Reproduction in Algae
Reproduction in Algae
Reproduction in Algae
Structure
4.1 Introduction
Ol?jeclives
4.2 Types of Reproduction '
V e g h t i v c l<cproduction
Asexual Reproduction
Sexual Reproduction
4.3 Reproductio~iand Life Cycle
C'lilar~~ydo~~~onus
~l/o/liri.~
~I/\~u
Lai~iinorrcr
, P rrcrrJ
4.4 Origin and Evolution of Sex
Origin o f Sex
E:volution o f Scx
4.5 Summary
4.6 Terminal Questions
4.7 Answers.
4.1 INTRODUCTION
In unit 3 you have learnt that algae vary in size from small microscopic unicellular
forms like Chlanzydonionas to large macroscopic multicellular forms like Lanzinaria.
The multicellular forms show great diversity in their organisation and include
filamentous. heterotrichous, thalloid and polysiphonoid forms. In this unit we
will discuss the types ofreproduction and life cycle in algae taking suitable
representative examples from various groups. Algae show all the three types of
reproduction vegetative, asexual and sexual. Vegetative method solely depend on
the capacity of bits of algae accidentally broken to produce a new one by simple cell
division. Asexual methods on the other hand involve production of new type of
cells, zoospores.
In sexual reproduction gametes are formed. They fuse in pairs to form zygote.
Zygote may divide and produce a new thallus or it may secrete a thick wall to form
a zygospore.
Yog will see that sexual reproduction in algae has many interesting features which
also throw light on the origin and evolution of sex in plants. This will be discussed in
the last section of this unit.
Objectives
After studying this unit you shoti:d be able to:
Akinete
Filamentous algae (like Ulothrii) may reproduce by producing motile cells called
zoospores (Fig. 4.2). The protoplast of a single cell divides many times by
mitosis to produce several zoospores. Each zoospore has 2-4 flagella with
which it swims for solnetime and then settles by its anterior end. It subsequently
divides into a lower cell which becomes the holdfast and the upper cell which
by further divisions becomes the vegetative filament. Zoospores are produced in
other algae also.
Zoospore
Zoospore
liberation
Cliluniydonronrrs
I Although this is a unicellular motile algae but it produces zoospores. The parent
1
cell divides inside the cell-envelop and each daugllter cell develops two flagella
each. These zoospores look exactly like the parent cell except they are smaller
in size. When the zoospores are fully developed the parent cell wall disolves,
. releasing them free into the surrounding water (Fig. 4.3 ).
Sometimes when there is less water outside, zoospores may lose flagella and
round up. These non-motile spores are called aplanospores which develop into
thick walled hypnospores.
On moist soil when zoospores can not be released due to lack of free water, they
get embedded within a gelatinous material formed from parent cell wall. Such
cells do not have flagella but whenever they become flooded with water they
develop flagella and swim away in the water. These gelatinous masses containing
thousands of non-motile cells are known as palmella stage of Chlamydonlonas.
Separation of
Daughter
cclls
D l v ~ s ~ oof
n
parent cell
Palmella Stage
the absence
(1x1
of water) .
Oedogonium
Zoospore are produced singly in a cell. Each has one nucleus and a crown of flagella
at the apex.
4.2.3 Sexual ~ e ~ r o d u c t i o n
Sexual reproduction in algae like in other organisms involves the fusion of two cells
from opposite sex called gametes, resulting in the formation of a zygote. Some ,
Gametes are always haploid and may or may not be different in morphology. If
both the sex cells look alike, they could be male called plus (+) or female called
minus (-) matingtypes o r strains. Gametes can fuse only when one is plus and
the other is minus.
a) Isogamy: When both the gametes are of the same size and morphology.
b) Anisogamy: The two gametes are distinctly different in size or shape, the
larger of the two is minus (female) type.
c) Oogamy: 'The female gamete, egg or ovum is big in size and has no flagella
hence it is non-motile. Male gametes are flagellated and highly motile. They are
also known as antherozoids, spermatozoids or sperms.
The male gameles are attracted by the female cells because of special hormones
called galnones (a volatile hydrocarbon) produced by them. Fusion of the gametes
leads to the formation of a zygote. If the conditions are unsuitable for growth, the
zygote may develop a thick wall and become a resting zygospore. Gametes being
haploid, are prclduced by mitosis in a haploid thallus. If the thallus is diploid as in
F ~ l c i the
~ s reproductive cells undergo meiosis or reduction division to form haploid
gametes.
Rcproduction in Algae
Isogamy
Oogamy
In haploid tliallus, after the fusion of gametes, the diploid zygote undergoes meiosis
during germination. However, in diploid algae a zygote may divide mitotically and
give rise to a diploid thallus (Fucus). Both haploid and diploid thallus are found in
Ulva. They look very similar in size and shape.
SAQ 4.1
a) Which of the following algae reproduce only by binary fission?
i) Volvox
ii) Chlumydomonas -
iii) Anacystis
i v) Microcystis
Sexual reproduction in this alga shows all the three different types depending on the
species (Fig. 4.6). Isogan-ly is found in C.moewtrsii, C.r-ehlhnrdii, C.,yynogam~aand
C.media
Oogamy
In clonal populations (cells obtai .ed by the repeated divisions of a single parent
cell) fusion may take place betwf an gametes which are homothallic or in self
compatible strains. For examplr , fusion occurs between any two cells of
C.gynogamu and C.niedia.
In C.moewusii and C.l.einha1 h i fusion of gametes can take place only when they
come from two different unrelated (heterothallic, self incompatible) strains.
The process of gametic attraction, fusion and related phenomena have been studied
in some detail in the laboratory. Under proper light condition and carbon dioxide
concentration, production of gametes can be initiated by nitrogen starvation. The
formation of male or female gametes (even in the case of isogamy) is attributed to
the varying concentration of gamones produced by them. The attraction between
gametes was found due to the presence of glycosidic mannose at the tips of the
flagella of one strain which in a complementary way binds with the substance
present in the flagella of the gamete of the opposite strain. Once this sticking of the
flagella of plus and ~ n i n gametes
~ ~ s takes place, flagella twist about each other
bringing the anterior ends of the gametes ciose. This is followed by cellular and
nuclear fusion.
The zygote secretes a thick wall and accumulates large amount of food materials like
starch, lipids and orange- red pigments. It is now known as zygospore which
remains dormant till the environmental conditions are favourable for its germination.
It has been shown that during germination of zygospore meiosis takes place
followed by mitosis resulting in haploid Chla~iiydo~~zoncrs
cells.
Life Cycle
Chlanzj~donzonasis unicellular, haploid and reproduces asexually many times by
forming zoospores. Under ~~nfavourable environmental conditions it produces
gametes which fuse to form diploid zygospores. During germination reduction
division takes place and haploid cells are fornled (Fig. 4.7).
Asexual
1 \ reproduction
Zoospores (n)
'f /'
Meiosis \
Gametes (n)
Fusion
(syngamy)
Fig. 4.7: Life cycle of Clrlrm~~~donronrrs.
It has been found that Ulothrix produces gametes when grown under long day
conditions while short day conditions initiate the formation of zoospores.
Life Cycle
Look at Fig. 4.8 showing the life cycle of Ulothrix.
Which is the diploid stage of the algae?
The thallus of Ulotlirix is haploid and the diploid stage is represented by the
zygote only.
Rcprod~~ction
ill Algae
We would like to draw your attention to the fact that in some species (Uspeciosa,
U.jlcca and in U.blpfexa) the zygote develops into an independent, unicellular
thailus which is diploid in nature. It produces zoospores asexually by meiosis. The
zoospores develop into haploid filaments.
Haplobiontic:
The thallus is haploid and only the zygote is diploid e.g. U.zonalu?
Diplobiontic:
In diplobiontic cycle, the alga consists of a haploid thallus that produces gametes and
a diploid unicellular stalked thallus which produces zoospores after meiotic division.
The two generations - hapIoid and diploid, alternate with each other. (alternation of
generations). Because the two thalli are very different in size and niorphology it is
known as heteromorphic, diplobiontic life cycle.
Isogamy
Alternation of Generations
The type of life cycle of an organism in which reproduction alternates with
each generation between sexual reproduction and asexual reproduction is
called alternation of generations. Tlie two generations are termed as
gametophytic and sporophytic generations. The gametophytic generation is
haploid(n) and the sporophytic generation is diploid (2n).
4.3.3 Ulva
The life cycle of Ulva is shown in Fig. 4.9. Note the thalli of sporophyte and
gametophyt/e. Both are morphologically alike. However, the gametophyte is
haploid (n) whereas the sporophyte is diploid (2n). The haploid gametophyte
produces gametes and the diploid sporophyte produces after meiosis zoospores that
germinate to form haploid gametophytes.
Fig. 4.9: Life cycle o f Ulvn. Note the ison~orphic rlternation o f generations.
The gametophytes of Ulva produce gametes which are isogalnous or anisogamous.
After fusion the zygote is formed which develops into a diploid sporophyte.
4.3.4 Laminaria
Sexual reproduction in Laminaria is oogamous type.
The mature diploid thalli of sporophytes produce sori or unilocular sporangia on the
surface of the lamina. Each sporangium divides by meiosis to give rise to 32
biflagellate zoospores which germinate to form male and female gametophytes (Fig.
4.10). 1
Unilocular
Sporangia
Female gametophyte
Male
gametophyte
Sporophyte
Zygote (2x1)
-
Female gametophyte~
Fig. 4.10: A) Life cycle oflanunurin. B) The sporophyte is mrcroscopic rnd the male and female
gametophytes are microsopic. '
The gametophytes of both sexes are microscopic with a few branches and their
fertility is controlled by environmental conditions.
Any cell of the female gametophyte can develop into an oogonium, the contents of
which form a single egg. The egg protrudes out when mature but remains attached to
the mouth of the empty oogonial cell.
Antheridia are produced singly as lateral outgrowths of the male gametophyte. Only
one sperm is produced from each antheridium, which is pear shaped and has two
flagella of unequal length.
After fertilization the zygote immediately divides mitotically without any resting
period and de;elops into a sporophyte ( Fig, 4.10).
Life Cycle
In Laminaria there is a distinct alteration of haploid gametophyte and a dominant
diploid sporophyte. II
1
Reduction division takes place in the sporangia of sporophyte before the formation
of zoospores, which germinate to form the male and female gametophytes.
The two dissimilar generations - one simple filamentous gametophyte and the other
highly differentiated, complex multicellular thallus - alternate with each other -
hence the life cycle is termed heteromorphic alternation of generations.
4.3.5 Fucus
Fucus has advanced type of reproductive structures, termed as receptacles, which
are swollen at the tips of branches (Fig. 4.1 I A).
Distributed over the miface of each receptacle are small pores, known as ostioles
which lead into the cavities, called conceptacles (Fig. 4.1 IB). Each conceptacle
may produce only eggs, only sperms or as in some cases both. A thallus may be
unisexual - either having male receptacle or only female ones.
Ostiole
I Conceptacle
At the base, inside the conceptacle is a fertile layer of cells whicl~develops into
oogonia (Fig. 4.12A and 4.14A). Each oogonium has a basal stalk cell and an upper
cell which undergoes reduction division and produces eight haploid eggs (4.12 C
and D). These are liberated in the conceptacle (Fig. 4.12E). Some of the cells inside
the conceptacle produce unbranched multicellular hairs called paraphyses which
emerge out of the ostiole as tufts.
Reproduction in Algac
;.. '.._
..;.. F..._,
B
..
...;:,.*;
..:.-,+
'
Antheridia
I
Sperm
@ C
- Rhizoidal
initial
Nucleus
Eye spot Cortex
1
. Medulla
Sperm
The release of the gametes is connected with the sea tides. At low tide, Fuctis fronds
shrink due to loss of water, and when such fronds are exposed to an on coming tide,
the eggs and sperms are released into the surrounding sea water.
The eggs of Fticus are known to attract sperms (Fig.4.13 A and B) by secreting a
gamone . ltnmediately after fertilizatiori a wall is secreted around the zygote. It has
been shown that unfertilized eggs can develop into gertnlings parthenogenetically if
treated with dilute acetic acid.
The four basic types of life cycles described above are sunimerised in Fig. 4.15
I
Meiosis I
I I
c/
Isomorphic
aitemation 9
(e.g. Ulva)
1
When the dominant pllase is the haploid gametophyte, the life cycle is termed as
haplontic life cycle. In this cycle diploid state or sporophyte is represented by zygote
which produces spores by meiosis that develop into gametophytes.
In diplontic cycle the main or dominant phase is the diploid sporophyte. The zygote
directly germinates into a sporophyte. Later meiosis takes place producing haploid
Algae
gametes that fuse to form the zygote. In the diplontic algae it is to be noted that no
free living haploid thalli are found.
When both the gametophyte and the sporophyte are equally developed and look
n~orpholoigicallysimilar, we have isomorphic alternation of generations. However,
ifga~netophyteis underdeveloped compared to the sporophyte the life cycle-is
known as heteromorphic alternation.
------
SAQ 4.2
a) In the following statements choose the correct alternative word given in the
parentheses.
i) Zygote of Chlanydon~onasundergoes (meiosis/mitosis) during
germination.
iii) In haplontic life cycle, the alga is (haploid/diploid), only the zygote is
(haploidldiploid).
What factors lead to the fusion of cells as such is not clear but fusion brings about
mixing of two different (but related) genomes together, one probably co'knpensating
for the deficiencies of the other. This particular feature is a biological advantage for
Iteprod~~ctiur~
in Algae
the survival of the species. It is no wonder that almost all organisms developed
sexual method of re~roduction.
Fig. 4.16 : Hypothetiral diagram illustra~ngth; origin ofsex in Ufotlrrir (Adapted from Kumar
and Singb. 1990).
Even in the case of prokaryotic cyanobacteria, and also in other bacteria different
mechanisms were discovered (para-sexual mechanisms) whose essential feature is
exchange or mixing of genes or complete genomes between a donor and a
recipient.
In all eukaryotic algae as in all plants and animals, fusion of cells is the method
by which sexual reproduction takes place. The question is how this fusion of cells
originated and further how this phenomenon was preserved and refined during
evolution. The study of the sexual processes found in the present day algae provide
some answers to the above questions.
In many algae one can not make out any difference in structure between a
zoospore and a gamete, except for their behaviour - a zoospore directly develops
into a filament whereas a gamete needs fusion with another gamete for further
development. If certain type of zoospores - small microzoospores can behave
like gametes, at times gametes which fail to fuse may behave like zoospores and
develop directly into thallus - a phenomenon called parthenogenesis reported to
be present in diverse organisms. Such observations indicate that gametes are
modified zoospores and gametic fusion originated through accidental fusion of
small and weak zoospores. As such fusions in general help by genetic
mixing, to acquire characters useful for biological survival, the essential features
of sex were retained and improved further during evolution.
4.4.2 Evolution of Sex
Isogamy, fusion of identical gametes seems to be the earliest state of sex. However,
the morphologically similar gametes may be different in origin, arising from two
different gametic mating types, plus and minus strains (heterothallic).
The simplest early state appears to be the fusion (not any more accidental but
regularised) of morphologicaIly similar gametes, perhaps arising from the same
thallus - homothallic isogamy. This is improved further by heterothallic
isogamous fusion, in which though gametes looked morphologically similar but
with genetical and biochemical differences to encourage fusion of opposite
mating types, plus and minus only.
Generally, the eggs are liberated into the surrounding water but there is a
tendency to retain the egg inside the oogonium itself, where fertilization also takes
place. The zygote or oospore may develop further inside the empty oogonium.
It is to be noted that the above account of the origin and evolution of sex is
entirely based on the study of reproductive process of various algae. Biologists
in recent years discovered that in algae, sex has genetic and biochemical
basis. In Chlanydon~onasgametes produce a volatile substance that attracts the
gametes of the opposite sex. The eggs of Fucus, Laminaria, Oedogonium and
many other algae have been shown to produce species-specific chemicals to
attract the spermatozoids. Such chemicals are known by a collective name
'garnones or pheromones' or sex hormones.
SAQ 4.3
a) Indicate which of the following statements are true or false. Write T for
true and F for false in the given boxes.
TERMINAL QUESTIONS
1. List the factors that control the life cycle of an algae.
2. With the help of a labelled diagram describe the three types of gametic
fusion in sexual modes of reproduction.
4.7 ANSWERS
Self-assessment Question
b) i) dimorphic
ii) Ulva, isomorphic,
iii) small pores, conceptacle
iv) biflagellate
/
v) gamone
vi) diplontic
Terminal Questions
Temperqture
Light
Availability of nutrients
Seasons
Wave action
Periodicity of tides
Isogamy
Anisogamy
Oogamy
There is mixing of two different but related genomes, one
compensating for the deficiency of the other. This is pal-titularly
advantageous for the survival of species.
In many algae one can not make out any difference in structure
between a zoospore and a gamete, except for their bellaviour - a
zoospore directly develops into a filament whereas a gamete needs f~rsion
with another gamete for further developnlent. If certain type of
zoospores - small niicrozoospores can behave like gametes, at times
gametes which fail to fuse may behave like zoospores and develop
directly into thallus - a phenomenon called parthenogenesis reported to
be present in diverse organisms. Such observations indicate that gametes
are modified zoospores and gametic fusion originated through accidental
fusion of small and weak zoospores.