu Mechanical isolation zygote does not survive.
species 1 Genitalia between hybrid
species are unsuitable offspring
Tem
Temporal isolation for one another. Hybrid sterility
Spec
Species reproduce at Hybrid survives but is
diff
diffe
different seasons or sterile and cannot
diff
fe
different times of day. reproduce.
Gamete isolation
species 2 Sperm cannot reach
Behavioral isolation
In animal species, or fertilize egg. F2 fitness
courtship behavior differs, Hybrid is fertile, but F2 hybrid
or individuals respond to has reduced fitness.
different songs, calls,
pheromones, or other
signals.
g
g
fro
fro
rog
rog
rel
og
rd
nf
df
pa
ke
llfr
Mating Activity
ee
o
leo
pic
wo
bu
gr
low
March 1 April 1 May 1 June 1 July 1
River female
Lake female
FIGURE 14.3B Sockeye salmon at Pleasure Point Beach, Lake FIGURE 14.3C Sockeye salmon in Cedar River. The river connects
Washington. with Lake Washington.
Some colonized an area of the lake near Pleasure Point Beach effect to adaptive changes can be reproductive isolation.
(Fig. 14.3B). Others migrated into the Cedar River (Fig. 14.3C). Another example is seen among Anolis lizards, which court
Andrew Hendry, a biologist at McGill University, is able to tell females by extending a colorful flap of skin, called a “dew-
Pleasure Point Beach salmon from Cedar River salmon because lap.” The dewlap must be seen in order to attract mates.
they differ in shape and size due to the demands of reproducing Therefore, populations of Anolis in a dim forest tend to evolve
in the river. In the river, males tend to be more slender than light-colored dewlaps that reflect light, while populations in
those along the beach. A slender body is better able to turn side- open habitats evolve dark-colored dewlaps. This change in
ways in a strong current, and the courtship ritual of a sockeye dewlap color causes the populations to be reproductively iso-
salmon requires this maneuver. On the other hand, the females lated, because females distinguish males of their species by
tend to be larger than those along the beach. This larger body the color of the dewlap.
helps them dig slightly deeper nests in the gravel beds on the
Ficedula Flycatchers As populations become reproductively
river bottom. Their deeper nests are not disturbed by river cur-
isolated, postzygotic isolating mechanisms may arise before
rents and will remain warm enough for egg viability.
prezygotic isolating mechanisms. Postzygotic isolating mecha-
Hendry has another way to tell beach salmon from river
nisms can keep species separate but they represent a large
salmon. Ear stones called otoliths reflect variations in water tem-
investment of energy to no avail. For example, the production of
perature while a fish embryo is developing. Water temperatures at
a hybrid requires an investment of energy that does not result in
Pleasure Point Beach are relatively constant compared to Cedar
the passage of genes to future generations. Therefore, natural
River temperatures. By checking otoliths in adults, Hendry found
selection would favor the evolution of prezygotic isolating mech-
that a third of the sockeye
anisms over postzygotic isolating mechanisms. The term rein-
males at Pleasure Point
forcement is given to the process of natural selection favoring
Beach had grown up in the
variations that lead to prezygotic reproductive isolation. An
river. Yet the distinction
example of reinforcement has been seen in Ficedula flycatchers
between male and female
of the Czech Republic and Slovakia. When the pied and collared
shape and size according to
flycatchers occur in close proximity, the pied flycatchers have
the two locations remains.
evolved a different coat color from that of the collared flycatch-
Therefore, these males
ers. The difference in color helps the two species recognize and
are not successful
mate with their own species.
breeders along the
Adaptation to new environments can result in multiple spe-
beach. In other
cies from a single ancestral species, as discussed in Section 14.4.
words, repro-
ductive isolation
dewlap has occurred. 14.3 Check Your Progress Knowing that the coat colors of lions
This example and tigers is adaptive to their habitats, construct a hypothetical
scenario by which they evolved from an ancestral species.
shows that a side
eu
creepers have a range of beak sizes don
estor
and shapes for feeding on various food
sources, including seeds, fruits, flowers, and
insects (Fig. 14.4).
Adaptive radiation has occurred in both plants and
* Kauai
animals throughout the history of life on Earth when a
akialoa
group of organisms exploits a new environment. For
Nukupuu
* Akialoa
FIGURE 14.4 Adaptive radiation in Hawaiian honeycreepers.
Genu
Genus Loxops s Hem
ignath
us
Great
amakihi
Anianiau
(green * Extinct species or subspecies
(lesser
solitaire)
amakihi)
14.4 Check Your Progress Five species of big cats are clas-
Amakihi
sified in a single genus: Panthera leo (lion), P. tigris (tiger), P. par-
dus (leopard), P. onca (jaguar), and P. uncia (snow leopard). What
evidence would you need to show that this is a case of adaptive
radiation?
FIGURE 14A Among the many of its uses today, corn is a fuel source, a component of many products that
absorb water, a feed for animals and a delicious food for people.
New species
new
Time
Time
no change species
ancestral species
ancestral species
FIGURE 14.6A Gradualistic model of speciation. FIGURE 14.6B Punctuated equilibrium model of speciation.
Opabinia
282
Thaumaptilon
Wiwaxia
Vauxia
Whether slow or fast, how could evolution have produced the or squids, so wouldn’t it seem
myriad of animals in the Burgess Shale and, indeed, in the history likely that all three types of
of life? Or, to ask the question in a genetic context, how can genetic animals evolved “eye” genes
changes bring about such major differences in form? It has been separately? Not so. In 1994,
suggested since the time of Darwin that the answer must involve Walter Gehring and his col-
developmental processes. In 1917, D’Arcy Thompson asked us to leagues at the University of
imagine an ancestor in which all parts are developing at a particu- Basel, Switzerland, discov-
lar rate. A change in regulatory gene expression could stop a devel- ered that Pax6 is required for
opmental process or continue it beyond its normal time. For eye formation in all animals tested
instance, if the growth of limb bones were stopped early, the result (see Fig. 11.4B). Mutations in the Pax6
would be shorter limbs, and if it were extended, the result would gene lead to failure of eye development in
be longer limbs compared to those of an ancestor. Or, if the whole both people and mice, and remarkably, Eye on fruit fly leg.
period of growth were extended, a larger animal would result, the mouse Pax6 gene can cause an eye to
accounting for why some species of horses are so large today. develop on the leg of a fruit fly.
Using new kinds of microscopes and the modern techniques
of cloning and manipulating genes, investigators have indeed dis- Increase in Complexity The developmental regulatory
covered genes whose differential expression can bring about genes called Hox genes have been much studied, and investiga-
changes in body shapes (Fig. 14.7A). More surprisingly, these tors tell us that the number of these genes increased twice during
same regulatory genes occur in all organisms. This finding sug- the evolution of animals. Both expansions are associated with an
gests that these genes must date back to a common increase in complexity, defined by the appearance
ancestor that lived more than 600 MYA (before the Despite millions of years of of different cell types. One expansion occurred
Burgess Shale animals), and that despite millions of divergent evolution, all ani- during the evolution of vertebrates. Invertebrates
years of divergent evolution, all animals share the mals share the same regula- have 13 Hox genes, while vertebrates, including
same regulatory switches for development. Previ- tory genes for development. humans, have four copies of the 13 Hox gene set.
ously in Section 11.4, p. 216, we pointed out how It appears that the set underwent a series of dupli-
the same regulatory gene, Pax6, turns on eye development even cations, and some of the duplicate genes may have taken on new
though the animal kingdom contains many different types of eyes, functions, a process mentioned in How Life Changes, p. 262.
and it was long thought that each type would require its own set Similarly, some other sets of regulatory genes that operate in
of genes. Flies, crabs, and other arthropods have compound eyes development were duplicated when vertebrates evolved. This
composed of hundreds of individual visual units. Humans and all increase in the number of regulatory gene numbers may have
other vertebrates have a camera-type eye with a single lens. So do contributed to the evolution of vertebrates and to their
squids and octopuses. Humans are not closely related to either flies complexity.
The limbs of these terrestrial mammals are shaped for running (or walking). The limbs of birds are shaped for flight.
FIGURE 14.7A Differential expression of the same regulatory genes during development can account for differences in vertebrate limbs.
FIGURE 14.7B
Differential expression of a
Hox gene causes (a) a chick
to have fewer vertebrae
than (b) a snake in a
particular region (colored
pink) of the spine.
Burke, A. C. 2000. Hox genes and
the global patterning of the somitic
mesoderm. In Somitogenesis. C.
Ordahl (ed.) Current Topics in Devel-
opmental Biology, Vol. 47. Academic
Press.
a. b.
2 MYA
4 MYA Equus
Neohipparion
Hipparion
12 MYA
Dinohippus
15 MYA Megahippus
Merychippus
17 MYA
23 MYA
25 MYA
35 MYA Miohippus
40 MYA
Palaeotherium
45 MYA
50 MYA
Hyracotherium
55 MYA
FIGURE 14.8 Simplified family tree of Equus. Every dot represents a genus.
Macroevolution, the study of the origin and history of model? The fossils of the Burgess Shale support the punctuated
the species on Earth, is the subject of this chapter and equilibrium model. How can genetic changes bring about such
the next. The biological species concept states that the members of major changes in form, whether fast or slow? Investigators have
a species have an isolated gene pool and can only reproduce with now discovered ancient regulatory genes (e.g., Hox genes), whose
one another. differential expression can bring about changes in body shapes and
The origin of species is called speciation. Speciation usually organs.
occurs after two populations derived from a larger one are separated Evolution is not directed toward any particular end, and the
geographically. If the members of a salamander population are sud- traits of the species alive today arose through common descent with
denly divided by a barrier, each new population becomes adapted adaptations to a local environment. The subject of Chapter 15 is the
to its particular environment over time. Eventually, the two popula- evolutionary history and classification of living organisms today.
tions may become so genetically different that even if members of
PUT THE PIECES TOGETHER
each population come into contact, they will not be able to produce
fertile offspring. Because gene flow between the two populations is 1. Scientists make observations and then formulate testable
no longer possible, the salamanders are considered separate spe- hypotheses to explain the observations. What testable hypoth-
cies. Aided by geographic separation, multiple species can repeat- eses have biologists made about speciation? (Hint: How might
edly arise from an ancestral species, as when a common ancestor species arise? What might cause them to arise?)
from the mainland led to many species of Hawaiian honeycreepers, 2. Paleontologists suggested the punctuated equilibrium model.
each adapted to its own particular environment. What data did they use?
Does speciation occur gradually, as Darwin supposed, or rap- 3. Is the study of evolution a scientific endeavor? Explain your
idly (in geologic time), as described by the punctuated equilibrium reasoning.
CHAPTER 1 can reduce the influence of pesticide resistant has one shell, which is complete with two electrons;
insects on their fields and farming practices. 3. b. Two. Oxygen has two shells with six valence
Check Your Progress The salmon must be successfully raised in an electrons in the outer shell. Therefore, oxygen
1.1 Observation data; observing that a worker ant environment that mimics nature as closely as requires two more electrons for a completed outer
produces eggs. 1.2 Test group: colonies exposed to possible, with exposure to pressures such as normal shell. 2.4 H+, OH-. 2.5 a. See Figure 2.6, page 32, in
the parasite; control group: colonies not exposed to predation and pathogens, for example, if they are text; b. This is the formula that gives each atom a
the parasite. 1.3 a. Levels of organization from cells to succeed in the wild. For instance, if the salmon completed outer shell. 2.6 One end of the molecule
to organism illustrate order; (b) unicellular organisms have no exposure to pathogens that they might is negative and the other end is positive because
divide, multicellular ones produce sperm and egg encounter in the wild, these salmon are at high risk oxygen attacts electron more than hydrogen does.
(c) metabolizes nutrients. 1.4 When organisms of infection upon introduction without the benefit of 2.7 Yes, because electropositive hydrogens are
reproduce they pass on a copy of their genes. building immune defenses from previous exposure. attracted to either electronegative oxygen or nitrogen.
1.5 Only by responding to stimuli can organisms 2.8 Hydrogen bonding causes water molecules
remain homeostatic. 1.6 Natural ecosystems absorb Testing Yourself to stick together and to other polar molecules.
pollutants. 1.7 The first suggests that humans are 1. a; 2. d; 3. c; 4. Scientific theories arise due to 2.9 The air loses heat as it causes water in the pad
apes; the second means that apes and humans are innumerable observations and experimentation. 5.b; to evaporate. 2.10 Polarity makes the emulsifiers
on their own evolutionary pathway. 1.8 a. Domain 6. c; 7. b; 8. c; 9. c; 10. d; 11. Each type organism hydrophilic. 2.11 The blocks of ice trap heat inside
Eukarya and kingdom Animals b. Fossil and has its own sequence of bases in its genes. 12. e; and prevent it from escaping to the environment.
molecular data 1.9 Better adapted members have the 13. b; 14. d; 15. a; 16. d; 17. d; 18. A college 2.12 a. H+; b. OH-. 2.13 a. Acidic; b. More H+.
opportunity through natural selection to reproduce campus has a location, as does an ecosystem. The 2.14 a. Down; b. Carbonic acid froms and releases H;
more, and in that way a species becomes adapted populations of students, faculty, and administrators therefore the pH decreases.
to its environment. 1.10 The hawk has levels of communicate with each other and the physical
organization; catches food for herself and offspring; environment (the buildings). 19. c; 20. e; 21. d; Form Your Opinion
remains homeostatic because she can respond to 22. d; 23. c; 24. a; 25. c; 26. c; 27. Evolution is Page 28: 1. Though controversy will undoubtedly
stimuli, lives in a semidesert ecosystem; and is related to all the other theories; for example, all arise over moral, monetary and other issues, science
adapted to flying. organisms are cellular because their common must be free to ask and answer questions about the
ancestor was cellular. 28. e; 29. f; 30. c; 31. g world in which we live. However, scientists should
Form Your Opinion conduct research with minimal risks to their own
Page 6: 1. No answer: The United States should not Thinking Scientifically safety and that of others. As technology advances,
continue to export its current farming technology. 1. a. Bacteria don’t die in sunlight when dye is present. the restrictions placed upon research should be
Exporting technology that is known to be detrimental b. Dye is protective against UV radiation. c. Experiment reevaluated to ensure their relevance. With careful
to ground water and top soil will eventually reduce consists of exposing control and test groups to UV and balanced monitoring, scientific endeavors
agricultural yields, resulting in a food shortage. The light. d. Hypothesis is not supported. 2. Plant the same should be supported and encouraged. 2. Safety
United States would be better served to encourage species of tomato plants in three large plots. All plots should be paramount in experimentation. The best
sustainable farming practices like crop rotation to receive the same treatment, except plot 1, your control, way to proceed is to aware of advisements regarding
foster long-term success in agriculture. Yes answer: receives no fertilizer; plot 2 receives the name brand safety and to learn from experience about how to
The United States should continue to export farming fertilizer in the same quantity as plot 3 which receives modify procedures to make them safe. 3. Yes answer:
technology to other countries in order to support the generic brand. Measure the tomatoes from each plot Depending on the benefits that an experiment might
the global food market. As solutions to the long- and calculate the average size to determine which plot have, I would consider being a guinea pig. However,
term problems of these technologies arise, the U.S. results in the largest tomatoes. research using people should be entirely voluntary
should make these available to other countries as and provide an explanation of the potential risks. I
well. 2. Circumstances such as labor costs, profits, Put the Pieces Together would be particularly willing to be an experimental
and marketing challenges might discourage a Page 21: 1. One example, that shows the relationship subject in the field of medical research, especially
farmer from growing organically. These obstacles between two theories, is the link between ecosystems if the results of experimentation had the potential
might be overcome through government subsidies and evolution. Different species of organisms are to prove widely useful. No answer: I would not
for sustainable farming practices, sharing best the result of evolution over time from a common be willing to serve as a guinea pig in experiments
practices through networking with successful organic ancestor. The evolution of new species creates new that may prove harmful to me. Because of known
farmers, and a world-wide movement towards eating populations that interact to form communities, which risks, and the fact that the risks associated with
primarily locally grown and seasonal produce. in turn interact to form ecosystems. Ecosystems experimentation on humans and animals cannot
Page 17: 1. Practical problems in agriculture, also evolve over time as new species evolve and always be predicted, I would be unwilling to subject
medicine, and conservation can be solved with a change the populations and communities within the myself to the potentially hazardous side effects of
knowledge of evolution. Because we know that ecosystem. 2. The evolution of antibiotic resistant being a human subject in experiments.
natural selection drives the evolution of pesticide bacteria results from natural selection. Mutations in Page 38: 1. The changes in lakes due to acid rain kill
resistant insects, changes to farming practices and bacterial DNA created resistant individuals. These fish and other wildlife and sometimes eliminate it
pesticide use are possible. In medicine, awareness bacteria survive to reproduce in the presence of altogether. Trees suffer as a result of acid rain’s effects,
of the evolution of antibiotic resistant bacteria antibiotics, whereas non-resistant individuals die becoming diseased or dying. If this continues, not
allows patients and doctors alike to make changes off. This process creates a population of antibiotic only will the atmosphere’s oxygen be affected, but the
to the way antibiotics are prescribed and used. In resistant bacteria. lumber industry will suffer. The food supply is at risk
conservation efforts, knowledge of evolution can if plants as well as fish are negatively affected by acid
be used to make informed decisions regarding
endangered species; directed evolution can also
CHAPTER 2 rain. 2. Human beings must take some responsibility
for what they do to the planet and make attempts
be used to select for organisms to clean up the Check Your Progress to control their negative impact. Driving less could
environment. 2. By naturally allowing for the 2.1 Carbon, nitrogen, phosphorus, sulfur. 2.2 See help to preserve lakes and the wildlife in them,
growth of nonresistant insect populations, farmers Figure 2.3B, page 29, in text. 2.3 a. One. Hydrogen forests, buildings made of limestone and marble, and
A-1
alike. No answer: We should not grow more corn Check Your Progress Testing Yourself
for ethanol conversion and decrease its use as food 15.1 When tracing the evolution of life, you need to 1. b; 2. d; 3. a; 4. c; 5. e; 6. e; 7. a; 8. d; 9. d; 10. a;
for humans and animals. We should develop better start with the past and move toward the present. The Ferns; b. produce seeds; c. naked seeds; d. needle-like
waste management practices and moderate our dates are in MYA; therefore, the larger numbers in the leaves, Conifers; e. fan-shaped leaves, Gingkos;
consumption of beef while continuing to use corn timescale are more distant from the present than are the f. enclosed seeds; g. one embryonic leaf, Monocots;
for food. If we can grow more corn, this is not an smaller numbers. 15.2 Yes, because dinosaurs evolved h. two embryonic leaves, Eudicots. 11. b; 12. d; 13. e;
either/or situation and we can have enough corn for at the start of the Mesozoic era, when the continents 14. b; 15. e; 16. b; 17. New and different stuctures
ethanol production and food uses. were all still joined. 15.3 Humans didn’t evolve until arise due to DNA differences. 18. b; 19. a, b, c;
Page 283: 1. Since science seeks to answer questions the Holocene epoch, long after the last mass extinction. 20. d, e; 21. b; 22. The three-domain system is based
about the natural world, the scientific community The possibility exists that humans could become extinct on differences/similarities in the sequencing of
should use phenomenon like the Burgess Shale to due to their own activities. 15.4 Only plants that rRNA. This is backed up by differences in structure.
help answer these questions. Since fossil evidence resemble virgina creeper are in the same genus. Domain
can help to fill in blanks in evolutionary history, Eukarya contains the plants, animals, fungi, and Thinking Scientifically
the fossils should be examined and evaluated in protists. 15.5 Animals share the same way of life; they 1. The tree shows that all life forms have a common
an attempt to figure out how they fit into the story are all motile and ingest their food. 15.6 Monkeys and source and how they are related, despite the
of our planet. The scientific community should gorillas share a more recent common ancestor than do occurrence of divergence, which gives rise to
do its best to preserve such phenomenon while reindeer and gorillas. 15.7 epidermal scales 15.8 Yes, different groups of organisms. 2. The specialized
objectively examining the history hidden inside. dinosaurs being vertebrates share the homologous environmental niche of these organisms is the same
Potentially detrimental “shots in the dark” should structures of vertebrates. as it was when they first evolved.
be avoided and in some cases, samples should be
preserved until technology can be developed Form Your Opinion Put the Pieces Together
to meet the demands of dissecting fragile fossil Page 299: 1. In some instances the ability to readily Page 307: 1. Yes answer: Putting the biosphere and
evidence. 2. Students should be presented objective identify any species of organism could be beneficial all of its organisms into a tree of life would put the
information about the unexplained. This can to society. In fields such as farming, education, and vast nature of the world in perspective and would
encourage students to understand the progressive medicine this technology would prove particularly make me want to preserve all species now alive.
nature of science, along with how much we do not useful for identifying organisms that propose a No answer: The overwhelming nature of a complete
know about the world in which we live. Exposure to threat to plants and/or human beings. For example, tree of life might decrease my interest in the
phenomena that is yet to be fully explained can help a farmer, doctor, or student could differentiate preservation of species by sheer over-stimulation.
students understand the scientific process. These poisonous or otherwise harmful insects or plant Seeing that so many species exist, some being very
students may someday be scientists themselves, species that differ only subtly in appearance from similar to one another might detract from the value
developing techniques or technology that will help non-harmful species. This could assist in medical of each species individually. 2. Yes answer: I do look
us learn more about these phenomena. diagnostics for rashes, insect or snake bites if forward this because determining the ancestry of
some portion of the organism were available for humans all the way to the first living source is an
Testing Yourself identification. Because of the need to scan either exciting concept that seems to fit naturally into the
1. c; 2. c; 3. b; 4. f; 5. a; 6. e; 7. h; 8. e; 9. c; 10. e; the organism or a portion of it, the ability to identify human desire to understand who we are and where
11. Genetic differences; 12. a. species 1; b. geographic a species may not always be possible, however. we came from. No answer: I do not look forward
barrier; c. genetic changes; d. species 2; e. genetic 2. The conversion of the CBOL methods to a criminal to tracing the ancestry of humans to the very first
changes; f. species 3. 13. d; 14. b; 15. Allopatric identification process would most likely require living source and do not trust the data that traces
Testing Yourself
other plants would burn. 2. Plant native grasses, trees,
and shrubs along ditches or other unusable parts
CHAPTER 23
1. c; 2. c; 3. b; 4. c; 5. a; 6. Terminal bud is at the of land to prevent runoff and erosion, and promote Check Your Progress
shoot tip and produces cells that add to the length of insects that are natural pollinators and predators. Use 23.1 Different second messengers can bring about
the stem and become new leaves and new axillary pesticides sparingly and with deliberate attempts to various cellular activities in the same or different types
buds. Axillary bud activity produces new branches avoid spraying during risky times for local insects, of plant cells. 23.2 Yes, because if the hormone
(and flowers). 7. A cereal grain is a dry fruit that fish, or other wildlife. Plant various crops including remained, it would continue to trigger a response long
contains a seed. 8. b; 9. c; 10. c; 11. b; 12. a; 13. d; those useful for ground cover. 3. I am certainly willing after the response was no longer needed. For example,
14. e; 15. c; 16. d; 17. a; 18. d; 19. b; 20. b; 21. b; to eat new grains in order for farmers to increase it would bring about bending after the light source
22. They all consists of parenchyma cells that fill the biodiversity. Maintaining biodiversity is good for the was no longer present. 23.3 The plant in (a) most
interior of an organ. 23. e; 24. c; 25. b; 26. b; 27. e; planet as a whole. If simple changes like eating habits likely doesn’t produce gibberellin, and the receptor in
28. c; 29. b; 30. Apical meristem (shoot tip and root can assist in keeping our planet a suitable home for (b) is more likely defective. 23.4 You could apply
tip meristem) is responsible for primary growth and many organisms, I’d be happy to do so. cytokinins to increase the number of cells and
the result is increase in the length of stem and root. Page 457: 1. Plants are a source of food, make gibberellins to increase the size of the cells. 23.5 a.
Vascular cambium is responsible for secondary oxygen, take in carbon dioxide, provide shelter, Abscisic acid maintains dormancy and closes stomata;
growth and the result is increase in girth. Girth serve as building materials, clean up nitrate b. Gibberellins have the opposite effect. 23.6 To be an
increases because secondary xylem builds up as pollution, grow materials used for clothing like effective hormone, a molecule needs only to combine
annual rings (wood). 31. c; 32. a. Broader expanse to cotton, produce or aid in production of medicines, with its receptor. 23.7 It is adaptive for roots to grow
collect sunlight; b. Prevents loss of water; c. Collects and serve to preserve soil. 2. Find some way to toward water because it enhances their ability to
sunlight; d. Allows gas exchange; e. Allows carbon prevent pets and other animals from having access extract water and dissolve minerals from the soil for
Check Your Progress government should not be involved in every choice Form Your Opinion
31.1 Carnivores are predators that attack and kill other we make during our lifetimes. Page 651: 1. (1) A hydra lives in an aquatic
animals. Digestion (both mechanical and chemical) of Page 644: 1. Our society admires thinness and environment, which ensures a moist exchange
meat is more easily accomplished than the digestion of to avoid being seen as overweight, a person surface. (2) Earthworms keep themselves moist
G-1
GLOSSARY G-2
G-3 GLOSSARY
GLOSSARY G-4
G-5 GLOSSARY
GLOSSARY G-6
G-7 GLOSSARY
GLOSSARY G-8
G-9 GLOSSARY
GLOSSARY G-10
G-11 GLOSSARY
GLOSSARY G-12
G-13 GLOSSARY
GLOSSARY G-14
G-15 GLOSSARY
GLOSSARY G-16
G-17 GLOSSARY
GLOSSARY G-18
G-19 GLOSSARY
GLOSSARY G-20
G-21 GLOSSARY
GLOSSARY G-22
G-23 GLOSSARY
GLOSSARY G-24
G-25 GLOSSARY
GLOSSARY G-26
G-27 GLOSSARY
GLOSSARY G-28
G-29 GLOSSARY
Note: Page references followed by f in pollinators, 248–249, 249f fungal diseases, 374, 374f Alternative splicing, 218, 218f
and t refer to figures and tables in small populations, 736 and introduction of alien species, 806 Altman, Sidney, 318
respectively. social behaviors and, 750–751, 750f, plant viruses and, 312 Altruism, 750–751
751f sustainable, 816, 816f Alu element, 240
Adaptive radiation, 278, 278f, 350 water use and, 815 Alveolates, 336f
A Addiction
definition of, 542
wildlife, value of, 802–803
AGS. See Adrenogenital syndrome
Alveoli, 654, 654f, 659f
Alzheimer disease, 163, 220, 530, 538
A (amino acid) site, 200–201, 202, 202f drug abuse and, 543, 544 AID. See Artificial insemination by donor Amantia, 374
Abdomen smoking and, 655 AIDS. See HIV/AIDS Ambulocetus natans, 256, 256f, 272f
of crayfish, 394f treatment of, 544 Air pollution, and acid rain, 38, 38f American Heart Association, 643
of insect, 396 Addison disease, 691, 691f Airsickness, 563 Ames test, 206
Abduction, 577 Adduction, 577 Alaska, 787, 787f Amino acids
Abiotic factors, in population control, Adductor longus, 579f Albumin, 53, 600 common, 53f
733, 733f Adenine (A), 56, 57f, 57t, 192, 192f, 198f Alcohol metabolism of, 666, 666f
ABO blood type, 182, 604 Adenoids, 632 and acidosis, 675 peptide formation, 52, 52f
Abortions, spontaneous, 164 Adenosine diphosphate (ADP), 58, 58f, and cancer, 154, 206 in proteins, 52–53, 53f
Abscisic acid (ABA), 456 88, 88f, 582–583, 583f and cirrhosis, 638 R groups of, 53, 53f
Abscission, 441 Adenosine monophosphate (AMP), 88 as diuretic, 674 sources of, 309
Absorption, in digestive process, 629, Adenosine triphosphate. See ATP effects of, 543, 597, 686 structure of, 53, 53f
629f Adenoviruses, 310, 310f fermentation and, 133, 133f, 134 synthesis of, 135
Absorption spectrums, of photosynthetic ADH. See Antidiuretic hormone functional groups and, 45 Amino groups, 45f
pigments, 109, 109f Adhesion, of water, 33 Alcoholic drinks, production of, 134 Amish, genetic abnormalities in, 263,
ABT-594, 406 Adhesion, of water in plant vessels, 33f, Aldosterone, 674, 675f, 691 263f
Abyssal plain, 792f, 793 454 Algae, 343–346 Ammonia
Acacia trees, 789 Adhesion junctions, in animal cells, blooms, 324, 775 and origin of organic molecules,
Acceptor end, 200 508–509 blue-green, 324 (See also 317–318, 317f
Accessory fruits, 497 Adipose tissue, 510f, 511 Cyanobacteria) as polar molecule, 32f
Acetabularia, 335f ADP (adenosine diphosphate), 58, 58f, brown, 337f, 344, 344f as waste product, 666, 666f
Acetic acid bacteria, 134 88, 88f, 582–583, 583f classification of, 335 Ammonium, in nitrogen cycle, 775, 775f
Acetylcholine (ACh), 530–531, 540, 543, Adrenal cortex, 683f, 684, 685, 690–691, diatoms, 343, 343f Amniocentesis, 179, 181
583 690f dinoflagellates, 343, 343f Amnion, 402f, 715, 715f, 716f, 717
Acetylcholinesterase (AchE), 531 Adrenal glands, 683f, 690–691, 690f green, 335f, 345–346, 345f, 346f, 793 Amniotic cavity, 716, 716f
Acetyl CoA, 128, 129, 129f, 135, 135f, 136 Adrenaline. See Epinephrine evolution of, 379 Amniotic egg, 399, 399f, 402, 402f, 403,
ACh. See Acetylcholine Adrenal medulla, 683f, 690, 690f mutualism in, 371 404
Acheulian tools, 420 Adrenocorticotropic hormone (ACTH), as plant ancestor, 352–353 Amniotic fluid, 25, 715, 717, 720, 720f
Achondroplasia, 180 683f, 686, 687f, 690, 690f, 691, life cycle of, 160, 160f, 344, 345, 345f Amniotic membrane, 720, 720f
Acid(s), 36, 36f 691f as photoautotroph, 770 Amoeba proteus, 335f, 339
Acid-base balance, regulation of, 658, 675 Adrenogenital syndrome (AGS), 691 red, 104–105, 335f, 336f, 337f, 344, Amoebic dysentery, 333, 339
Acid deposition, 38, 38f, 775, 807 Adult stem cells, 142, 145, 149, 151 344f, 793 Amoeboids, 335, 336f, 337f, 339, 339f
Acidosis, 37, 658, 675 Advertising, and classical conditioning, Algin (alginate), 344 Amoebozoans, 336f
Acid rain. See Acid deposition 747 Alien species, and extinctions, 799, 805f, AMP (adenosine monophosphate), 88
Acoelomates, 387 Aerobic exercise, 136, 136f 806, 806f Amphetamines
Acon terminal, 530, 530f Aerobic process, 125 Alkalosis, 37, 658, 675 amphetamine psychosis, 543
Acromegaly, 688, 688f Aerobic prokaryotes, 323 Allantois, 402f, 715, 715f, 716f, 717 effects of, 597
ACTH. See Adrenocorticotropic hormone AF-2, 206 Allele(s), 144, 173, 173f urine tests for, 673
Actin Afferent arteriole, 671, 671f, 672f dominant, 173 Amphibians
and cell motion, 78 Afferent neurons. See Sensory (afferent) harmful, heterozygote advantage and, characteristics, 384, 401, 401f
in cell structure, 68f, 78, 78f neurons 266 circulatory system, 593, 593f
in muscle contraction, 89, 89f, 512, African apes, 413f multiple, 182 evolution of, 399f
582–583, 582f, 583f African ethnicity, skeletal remains and, in natural selection, 177 excretory system, 666
Actin filaments, 78, 78f 569 recessive, 173, 261 kidneys, 670
Action potential African sleeping sickness, 333, 338 x-linked, 185, 185f reproduction, 698
mechanism of, 528–529, 528f, 529f Afterbirth, 720, 720f Allele frequency respiration, 651, 656
speed of, 529, 529f Agar, 344 calculation of, 260, 260f Amplexus, 698, 698f
Active immunity, 615 Agaricus bisporus, 374 changes in, causes, 259–261 Ampulla, 562, 562f
Active site, of enzyme, 90, 90f Age Hardy-Weinberg principle, 260 Amygdala, 538, 538f
Active transport, 97t age structure diagrams, 731, 731f Allen’s rule, 424 Amyloid plaques, 538
Acyclovir, 708 and risk of Down syndrome child, 163 Allergic reactions, 622, 622f, 622t Anabaena, 324f, 360
ADA (adenosine deaminase), 237 Age-associated nerve deafness, 561 Alligators, 290, 402, 402f, 698–699 Anabolic steroids, 50
Adam’s apple, 654, 703 Agent Orange, 309 Allopatric speciation, 276–277 Anabolism, 135
Adaptation Agglutination, of blood, 604, 604f Alloploidy, 279, 279f Anaerobes
in angiosperms, 433, 433f Aggregate fruits, 497 All-or-none manner, of action potential, facultative, 323
as characteristic of life, 16 Agnatha. See Jawless fishes 528, 528f obligate, 323
early theories of, 251 Agranular leukocytes, 601 Allosteric site, 92, 92f Anaerobic process, 125
innate behaviors and, 744–745 Agriculture Alpha 1 antitrypsin, 406 Anal canal, 636
to land environment, in plants, contour farming, 816 Alpha helix, 54, 54f Analogous structures, 303
354–355 development of, 422, 422f Alpine tundra, 783, 783f, 786 Analogy, 303
learned behaviors and, 746–747 diversity in, 816, 816f Alternation of generations, 160, 354, 354f Anandamide, 544
I-1
I-2 INDEX
B 620
BCR. See B cell receptors
228f, 229f
Biomass, ecological pyramid and, 773,
Blastula, 390, 710, 710f
Blaylock, Mike, 457
Baboons, 415, 752–753 Beagle (ship), 250, 251f 773f Blending model of inheritance, 170
Baby boom, 731 Beaks, of birds, 254, 254f, 403, 403f Biomolecules Blepharisma, 335f
Bacillus anthracis, 297, 328 Bean seed, structure of, 495f, 498f carbon in, 44 Blindness, causes of, 555
Bacillus (bacilli), 320f, 321 Bears, grizzly (Ursus arctos horribilis), classes of, 44 Blind spot, 554
Bacteria 810, 810f functional groups and, 45, 45f Blood. See also Clotting; Red blood cells;
antibiotic resistance, 252, 322, 326 Bee(s) Bioreactors, 233 White blood cells
as biological weapons, 328 beeswax, 50 Bioremediation, 309, 309f, 759f calcium level, regulation of, 689
chemoautotrophic, 319, 793 biological clock in, 753 Biorhythms, 693 color of, 588–589
chromosomes, 320 honeybees (Apis mellifera), 753, 753f, Biosphere composition of, 511, 511f, 600, 600f
commercial uses, 323 803 aquatic ecosystems, 791–794 as connective tissue, 511
cyanobacteria (See Cyanobacteria) mimicry in, 763, 763f definition of, 12 functions of, 511, 600
denitrifying, 775 as pollinators, 249, 249f ecological study of, 728 glucose levels, regulation of, 518, 638
and disease, 309, 326, 326t Beer, brewing of, 134, 373, 435 human impact on, 12 and homeostasis, 518, 519
as domain, 14, 14f, 297 Beetles, 493, 701, 701f, 759, 759f, 763, as level of organization, 8f pH level, 37, 675
endospore formation, 321, 321f 763f terrestrial ecosystems, 785–790, 785t regulation of, 518, 600, 658, 675
evolution of, 337f Behavior Biosphere 2, 780–781 phosphate levels, regulation of, 689
functions performed by, 14 courtship, 753 Biosynthesis stage, of viral life cycle, 311, salt-water composition of, 520
generation time, 321 definition of, 744–748 311f, 314, 314f, 315, 315f water content of, 33
genetically modified, 233, 233f genetically-based, 744–745, 744f, 745f Biotechnology, definition of, 229, 230 Blood clots, immune system and, 614
gene transfer in, 322, 322f genetic vs. environmental factors in, Biotechnology products, 233. See also Blood doping, 648
habitats of, 14 183 Genetically modified organisms Blood flukes, 388, 388f
in human intestines, 766 imprinting, 746 Biotherapy, 603 Blood pressure, 599, 599f
human uses of, 309 learning, in animals, 746–747, 746f, Biotic community, in chemical cycling, arterioles and, 595, 595f
kingdoms of, 14 747f 774, 774f autonomic system and, 526
INDEX I-3
I-4 INDEX
INDEX I-5
I-6 INDEX
INDEX I-7
I-8 INDEX
INDEX I-9
F 491f, 494f
Filarial worm, 393
Flukes, 388, 388f
Fluoride treatments, for teeth, 633
Fossil record
continental drift and, 294–295
Facial bones, 573, 573f Filter feeders, 385, 628, 628f Flycatchers, speciation in, 273, 273f, 277 data, 303
Facilitate diffusion, 96, 96f, 97t Fimbriae, 704, 704f Focusing, of eye, 555, 555f evolution, evidence of, 255
Facilitation model of succession, 767 Finches Folacin, dietary requirements, 642t geological timescale and, 292, 293t
Facultative anaerobes, 323 on Galápagos islands, 254, 254f, 733, Folic acid, dietary requirements, 642t hominids, 418, 419f
FAD (flavin adenine dinucleotide), 124, 733f, 760, 760f Foliose lichens, 371, 371f phylogenetic research and, 303, 303f
129, 129f, 130, 130f, 132 speciation in, 278 Follicle, in oocyte production, 705, 705f, primates, 415, 415f
FADH2, 125, 125f, 129, 129f, 130, 130f, Fingerprints, 515 706, 706f Founder effect, 263, 263f, 423
131, 131f, 132, 132f Fire ants, 2–3, 2f, 3f Follicle-stimulating hormone (FSH), 683f, Fovea centralis, 554, 554f, 557
Fall equinox, 782f First law of thermodynamics, 87, 771 686, 687f, 703, 706, 706f Fowl pest disease, 328
Fallopian tubes. See Oviducts First messenger, 684, 684f Follicular phase, of ovarian cycle, 706, 706f Fox, Sidney, 319
Familial hypercholesterolemia, 182, 237 First polar body, 705, 705f Fontanels, 572, 718 Foxes
Family, as taxon, 14, 14t, 296, 298f Fischer lovebirds (Agapornis fischeri), Food. See also Diet digestion in, 631, 631f
FAP. See Fixed action patterns 744, 744f allergic reaction to, 622 domesticated, 252, 252f
Farming. See Agriculture Fish angiosperms and, 368, 368f Frameshift mutations, 204
Farsightedness (hyperopia), 556, 556f bony, 399f, 400–401, 400f, 668, 668f caloric value of, 86 Francisella tularensis, 328
Fast-twitch muscle fibers, 584, 584f cartilaginous, 399f, 400, 400f, 668, carcinogens in, 206 Franklin, Rosalind, 194, 194f
Fats 668f corn, development of, 280, 280f, 368 Free-diving, by humans, 648, 649
in body characteristics of, 384 fermentation and, 134, 134f, 373, 374 Free radicals
burning, with exercise, 136, 136f circulatory system, 593, 593f foraging for, 748, 748f detoxification of, 154
formation of, 135 in diet, 597, 640, 643 fungi, 373, 374 effect of, 206, 597
in diet, 597, 636, 640, 640t evolution of, 292, 293t, 399f genetically engineered, 233, 234, 235, Freshwater bony fishes, 668, 668f
digestion of, 634, 634f excretory system, 666, 666f 235f Fresh water ecosystems, 791, 791f, 804,
as energy source, 135, 136 group living in, 750, 763 macromolecules in, 46, 46f 805
structure of, 49, 49f jawless, 399f, 400, 400f polyploid plants, 279, 279f Fresh water supplies, 815, 815f
Fatty acids, 49, 49f kidneys of, 670 Food and Drug Administration, U.S., 233, Frogs, 401, 401f
FBN1 gene, 184 lateral line system in, 400, 564, 564f 235, 406 development, 380, 380f, 710, 711, 711f,
Feathers, in birds, 403 lobe-finned, 401 Food chains, 773 712, 712f, 714, 714f
Feather star, 397f in ocean ecosystem, 793, 793f Food vacuole, 339, 339f, 340, 340f egg, structure of, 713, 713f
Feces, composition of, 636 overfishing, 809, 809f Food webs reproduction, 698, 698f
Feeding strategies, of animals, 628, 628f and pet trade, 809 alien species and, 806 reproductive barriers in, 274
Female(s), human ray-finned, 400–401 detrital, 772f, 773 Fronds, 360, 360f
external genitals, 704, 704f reproduction, 698, 700 and energy flow in ecosystems, Frontal bone, 572, 572f, 573f
and iron in diet, 641 respiration in, 651, 651f, 652, 652f 772–773, 772f Frontal lobe, 535, 535f, 538, 539f
and osteoporosis, 578 speciation in, 276–277, 277f grazing, 772–773, 772f Fruit, 366f–367f, 367
reproductive system, 704–706, 704f, symbiosis in, 766, 766f Foot, of mollusc, 391, 391f anatomy of, 497, 497f
705f Fish hatcheries, 812, 812f Foot-and-mouth disease, 328 development of, 490, 494f, 497
and sexual selection, 749 Fishing, 738, 809, 809f Foraging, 748 in diet, 642
Female gametophyte, 363, 363f, Fish oil supplements, 643 Foramen magnum, 572 evolution of, 365
366f–367f, 490, 490f, 494, 495 Fitzroy, Robert, 250 Foraminiferans, 336f, 337f, 339, 339f types of, 497, 497f
Femur, 572f, 574, 574f Fixed action patterns (FAP), 746, 746f Forebrain, 525, 533, 533f Fruit flies. See Drosophilia melanogaster
Fer-de-lance pit viper, 406 Flagellates, 336f Foreign antigens, 615 Fruiting body, 342
Fermentation Flagellin, 320 Forelimbs Fruticose lichens, 371, 371f
ATP yield from, 125, 133 Flagellum (flagella), 67, 67f, 79, 79f, 80t, mobile, in primates, 412–413 FSH. See Follicle-stimulating hormone
foods produced by, 134, 134f, 373, 374 320, 320f of vertebrates, as homologous Functional genomics, 240–241
in metabolism, 125, 133, 133f in Chlamydomonas, 345, 345f structures, 303 Functional groups, 45, 45f
and muscle contraction, 584 in sponge collar cells, 385, 385f Forensics, 568–569 Fungus (fungi), 370–374
Ferns, 360–361, 360f tinsel, 338 Foreskin, 702, 702f animals vs., 370
in Carboniferous period, 364, 364f Flame cells, 387, 387f, 667, 667f Forest(s) characteristics of, 370, 372
characteristics of, 353 Flamingo, 403f acid rain and, 38, 38f classification of, 297
diversity of, 360, 360f Flat-head Lake, Montana, 801, 801f boreal (northern), 786 club, 371f, 373
economic value, 360–361 Flatulence, 637 and carbon cycle, 776 and disease, 374, 374f
evolution of, 492, 492f Flatworms, 389, 389f in Carboniferous period, 364, 364f economic importance, 374
life cycle, 354–355, 361, 361f characteristics of, 384 climate amelioration by, 804 evolution of, 336f, 337f, 371f
uses of, 360 chemoreceptors in, 552 coniferous, 783, 783f, 785t, 786, 786f as kingdom, 15, 15f
Fertile Crescent, 422 circulatory system, lack of, 590, 590f early, 443, 443f life cycle, 160, 370, 372, 372f
Fertility rate, U.S., 738 evolution of, 382f economic exploitation of, 803 main phyla, 372–373
Fertilization free-living, 387 energy flow in, 772–773, 772f mutualism in, 370, 370f, 446, 446f
chromosome number and, 155 hydrostatic skeleton, 570 montane coniferous, 783, 783f, 786 mycorrhizal, 371, 371f, 464, 464f, 766
double, in angiosperms, 366f–367f, parasitic, 388, 388f temperate deciduous, 783, 783f, 785f, parasitism in, 764
367, 494f, 495, 495f reproduction, 700 785t, 788, 788f plants vs., 370
external, 700, 700f respiration in, 651 temperate rain, 785f, 786 sac, 371f, 372–373, 372f
and genetic variation, 156 Flexor carpi group, 579f tropical deciduous, 785f zygospore, 371f, 372
in humans, 704, 705, 716 Flight, in birds, 403, 403f tropical rain, 783, 783f, 785f, 785t,
Fertilizers, and chemical cycling, 774–775 Flooding, natural regulation of, 804 790, 790f, 805, 805f, 810
Fetal development, in humans, 715,
718–719, 718f, 719f, 719t
Florida scrub jay, 751
Flounders, 763, 763f
and global warming, 118, 118f
tropical seasonal, 785f G
Fever, 613 Flower(s) water-holding ability, 804, 804f GABA, 531, 542, 542t, 543
Fever blisters, 314 anatomy of, 490–491, 490f Formed elements, of blood, 600, 600f GABA transaminase, 531
Fiber, in diet, 639 complete and incomplete, 365, 365f, Formosan termite, 806 Galápagos Islands
Fibers, plant, 437 491 fosB gene, in mice, 745, 745f alien species on, 806
Fibrillin, 184 in eudicots, 434, 434f, 491, 491f Fossil(s) birds on, 251, 254, 254f, 278, 733,
Fibrin, 603, 603f evolution of, 352f, 493, 493f Burgess shale, 282–283, 282f–283f 760, 760f
Fibrinolysin, 706 in monocots, 434, 434f, 491, 491f early research on, 250f, 251 lizards on, 251, 276
Fibroblasts, 510, 510f perfect and imperfect, 491 as evidence for evolution, 255–256, 255f resource partitioning on, 760, 760f
I-10 INDEX
INDEX I-11
I-12 INDEX
INDEX I-13
K Language
in animals, 752–753, 753f
elements basic to, 26, 26f
history of, 292, 293t
glycogen storage in, 48
and homeostasis, 518
Kanamycin, 561 human development of, 420, 422 levels of organization, 8f structure of, 638
Kangaroos, 404 La Niña, 794, 794f molecules of (See Organic molecules) Liverworts, 356, 356f
Kaposi sarcoma, 609f Lantern flies, 763, 763f origin of, 24–25, 317–319 Lizards, 402, 402f
Karyotypes, 144, 144f, 163, 164 Lanugo, 719 on other planets, 25 on Galápagos Islands, 251, 276
Katydids, 763 Large intestine, human, 636, 636f, 637 pH range required by, 37 reproduction, 700
Kelp, 106f, 344 Larva water and, 24–25, 25f, 33–35 speciation in, 276, 277, 277f
Keratin, 515 fire ant, 3, 3f Life cycle LM. See Light micrograph
Ketamine (special K), 543 Larynx (voice box), 633, 654, 654f algae, 160, 160f, 344, 345, 345f Loam, 462
Keystone species, 810 LASIK surgery, 556 angiosperms, 365–368, 366f–367f, Lobe(s), of brain, 535, 535f
Kidneys Latent operant conditioning, 747 490–491 Lobe-finned fishes, 401
and acid-base balance, 675 Lateral line system, 400, 564, 564f bacteriophages, 311, 311f Lobsters, 564, 564f, 589, 589f
artificial, 676, 676f Latissimus dorsi, 579f blood flukes, 388, 388f Logistic growth, 732, 732f, 735
evolution of, 668–669 Laughing gulls, 746, 746f, 753 bryophytes, 356, 357, 357f Longhorn beetle, 763, 763f
and homeostasis, 518 Laurasia, 294f ferns, 361, 361f Long-term memory, 538
human Law of independent assortment, fungi, 160, 370, 372, 372f Long-term potentiation (LTP), 538
function of, 670, 670f 174–176 gymnosperms, 363, 363f Loop of Henle. See Loop of the nephron
structure of, 670–671, 671f Law of segregation (Mendel), 172–173, humans, 160, 160f Loop of the nephron (loop of Henle), 670,
in urinary system, 670, 670f 172f, 182 mitosis and meiosis in, 160, 160f, 354, 671f, 672f, 674, 674f
urine concentration, 674–675, Laws of probability, Mendel’s laws and, 354f Loose fibrous connective tissue, 510, 510f
674f, 675f 175 mosses, 357, 357f Lophotrochozoa, evolution of, 382, 382f
urine formation, 573f, 672–673 Laws of thermodynamics, 87 pine trees, 363, 363f Lordosis (swayback), 573
structure of, 670–671, 671f LDCs. See Less-developed countries plants, 160, 160f, 354, 354f Lorenzo’s Oil (film), 74
Kidney stones, 641t, 670 LDL (low-density lipoprotein), 597, 643 tapeworms, 388, 388f Low-density lipoprotein (LDL), 51, 597,
Killer whale (orca; Orcinus orca), 272f, Leaf curl fungi, 372 viruses, 311, 314, 314f, 315, 315f 643. See also Cholesterol
405f, 809 Leafhopper, 396f water molds, 342 LTP. See Long-term potentiation
Kilocalories, 86 Leaf (leaves) Life history, 735 Lucy, 410, 418, 418f
Kinesin, 79, 79f cells, 444f Life tables, 730, 730f Lucy’s baby, 418
Kinetic energy, 86, 86f color of, 104–105, 109 Ligaments, 510f, 511, 571, 577, 577f Lumbar vertebrae, 572f, 573
Kinetochores, 144, 147, 147f, 148–149 eudicots, 434, 434f, 438, 438f, 439f, Ligers, 270–271, 270f Lumber, 369
Kingdom, 14–15, 14t, 296, 298f 444, 444f Light Lumen
Kingdoms, 296f monocots, 434, 434f, 444 and photosynthesis, 104–105 of blood vessels, 509
Kinins, 601 sessile, 433 Light micrograph (LM), 66 of ER, 71, 71f, 72, 72f
Kinocilium, 563, 563f structure and function, 432–433, 432f, Light microscopes, 65, 65f, 66, 66f Lungs
Kin selection, 750–751 433f, 444, 444f Light reactions, in photosynthesis, cancer of, 143f, 154, 206, 655, 660,
Klinefelter syndrome, 163, 163t tissues, 438, 438f, 439f 108–113 660f
Klipspringers, 752 of vascular plants, structure of, 355, absorption of solar energy, 109, 109f evolution of, 401
Knop, William, 460–461 355f overview, 108, 108f gas exchange in, 96
Koalas, 404, 404f veins in, 107, 107f, 434, 434f, 438, production of ATP and NADPH, human, 654, 654f
Kovalick, Walter W., 457 439f, 444, 444f, 445f 110–113, 110f–113f diseases of, 660, 660f
Kress, John, 299 and water transport in plants, 455, Lignin, 437 smoking and, 655, 655f
Krill, 395, 628 455f Limb(s). See also Appendages; Joint(s) structure of, 650, 651, 651f
Kudzu, 806, 806f Leaf primordia, 441, 441f development of, 284 Lupus, 621, 621f
Kyphosis (hunchback), 573 Leaf scars, 441, 441f forelimbs Luteal phase, of ovarian cycle, 706, 706f
Learning mobile, in primates, 412–413 Luteinizing hormone (LH), 683f, 686,
in animals, 746–747, 746f, 747f of vertebrates, as homologous 687f, 703, 706, 706f
I-14 INDEX
INDEX I-15
I-16 INDEX
INDEX I-17
I-18 INDEX
INDEX I-19
I-20 INDEX
INDEX I-21
I-22 INDEX
INDEX I-23
I-24 INDEX
INDEX I-25
I-26 INDEX