Chlorophyceae

Introduction to Chlorophyceae (Green Algae):

Chlorophyceae (chloros, green; phyceae, algal organisation) is commonly known as
green algae’. Fritsch (1935) considered to include the green algae under the class
Chlorophyceae, which have been raised to the rank of division Chlorophyta by Smith
(1938), Tippo (1942) and Bold (1950).

Later Prescott (1969) and Round (1973) considered it to the rank of phylum
Chlorophyta. Papenfuss (1946) included the suffix ‘phyco’ to the divisions of algae and
named chlorophyta as Chlorophycophyta. Later Bold and Wynne (1978) also followed
the same suggestion. Considering more appropriate, the classification of Fritsch (1935)
is followed in this book.

This class consists of 425 genera and about 6,500 species but, later Prescott (1969)
reported that the number of species may be as many as 20,000; with more being
discovered continuously. The name green alga is given because of the presence of
dominant pigments like Chlorophylls a and b over the carotenoids and xanthophylls.
They are all eukaryotes.

Occurrence of Chlorophyceae (Green Algae):

The members of Chlorophyceae generally grow in fresh water (about 90%) and the rest
in saline water, terrestrial habitat etc. The fresh water members such as Volvox,
Oedogonium, Spirogyra etc. grow in ponds, pools and lakes.

Members of conjugales (e.g., Spirogyra, Zygnema etc.) and Oedogoniales (e.g.,

Oedogonium etc.) are strictly fresh water, but the members of Ulvaceae and Siphonales
are predominantly marine. Some members of Volvocales, Chaetophorales and
Cladophorales grow both in fresh and saline water.

Some species of Ulothrix and Vaucheria are subaerial and grow on damp soil. Some
members may be terrestrial and grow as epiphytes on tree trunk, leaves etc. (e.g.,
Trentepohlia); as epizoic i.e., (growing on animal bodies (species of Characium and
Cladophora); as endophytes (e.g., Chlorella), as parasites (e.g., Cephaleuros,
Rhodochytrium and Phyllosiphon) and also cause diseases.
They can also grow in further different habitats like hot springs (Chlorella), snow
(Chlamydomonas yellowstonensis), saline water (C. ehrenbergi) and some remain as
partners in lichen associations.

Important Characteristics of Chlorophyceae (Green Algae):

1. Members of Chlorophyceae grow mostly in fresh water, a few in brackish and saline
water and a few are terrestrial.

2. They show wide range of variations in their thallus structures like unicellular motile
(Chlamydomonas) and non-motile (Chlorella), coenobium (Volvox), palmelloid
(Tetraspora), dendroid (Ecballocystis), filamentous branched (Cladophora) and
unbranched (Spirogyra), heterotrichous (Coleochaete), siphonaceous (Vaucheria) and
parenchymatous (Ulva).

3. Flagella are 1-many, equal in size and inserted either apically or sub-apically. The
flagella show typical 9+2 arrangement when viewed under E.M.

4. The cells are eukaryotic in nature. Usually there is only one nucleus in each cell, but
in Siphonales and Cladophorales many nuclei are present in their coenocytic body.
Normally the number of nucleolus is one per nucleus, but several nucleoli are present in
the members of Conjugales.

5. The cell wall is mainly made up of cellulose, which comprised of hydroxyproline

glycosides or xylans and mannans. In Chara the cell wall is encrusted with calcium and
magnesium carbonate.

6. Inner to the cell wall, semipermeable cell membrane is present which encircles the
protoplast. The cytoplasm contains many small vacuoles which pushes the nucleus with
cytoplasm towards the periphery and called primordial utricle.

7. The flagellate cells have eye-spot or stigma in the anterior portion, which remain
inserted at one side of the chloroplast.

8. The pigments are located in the chloroplast. Chkiroplast generally contains

pyrenoid(s).
9. The main pigment is chlorophylls a and b; those dominate over α-and β-carotenes
and xanthophylls.

10. Phycobilins are absent.

11. The reserve food is starch, composed of amylose and amylopectin.

12. They reproduce by all the three means i.e., vegetative (cell division and
fragmentation), asexual (zoospore, aplanospore, akinete etc.) and sexual (isogamy to
oogamy). The sexual reproduction is absent in some members of Chlorococcales.

13. The zygote or oospore is the only diploid structure in their life cycle.

The range of thallus structure of Chlorophyceae is as follows:

(i) Unicellular motile forms—e.g., Chlamydomonas.

(ii) Unicellular non-motile forms—e.g., Chlorella.

(iii) Colonial motile form—e.g., Volvox, Eudorina, Pandorina.

(iv) Colonial coccoid forms—e.g., Hydrodictyon, Pediastrum.

(v) Palmelloid forms—e.g., Tetraspora.

(vi) Dendroid forms—e.g., Prasinocladus.

(vii) Un-branched filaments—e.g., Ulothrix, Oedogonium.

(viii) Branched filaments—e.g., Cladophora.

(ix) Heterotrichous forms—e.g., Coleochaete.

(x) Siphonaceous forms—e.g., Vaucheria.

(xi) Parenchymatous forms—e.g., Ulva, Codium, Enteromorpha.

Habit and structure: Range of thallus

organization
In Chlorophytes there is no differentiation of root, stem and leaves; hence the plant body
is thalloid in nature. The thallus may be single-celled or many-celled and
shows evolutionary progression from simple to more complex types of construction.
Chlorophytes are a heterogenous group exhibiting a wide range in their thallus structure and
morphology beginning from simple microscopic motile unicellular forms through
multicellular flagellated or non flagellated colonies, palmeloid forms, dendroid forms,
filamentous forms, heterotrichous forms, siphonous forms to well developed
parenchymatous thalli.

I. Unicellular thallus

It is the simplest type of construction within Chlorophyta. Unicellular thallus is of two

types:

(i) Unicellular motile thallus: Vegetative cells have two or four flagella and are motile
(e.g., Chlamydomonas, Tetraselmis, Sphaerella)

(ii) Unicellular non-motile thallus: These cells do not possess flagella, eyespot etc., meant
for locomotion. (e.g., Chlorella, Chlorococcus).

II. Colonial thallus

The colonial habit is achieved by aggregation of the products of cell division within a
mucilaginous mass, by aggregation of motile cells or juxtaposition of cells subsequent to cell
divisions.

(i) Coenobium: A colony with a definite shape, size and arrangement of cells. The number
of cells in a coenobium is determined at the juvenile stage and subsequently there is only
increase in size. Coenobia may be motile with flagellated cells (e.g., Eudorina, Pandorina,
Volvox) or non-motile having cells without flagella (Pediastrum, Hydrodictyon).

(ii) Palmelloid: In contrast to coenobial forms, in a palmelloid colony the number of cells,
their shape and size is not definite. The cells remain irregularly aggregated within a
common mucilaginous matrix, but they are independent and function as individuals. In some
palmelloid forms it is a temporary phase (e.g. Chlamydomonas), whereas in others it is a
permanent feature (e.g., Tetraspora)

(iii) Dendroid: The colony appears like a microscopic tree. The number, shape and size of
cells is indefinite and a mucilaginous thread is present at the base of each cell. Threads of
different cells are united to form a branched structure (e.g., Ecballocystis).

III. Filamentous forms

(i) Filamentous unbranched: Simple unbranched filaments may be free floating

(e.g., Spirogyra) or attached to some substartum (e.g., Ulothrix, Oedogonium)

(ii) Filamentous branched: In Cladophora simple branched filaments remain attached to the
substratum by a basal cell. In such filamemnts branches arise just below the septa between
any two adjacent cells except the basal cell.

(iii) Heterotrichous: The thallus is very much evolved and differentiated into prostrate and
erect systems (e.g., Fritschiella, Coleochaete, Stigeoclonium, Draparnaldiopsis). Both the
prostrate and the erect systems may be well developed or there is progressive elimination
of prostrate or erect systems.
(iv) Siphonaceous: The thallus is made up of branched, aseptate, coenocytic, tubular
filaments as the nuclear divisions are not accompanied by wall formation (e.g. Protosiphon,
Codium)

(v) Parenchymatous: Thallus is formed by the divisions of cells in two or more planes. The
daughter cells do not separate from the parent and give rise to parenchymatous thalli of
various shapes (e.g., Ulva, Enteromorpha). Leaf like or foliaceous thallus as seen in Ulva has
evolved due to transverse as well as longitudinal septation in the filament

Thallus organisation Genera

Motile unicells Chlamydomonas

Motile colourless unicells Polytoma

Nonmotile unicells Chlorella

Encapsulated unicells Phacotus

Motile colony Pandorina

Dendroid colony Tetraselmis

Palmelloid colony Tetraspora

Coccoid (Zoosporic) Chlorococcum

Coccoid (Azoosporic) Chlorella

Simple filament Ulothrix

Heterotrichous filament Stigeoclonium

Discoid (prostrate type) Coleochaete

Crusts or cushions Pseudopringsheimia

Erect type Draparnaldia

Pseudoparenchymatous (uniaxial) Dasycladus

Pseudoparenchymatous multiaxial Codium

Foliose, parenchymatous Ulva

Tubular, parenchymatous Enteromorpha

Pigmentation and Photosynthetic Apparatus:Thylakoids

green algae, red algae, brown algae or blue-green algae-

chlorophylls, carotenoids and biloproteins.

Chlorophylls:

Chlorophyll a, b, c, d and e types.

Chlorophyll a is present in all classes of algae.

Chlorophyll a , chlorophyll b –Chlorophyceae

Carotenoids:

ϒ Carotene and lycopene

chloroplast may be discoid, plate-like, reticulate, cup-shaped, spiral or ribbon

shaped

Cup-Chlamydomonas, girdle shaped e.g., Ulothrix,

reticulate e.g., Cladophora,

stellate e.g., Zygonema,

spiral e.g., Spirogyra,

discoid e.g., Chara or

parietal e.g., Draparnaldiopsis.

Classification of Chlorophyceae (Green Algae):

Class. Chlorophyceae
Orders:
1. Volvocales
2. Chlorococcales
3. Ulotrichales
4. Cladophorales
5. Chaetophorales
6. Oedogoniales
7. Conjugales
8. Siphonales
9. Charales

1. Order. Volvocales:
The order Volvocales includes 60 genera and about 500 species.

Important characteristics:
i. They are commonly found in fresh water. Some are grown in brackish water, marine
water and also on soil.

ii. The plant body is unicellular or multicellular and the multicellular ones are colonial in
habit.

iii. Both unicellular and colonial members are motile, either throughout or some part of
their life cycles.

iv. They reproduce both asexually and sexually. Asexual reproduction takes place by
zoospores, aplanospores, hypnospores etc. and sexual reproduction by iso-, aniso-,
and oogamy.

2. Order. Ulotrichales
Important characteristics:
The order Ulotricales includes 80 genera and about 430 species. Most of them are fresh
water, while a few are marine (e.g., Ulva).

The important characteristics of the order are:

i. They are commonly found in fresh water (e.g., Ulothrix) or on soil, but a few are
marine (e.g., Ulva, Enteromorpha).

ii. Plant body is commonly an unbranched filament; but in Ulvaceae it is

parenchymatous or foliaceous.

iii. Cells are uninucleate and contain chloroplast of different types like C-shaped, pari-
etal, axial etc.
iv. Each chloroplast contains one or more pyrenoids.

v. Asexual reproduction takes place by means of bi- or quadriflagellate zoospores,

aplanospore and akinetes.

vi. Sexual reproduction takes place by gametic union and may be iso-, aniso-, or
oogamous type.

3. Order. Chaetophorales:
Chaetophorales are the plants with hair or setae.

The important characteristics of the order are given below:

i. Members are generally found in fresh water.

ii. Plant body is filamentous and shows prominent heterotrichous habit; however, in
Coleochaete, the prostrate system is well- developed and in Microthamnion the erect
system is well-developed.

iii. Some members have setae (Coleochaete) or hairs (Stigeoclonium) of different types.
The hairs may be in the form of single elongated cell or rows of fine and elongated cells.

iv. The cells contain a parietal chloroplast with many pyrenoids.

v. Erect system bears reproductive structures.

vi. Vegetative reproduction takes place by fragmentation.

vii. Asexual reproduction takes place by bi- or quadriflagellate zoospores, aplanospores

or akinetes.

viii. Sexual reproduction is commonly isogamous (Fritschiella, Stigeoclonium),

anisogamy (Aphanochaete) and oogamy (Coleochaete) are found occasionally.

4. Order. Oedogoniales
Important characteristics:
i. Most of the members grow in fresh water. The order is represented by only three
genera, Oedogomium, Oedocladium and Bulbochaete.
ii. They are filamentous and the filaments may be branched (Oedocladium and Bulbo-
chaete) or unbranched (Oedogomium).

iii. The plant body is differentiated into apical and basal region.

iv. It consists of cylindrical cells and the cells are longer than breadth.

v. Cells are uninucleate and have reticulate chloroplast with pyrenoids.

vi. Cell division is elaborate and a cap is formed at the upper end of the daughter cell.

vii. Asexual reproduction takes place by pyri- form, multinucleate and multiflagellate
zoospores. Flagella are arranged in a ring around the beak-like anterior end.

viii. Sexual reproduction is advanced oogamous type.

ix. Both androspores and antherozoides are multiflagellate.

x. Male gametes are similar to zoospore but smaller in size.

xi. Heterothallic or dioecious species are of two types: macrandrous (where male and
female filaments are of normal size) and nannandrous type (where male is very small
i.e., dwarf male or nannandrium and the female one is of normal size).

5. Order. Siphonales:
Important characteristics:
i. Most of the members of Siphonales are marine. A few members are freshwater. Some
members grow as epiphytes or endophytes.

ii. Thalloid plant body is variously branched, aseptate and multinucleate i.e., coenocytic.

iii. Plant body may be simple vesicular type (Protosiphon) to much branched
filamentous type.

iv. Numerous small and discoid chromato- phores are arranged peripherally inside the
thallus.

v. Nuclei are present towards the inner layer.

vi. The characteristic pigments of this order are siphonin and siphonoxanthin.

vii. Presence of siphon-like central vacuole throughout the plant body, which remains
filled with sap. Cytoplasm is present between the outer wall and vacuole. The order is
named “Siphonales” because of the presence of siphon-like vacuole.

viii. The plant reproduces by all the three means vegetative, asexual and sexual.
Vegetative reproduction takes place by fragmentation, asexual reproduction by
multiflagellate zoospore, aplanospore or hypnospore and sexual reproduction by
oogamy. Rarely they perform iso- and anisogamy.

6. Order. Charales
Important characteristics:
i. Members of this order are distributed throughout the world.

ii. Commonly they are found in fresh water with muddy or sandy bottom and also in
water flowing over limestone.

iii. Plants are macroscopic, much branched, and erect and commonly up to 30 cm in
length.

iv. The plants are differentiated into nodes and internodes. Some of the nodes bear
branches of unlimited growth, those are again divided into nodes and internodes. Each
node of the main axis and branch of unlimited growth bear a number of branches of
limited growth.

v. Cells are very long, uninucleate and contain many discoid chloroplasts.

vi. Most of the species show cortication in the internodes. The cortex consists of
vertically elongated row of cells.

vii. Sexual reproduction is highly advanced, oogamous type.

viii. The male and female reproductive bodies are globule and nucule, respectively.
Globule develops many antherozoids and nucule contains only one egg.
ix. Zygote is produced after sexual reproduction. It shows very much elaborate post-
fertilization changes. During germination, zygote undergoes meiosis and gradually it
forms the plant body.

Methods of reproduction in Chlorophyceae

In Chlorophyceae reproduction takes place by any one or all of the following three
methods, vegetative, asexual and sexual.

1. Vegetative Reproduction: Vegetative reproduction in Chlorophyceae takes place by

the fragmentation, cell division, akinete formation. It involves only vegetative cells and
the parent cell wall is retained after reproduction. Many forms multiply largely by
vegetative means such as:

(i) Fragmentation: In Oedogoniales, vegetative reproduction takes place by the

development of detached fragments of the thalli into a new individual. Fragmentation
involves breaking up of the thallus into two or more pieces or fragments in colonial
(Dictyosphaerium) and filamentous forms (Spirogyra, Ulothrix, Oedogonium etc.). Each
fragment functions as a reproductive unit and grows into a new filament after
undergoing cell division and the subsequent growth of component cells. Fragmentation
may be because of some external mechanical pressure or by dying out of intercalary
asexual or sexual reproductive bodies.

(ii) Cell division: Multiplication by ordinary cell division is a characteristic feature of

some Chlorophyceans. It is also called fission and is a common method of
reproduction in unicellular forms. The cell division is preceded by mitotic division of the
nucleus. The nuclear division is simultaneously followed by the cleavage of the
cytoplasm, which starts and continues by a median constriction of the cell. The
constriction deepens and finally cuts the cytoplasm into two parts.

(iii) Akinetes: An akinete is a large, one-celled, oblong, non-motile, thick walled, spore
like modified resting vegetative structures derived by the thickening of the wall of a
vegetative cell. Akinetes have very thick cell wall and are enriched in food materials.
Akinete can withstand unfavourable conditions and with the onset of favourable
conditions, they can germinate to form a new individual. Akinetes are commonly formed
in Pithophora, Ulothrix, and Chlamydomonas.

2. Asexual reproduction: It takes place by the formation of some specialized

reproductive cells called spores. The spores are produced endogenously in some
specialized bodies called sporangia. Most commonly they are formed by
repeated mitotic division of protoplasts and hence may be called mitospores.
Mitospores are asexual spores and are different from the sexual spores which are
formed meiotically and thus called meiospores. Mitospores may be motile or nonmotile.
Various kinds of spores are zoospores, aplanospores, hypnospores, autospores etc.

(i) Zoospores: Asexual reproduction by zoospores is very widespread in

Chlorophyceae. Zoospores are flagellated asexual spores. They may be biflagellate
(Chlamydomonas), bi-as well as quadri-flagellate (Ulothrix), and multiflagellate with a
ring of flagella (Oedogonium). Zoospores develop either in modified cells
called zoosporangia or in vegetative cells. Zoospores are often formed during night
and are then liberated in the morning from the parent cell through a pore in the
surrounding cell wall or by rupturing of the cell wall (Chapman and Chapman, 1973) and
remain motile from 3 min. to 3 days. The liberated zoospore from zoosporangia contains
a nucleus, an eyespot, one or more chloroplast and 2 to many flagella of equal length
(isokont) which are inserted at the anterior end. On settling over a suitable substratum,
the zoospore germinates into a new thallus similar to that of its parent. Zoospore
formation is the most effective and rapid way of reproduction under favourable
environmental conditions.

(ii) Aplanospores: Non-flagellate, non-motile zoospores which secrete a thin wall of

their own, are called aplanospores. Aplanospores are arrested zoospores which have
skipped the motile phase. Each aplanospore germinates into a new individual
resembling the parent.

(iii) Hypnospores: Under certain conditions the aplanospores secrete thick walls
around them and store abundant food reserves. These thick walled aplanospores are
called the hypnospores. Because of the thick wall, hypnospore may undergo a long
resting period and may survive even in some unfavourable conditions (Ulothrix).

(iv) Autospores: The aplanospore which appears identical to the parent cell are called
autospores (Chlorella).

5) Palmella stage:

Hundreds and thousands of cells are embedded within a common gelatinous matrix. In
Chlamydomonas, the Palmella stage occurs temporarily whereas this is permanent
feature in Palmella.

3. Sexual Reproduction: Sexual reproduction occurs in almost all members of

Chlorophyceae. It involves the fusion of two specialized reproductive cells called
the gametes. The fusion of the cytoplasm of the gametes is known
as plasmogamy and the fusion of the gamete nuclei is termed as karyogamy or
fertilization. Both the gametes fuse and form a zygote which has a diploid nucleus.
In monoecious or homothallic species the two fusing gametes are produced by the
same thallus whereas in dioecious or heterothallic species the fusing gametes are
produced by the two separate thalli. Sexual reproduction always involves three steps:

(i) Gamete production

(ii) Gamete fusion (syngamy and fertilization)

(iii) Zygote germination

Sexual reproduction is of three kinds: isogamy, anisogamy and oogamy.

(i) Isogamy: It is a primitive type of sexual reproduction and mostly occurs in lower
forms. The fusing gametes are morphologically similar in size, form and structure and
are called isogametes. Isogamous sexual reproduction is common in some species
of Chlamydomonas. The isogametes are usually naked, without any cell wall and are
often flagellated. They are produced in ordinary vegetative cells called the gametangia
by the division of the cell protoplast into several daughter protoplasts. Each daughter
protoplasts aquire flagella and is called an isogamete. These isogametes are smaller
than zoospores. The gametes digest the parental cell wall with the help of certain
enzymes (wall autolysins) and are released into the surrounding water.
(ii) Anisogamy: In anisogamy the fusion takes place between dissimilar gametes,
the anisogametes. Anisogamy can be categorized into physiological anisogamy and
morphological anisogamy. In physiological anisogamy (common in Chlamydomonas
monoica, Spirogyra) the fusing gametes are morphologically identical but different in
their behavior e.g. one gamete may be more active than the other. The gametes
of Spirogyra are discernible by their degree of motility. In morphological anisogamy
(common in Chlamydomonas braunii) the fusing gametes differ noticeably in size. In
most of the members of Chlorophyceae the anisogametes are of two different kinds and
can be designated as male and female. Gametes are produced in specialized cells
called gametangia. The male gamete is small and active whereas the female gamete is
large and passive. These gametes fuse externally after their release from the parent cell
wall. Physiological anisogamy is a primitive type of anisogamy whereas morphological
anisogamy is an advanced type of anisogamy.

(iii) Oogamy: Oogamy is the most advanced type of sexual reproduction and occur in
higher forms. The fusing gametes (male and female) in oogamy differ from each other in
every aspect such as size, motility, behavior and structure and are called
heterogametes. Distinct sex organs are formed. Male sex organ is called
an antheridium and the female is termed as oogonium. Inside the antheridia small,
active flagellated gametes are produced which are called as sperms. Sperms are
produced in huge numbers and on maturity they are released in the surrounding water.
The female gametes are produced singly and are large and passive. The female
gamete is retained within the oogonium. The sperm swims in the direction of the egg
and fuses with it. The resulting fusion cell is called the zygote. Oogamous sexual
reproduction is common in Chlamydomonas coccifera, Volvox, Chlorogonium,
Pandorina, Oedocladium etc. The order Oedogoniales is purely oogamous.

Parthenogenesis:

Sometimes the perfect spores are attained without fusion and are called azygospores or
parthenospores. Here the female gamete may give rise to a new plant without fusing
with male gamete. The process is called parthenogenesis. This process has been
recorded from several Chlorophyceae. Sometimes one species of the genus reproduces
parthenogenetically and the other species does not, e.g., Ulva lactuca is
parthenogenetic species and U. lobata is not.

Zygote and its germination:

The zygote is a very resistant perennating body which usually faces the adverse
environmental conditions for its survival. The zygotes of Chlorophyceae may be thin-
walled or thick-walled. The thin-walled zygotes germinate within a day or two after their
formation. The thick-walled zygotes remain dormant for a short or long period and then
germinate.

Iso - and anisogamous species form thick-walled zygotes which possess flagella on
them for a short time and the zygote remains motile during this period. Later on, these
flagella are withdrawn and the zygote secretes a thick wall around it.

All oogamous species possess non-motile zygotes from the very beginning.

The rest period of the zygotes of various Chlorophyceae varies from genus to genus.
The zygotes of Chlamydomonas germinate in ten days; those of Ulothrix in 5 to 9
months, and the zygotes of Oedogonium take the largest period in their germination,
i.e., 12 to 14 months.

At the time of the germination of the zygote, meiosis occurs to bring the haploid phase
again.
Economic Importance of Chlorophyceae (Green Algae):
The green algae are not so economically important except a few members.

Among them Chlorella is very important because of its high protein content, presence of
vitamins and its use in baking industry in the preparation of cake, pastries etc. It is also
used in the preparation of an antibiotic, chlorellin; which is used to control bacteria. It is
also used in different physiological experiments.

Another important member, Chara, is very useful to control malaria for its larvicidal
properties. It is used as fertiliser and in the preparation of polishes.

Monostroma is used to prepare the common food ‘aonori’ in Japan.

In South India, Green Laver, a kind of food, is prepared from Spirogyra and
Oedogonium.

Ulva and Enteromorpha are also eaten’ by some people.

Many members are used as a source of food and O2 for many aquatic animals.