A TEXTBOOK ON ALGAE

Macmillan International College Editions (MICE) will bring to university,

college, professional and school students in developing countries low cost but
authoritative books covering the history and cultures of the Third World, and the
special aspects of its scientific, technical, social and economic development. In
other parts of the world MICE will satisfy the growing demand for information
about the developing countries.
The MICE programme contains many distinguished series in a wide range of
disciplines, some titles being regionally biassed, others being more international.
Library editions will usually be published simultaneously with low cost paperback
editions.
For full details of the MICE list, please contact the publishers.

Macmillan Tropical Biology Series

This is a new series of textbooks for undergraduate biology students in the tropics.
A TEXTBOOK ON
ALGAE

H D Kumar and H N Singh

M
 © Affiliated East-West Press Private Limited
Softcover reprint ofthe hardcover 1st edition 1979 978-0-333-31541-5
All rights reserved. No part of this publication may be reproduced or
transmitted, in any form or by any means, without permission.

First Indian edition published by Affiliated East-West Press Private Limited 1971
Second Indian edition 1976
First Macmillan International College Edition (taken from second Indian edition)
1979

This edition is published by

THE MACMILLAN PRESS LID
London and Basingstoke
Associated companies in Delhi Dublin
Hong Kong Johannesburg Lagos Melbourne
New York Singapore and Tokyo

British Library Cataloguing in Publication Data

Kumar, Har Darshan

A textbook on algae. - 2nd ed. - (Macmillan
tropical biology series).
1. Algae - Tropics
I. Title 11. Singh, Hriday Narain.
589'.39'093 QK566

ISBN 978-0-333-26072-2 ISBN 978-1-349-16144-7 (eBook)

DOI 10.1007/978-1-349-16144-7

This book is not for sale in India, Sri Lanka, Pakistan and Bangladesh.

This book is sold subject to the standard conditions of the Net Book Agreement.

The paperback edition of this book is sold subject to the condition that it shall not,
by way of trade or otherwise, be lent, resold, hired out, or otherwise circulated
without the publisher's prior consent in any form of binding or cover other than
that in which it is published and without a similar -condition inc1uding this
condition being imposed on the subsequent purchaser.
 Contents

Foreword by G. E. Fogg vi
Preface VII

Part I COMPARATIVE ACCOUNT OF ALGAE

Introduction 3
2 Comparative Morphology 11
3 Reproduction and Perennation 26
4 Economic Importance 38

Part II STRUCTURE AND REPRODUCTION OF

SELECTED ALGAE

5 Cyanophyta 51
6 Chlorophyta 69
7 Xanthophyta 123
8 Bacillariophyta 131
9 Phaeophyta 141
10 Rhodophyta 161
11 Field and Laboratory Techniques 178
Suggestions for Further Reading 203
Glossary 206
Index 212
 F oreword

The study of algae is a rapidly expanding branch of botanical science.

Some of these simple plants are being used as experimental material for
advanced research in biochemistry, genetics and molecular biology; others are
of great economic importance both as nuisances and as contributors to the
pr6ductivity of soils and fisheries. Beyond this, most algae are extremely
beautiful objects in themselves and perhaps in the ultimate analysis this is
the main source oftheir attraction for scientists. Nevertheless, however good
the reasons for studying the algae, they may be nullified ifthe student is faced
with inadequate, uninspiring or intimidating textbooks. I am therefore very
glad that two such active and distinguished Indian workers on algae as
Professor H. D. Kumar and Dr. H. N. Singh have found time to write this
introductory text. By providing a clear and sound background ofthe morpho-
logy which every student of the algae requires but at the same time indicat-
ing the other aspects and showing how one may begin practical work on the
algae oneself, I believe that they have produced a book wh ich will contribute
greatly to the future development of the subject.

G. E. Fogg, Se.D., F.R.S.

The Marine Science Laboratories (Menai Bridge)
University College of North Wales, Bangor
 Preface

The need for a suitable textbook on algae for undergraduates has been feit
in many universities in recent years. Our main objective in writing this book
is to fulfil tbis need and to present an updated account of morphology and
reproduction of the more important genera, supplemented with brief descrip-
tions ofrecent contributions on their physiology, biocbemistry, cytology and
genetics.
Any attempt at improving the standard of teaching of algae must begin
in practical cIasses and field studies. One complete chapter is therefore excIu-
sively devoted to field and laboratory techniques. It incIudes some of the
simpler metbods ror culturing, microchemical study, separation of pigments
by chromatographic techniques, preparation of herbarium sheets and per-
manent slides, estimation of growth and primary productivity, and ecologi-
cal study of soil and freshwater forms.
The remaining chapters incorporate brief and simple accounts of some
important and representative types. The discussion also covers topics of
current interest such as the concept of Procaryota and Eucaryota; the ultra-
structure of algal cells and their organelles; the developmental cycle and gene-
tic recombination in certain blue-green algae; the physiology and biochemistry
of nitrogen fixation and heterocyst differentiation; the nature of sexual sub-
stances (gamones) produced by algae; genetics in Chlamydomonas; and the
physiology of growth and differentiation in synchronous cultures of Chlorella.
The genera described in Part 11 are those usually prescribed for under-
graduates. Minute variations in structure, reproduction and life history have
been avoided so as to give proportionately greater coverage to micromor-
phology and modern experimental studies. It is boped that the general com-
parative topics dealt with in Part I will be equally useful for advanced study.
Test questions and selected references for further reading are incorporated
at appropriate places. Exhaustive Iists of references have been avoided but
care has been taken to include the latest references as far as possible, since
the reader can always find most of the earlier works of individual authors
in the citations given.
We are greatly indebted to Professor G. E. Fogg for many valuable
suggestions and for critically reviewing the manuscript. We thank Professors
R. N. Singh, T. V. Desikachary, H. A. Von Stotsch, M. B. E. Godward,
B. M. Johri, Drs. V. P. Singh and F. E. Round for their helpful comments.
We are grateful to Dr. Norma J. Lang ofthe University ofCalifornia, Davis,
for providing the electron micrograph of Anabaenopsis circularis (frontis-
piece).

Varanasi H. D. Kumar
H. N. Singh
 PART I

COMPARATIVE ACCOUNT
OF
ALGAE
 1
Introduction
DEFINITION

The a]gae comprise a ]arge and heterogeneous assemblage of relatively

simple p]ants which have Iitt]e in common except their characteristic oxygen
evo]ving type of photosynthesis. They exhibit great diversity in size and
appearance and are found in freshwater of all kinds, barks, soils, rocks and
marine habitats. Many unice]Ju]ar forms, such as desmids and diatoms, are
extremely beautiful microscopic objects. Some of the ]arger forms, espe-
cia]Jy the red seaweeds, are highly attractive and when mounted and dried
on herbarium sheets appear like paintings.
Algae differ from fungi in possessing chlorophyll and characteristically
holophytic (plant-like), photosynthetic mode of nutrition; parasitic, hetero-
trophie and even holozoic (animal-like) mo des are, nevertheless, known in
certain forms. They differ from bryophytes in lacking a jacket of sterile
cells around their reproductive organs which are mostly unicellular. In
brown algae (Phaeophyceae) muIticellular reproductive organs are produced
but all the cells of such a multicellular (plurilocular) sporangium or gametan-
gium are fertile. The Charophyceae (green algae) constitute another excep-
tion since their reproductive structures are apparently multicellular and the
antheridium is surrounded by an envelope of sterile cells. The female repro-
ductive organ of algae is not organized into an archegonium (egg-producing
muIticellular organ) as is characteristic of bryophytes and pteridophytes.
With the exception of blue-green algae, most others can be sharply dis-
tinguished from bacteria. The main difference between blue-green algae
and bacteria on the one hand, and the remaining algae, fungi and higher
plants on the other, is the procaryotic n~ture of cells in the former group
as contrasted with the eucaryotic nature in the latter. Procaryota lack
organized nuclei, chromatophores and mitochondria. Their cell walls contain
certain characteristic mucopeptides (or muropeptides), and muramic acid,
not found in those of Eucaryota. They are susceptib]e to infection and
lysis by viruses ofsimilar morphology. However, b]ue-green algae resemble
other algae and green plants in their oxygen evolving type of photosynthesis
whereas in photosynthetic bacteria no oxygen is evolved.

VALIDITY OF DIVISION THALLOPHYTA

Algae, fungi and lichens have ]ong been grouped in Thallophyta on account
of the following characters which they share; (I) the zygote does not develop
4 A TEXTBOOK ON ALGAE

into a multicellular embryo while still contained inside the oogonium, and
(2) the reproductive organs, whether spore-producing or gamete-producing,
are not enveloped by a wall of sterile cells.
Smith (1955) believes that the common features of algae and fungi are
due to parallel evolution and not because of phylogenetic relationship.
On this basis, therefore, algae and fungi should not be cIassified togetherin
the Division Thallophyta. This idea is also shared by some mycologists such
as C. J. Alexopoulos, G. C. Ainsworth and A. S. Sussman.
Researches carried out during the past decade (see Klein and Cronquist
1967) have, however, provided some indication of a phylogenetic connection
between algae and fungi. Total absence of photosynthetic apparatus in
fungi rules out the possibility that algae might have evolved from fungi.
That fungi might have evolved from algae by the logs of chlorophyll is
supported by: (1) similarity in certain structural and functional characteristics
between the lower fungi (Phycomycetes, especially Oomycetes and Chytri-
diales) and some algae belonging to the Xanthophyta, Euglenophyta and
Pyrrophyta; (2) similar flagellation, i.e., two unequal flagella-one whiplash
type and the other pinnate type in both Oomycetes and Xanthophyta;
(3) chemieal resemblance of the cell wall polysaccharides of Vaucheriaceae
(Xanthophyta) and Saprolegniaceae (Oomycetes); and (4) the presence in
Euglenophyta and Pyrrophyta, like Chytridiales, of one whiplash flagellum
and a nucleus in which the nuclear membrane persists throughout the
division cycle.
The common features between Oomycetes and Xanthophyta, and among
Chytridiales, Euglenophyta and Pyrrophyta, indicate that fungi might have
evolved from algae and that their evolution was probably diphyletic: the
Xanthophytan line giving rise to Oomycetes and the Euglenophytan and
Pyrrophytan lines leading to Chytridiomycetes. Therefore, the cIassifica-
tion of both algae and fungi in the same group Thallophyta does not seem
unjustified. Nevertheless, a much better and rational way of cIassifying the
lower plants is to group blue-green algae and bacteria as Procaryota, and
other algae, fungi and higher farms in Eucaryota.

CLASSIFICATION

The algae have been cIassified mainly on the basis of the photosynthetic
pigments (Table I), storage products of vegetative cells and the form of
motile reproductive cells. Unlike bryophytes, pteridophytes and higher
plants, the morphology of the mature vegetative stages does not afford a
reliable criterion for distinguishing different cIasses or phyla of algae.
Consequently, several examples are known where morphologically similar
algae come to be placed in different cIasses, because they differ in their
physiological and biochemical characters. Such a cIassification of morpho-
logically similar but physiologicalIy dissimilar forms in different cIasses is
based on the view that there has been a parallelism in their evolution.
The main criteria (see Tables 11 and III) on the basis of which the algae
 Table I. Distribution of Photosynthetic Pigments in Algae

Phylum Chlorophylls Carotenes Xanthophylls Phycobili·

proteins

c:l >.
.s:: c:l
0. c:l
c:l 0 ·5
:z c:: c:l :zc:l c:: ·5
<I)
:z c:l :zc:: ...
c:: c:: .c:: c:l c: .s:: :z c: ~ ->:::
<I) <I)
a b c d e Ot ~ e 0 :z c: c:l
Y c:: c:: c::
:z <11<11>< :zc: :zc:<11 c:<11>< ~c: ~
<I) <11 ~
·2
c:<11 c:<11 <11<11>< :zc: c:<11><
·ü <I) <11 '11
<I) >< >< c:l
0. <11 c:: c:: <11 0 0 <:)
c: >< c:<11 ... <11>< 0>< 0c:l <11>< 0 >< ~
0 ~ >< <11 a" <>
U > "., ";:: >< .s:: "ü ;:....
<11 <I) <I) >. >. 0.
<I)
>.
:zu "5 <11>< Ö<11 0>< .c::c: :su0 -; ~<11 0c:l ~
J,.l.l ...l '" ll..,
....l f1: N ;; Z <: ~ Ci Ci Ci c. ~ ~ Vl <:
- 0'" ~

Cyanophyta + - - - - - + - + + + ? + - - - + + + + +
ChIorophyta +
Chlorophyceae + + + + + + + + - - - - - - - - +a +
Charophyceae + + + + + + + + +
Xanthophyta + + + +? +?-
Chrysophyta + + + + +
Bacillariophyta + + + + + + +
Pyrrophyta + + + + + +
Cryptophyta + + + +? - + + +
Euglenophyta + + + + + + + +
Phaeophyta + + + + + + +
Rhodophyta + + + + + + + - - - - - - - - + +
? - Present in traces only a - Siphonales
 Table 11. Diagnostic Features of Various Aigal Phyla

Phy!um Flagel/a Genetic apparatus Cell wall Food reserves

Cyanophyta absent nucleus absent; naked DNA mucopolymeric proteinaeeous

fibrils (i.e., not associated with cyanophycin and
histones) present in nucleoplasm cyanophyeean
starch

Chlorophyta 1, 2 or 4 (or more), eommonly nuc1eus with nuc1eolus; spindie cellulosic true stareh
equal, anterior, aeronematie, and eentromeric chromosomes
smooth made up of protein and DNA
fibrils associated with his tones

Xanthophyta 2 (rarely more), unequal, anterior, as in Chlorophyta when present either peetinaceous oil and leucosin
1 aeronematie and 1 pantonematie, or eellulosic (Heterosiphonales)
1 with stiff hairs and 1 smooth

Chrysophyta 2, anterior, equal or unequal, both as in Chlorophyta organisms periplastic or with leu eosin and
aeronematic or 1 acronematic and silicified scales, or calcified plates chrysolaminarin
1 pantonematic, 1 with stiff hairs (coeeoliths); non-eellulosic
and 1 smooth

Bacillariophyta generally 1, anterior, pantonematic as in Chlorophyta silicified, in 2 pieces, each with leueosin or
with stiff hairs eomplex perforations chrysolaminarin
Pyrrophyta 2, terminal, inserted in a girdle in nuc1eus with nucleolus, nuclear cells periplastic or with cellulosic true starch or oil
some forms, 1 acronematic and 1 membrane persistent; chromo- wall divided into a number of or both
pantonematic (1 spiral with fine somes mainly made up of DNA plates or 2 halves
hairs) fibrils; no evidence of spindIe

Cryptophyta 2, terminal, sometimes lateral, nucleus with characteristic chro- periplastic as in Pyrrophyta
either both pantonematic or 1 mosomes
pantonematic and 1 acronematic,
when 2, both have stitl' hairs

Euglcnophyta generally I, anterior, pantone- nucleus with one or more endo- periplastic paramylum
matic, sometimes 2, equal or un- somes and many acentromeric
equal, having fine hairs chromosomes; spindIe absent;
nuclear membrane persistent

Phacophyta 2, lateral, unequal, 1 acronematic as in Chlorophyta cellulosic with alginic and fucinic laminarins,
and 1 pantonematic, 1 with stitl' acids as characteristic components leucosin and
hairs and 1 smooth mannitol

Rhodophyta absent as in Chlorophyta non-cellulosic polysaccharides and floridean starch,

polyuronic acids; or with xylan, galactan sulphatc
polyxylan, galactose and xylose; polymers and
occasionally cellulosic f1oridioside
8 A TEXTBOOlC ON ALGAE

Table IU. Dlagnostlc Cbaracterlstlcs oe Varlous Alpl Pbyla

Phylum Photosynthetic apparatus Eye spot

Cyanophyta chromatophores absent, thyla- absent

koids occur singly; pyrenoids
absent

Chlorophyta chromatophores (chloroplasts) part of chloroplast but distinct

with 2-6 thylakoids grouped in a from ftagella
band; pyrenoids with starch
sheath

Xanthophyta chromatophores with 3 thylakoid part of chloroplast but situated

bands; pyrenoids naked adjacent to ftagella

Chrysophyta as in Xanthophyta as in Xanthophyta

Bacillariophyta as in Xanthophyta absent

Pyrrophyta as in Xanthophyta variable

Cryptophyta chromatophores with bands of 2 either as in Chlorophyta, or

thylakoids each; pyrenoids naked found in ternally in spur of
chloroplast, at some distance
from ftagella

Euglenophyta chromatophores with 2-6 thyla- not part of chloroplast but asso-
koids grouped into a band; pyre- ciated with flagella
noid projecting and with an
apical sheath of paramylum

Phaeophyta as in Xanthophyta; pyrenoids as in Xanthophyta

projecting and naked

Rhodophyta chromatophores with single and absent

widely separated thylakoids;
pyrenoids naked

have been classified are: (I) tbe cbemical composition and relative amounts
of the pbotosyntbetic pigments; (2) tbe cbemical nature of tbe food reserves;
(3) the chemical and physical nature of the cell wall (precise details of tbe
physical features can be revealed only under the electron microscope; and
(4) the number, morphology and orient at ion of flagella, especially tbose of
motile reproductive bodies, zoospores or gametes.
These principles are generally accepted as broad guidelines for tbe
primary classification of algae. However, tbe systems of classification
differ widely as regards tbe rank and status assigned to various taxa. Fot
instance, Fritscb (I93S, 1944) divides tbe algae into 11 classes wbereas
others classify them into 7-10 pbyla. Tbe number of orders within a class
also varies considerably in different systems of cla~sification.
 INTRODUCTION 9

The broad system of classification followed in this book is derived from

the ideas of Stanier and van Niel (1962) and of Round (1965) except that
we propose a total of 10 phyla whereas Round recognizes only eight. Our
scheme also indicates the comparative position ofviruses, bacteria and fungi.
The Subkingdom Eucaryota includes bryophytes, pteridophytes, higher
plants ami animals as weIl.

LlVING ORGANISMS

Kingdom 1 ACELLULAR ORGANISMS (Viruses)

Kingdom 2 CELLULAR ORGANISMS
Subkingdom PROCAR YOTA
Phylum SCHIZOMYCOPHYTA (Bacteria)
Phylum CYANOPHYTA (Blue-green algae)
Subkingdom EUCAR YOT A
Phylum CHLOROPHYTA
Phylum XANTHOPHYTA
Phylum CHRYSOPHYTA
Phylum BACILLARIOPHYTA
Phylum PYRROPHYTA
Phylum CRYPTOPHYTA
Phylum EUGLENOPHYTA
Phylum PHAEOPHYTA
Phylum RHODOPHYTA
Phylum MYXOMYCOPHYTA (Myxomycetes, slime-molds)
Phylum EUMYCOPHYTA (Fungi)

Each of the 10 algal phyla contains a single Class (-phyceae) except for
Chlorophyta which includes two (Chlorophyceae and Charophyceae) and
Chrysophyta which is divided into Haptophyceae (with 2 equal ßageIIa)
and Euchrysophyceae.
Some of the algal phyla show certain resemblances and interrelationships
with other algal phy la. Thus, the Rhodophyta and Cyanophyta are charac-
terized by the absence of ßagellated bodies, and presence of biliprotein
pigments and protoplasmic connections between adjacent vegetative cells.
Despite these simiIarities, the red and blue-green algae do not seem phylo-
genetically related in view of the marked differences in their cell structure.
Some relationship is possible between Chlorophyta and Euglenophyta since
both contain chlorophyll-b, ß-carotene and another pigment lutein, and also
because some members of the Chlorophyta (Volvocales) possess an eye spot
of the type found in Euglenophyta. The presence of chlorophyll-c is a
feature which seems to link together the members of Pyrrophyta, Bacillario-
phyta, Cryptophyta and Phaeophyta. Similarly, the presence of the
perfplastfc envelope, the cytopharyngeal apparatus associated with the
gullet, and the contractile vacuoles seem to bring together the members of
Cryptophyta, Pyrrophyta and Euslenophyta. Some phylosenetic connec-
10 A TEXTBOOK ON ALGAB

tion may exist between Pyrrophyta and Cryptophyta since in both the
nuclear membrane persists through the division eycle.
The reeurrence of eertain characters in different algal phyla, however,
does not necessarily mean that these are phylogenetically related. It might
rather be due to parallel evolution of these features in different phyla.
Algae in general seem to have evolved from some photosynthetie procaryo-
tic ancestors closely related to, but not identical with, the present day blue-
green algae. This view is based on the hypo thesis that chloroplasts of algae
and higher plants are celluJar symbionts which existedas photosynthetie
procaryotes in the remote past (see Raven 1970).

TEST QUEST/ONS

1. How would you distinguish cellular from acellular organisms?

2. In spite of the functional equivalence between procaryotic and eucaryotic cells, the
former are considered more primitive than the laUer. Why?
3. How would you justify the inclusion of Cyanophyta and Charophyceae among
algae?
4. What objections would you have if algal flagellates were to be included in the
Phylum Protozoa ?
5. Why is a system of c1assification based on a single feature rather than a combina-
tion of characters considered invalid and unreliable ?
6. Zoologists generally group all unicellular animals in the single Phylum Protozoa
but botanists do not put all unicellular plants in one major taxon. How can you
account for this disparity ?

SELECTED REFERENCES

Fritsch, F. B., Tbe Structure and Reproduction of the Algae, Vol. I, Cambridge Univ.
Press, London (1935).

Pritsch, F. B., The present day c1assification of algae, Bot. Rev., 10,233-77 (1944).

Klein, R. M. and Cronquist, A., A consideration of the evolutionary and taxonomie

significance of some biochemical, micromorphological and physiological charac-
ters in the thallophytes, Q. Rev. Bio/., 42, 105-296 (1967).

Raven, P. H., A multiple origin for plastids and mitochondria, Science, Wash., 169,
641-46 (1970).

Round, F. B., The Biology of the A1gae, Bdward Amold (Publishers), London (1965).
Smith, G. M., Cryptogamic Botany, Vol. I, Algae and Fungi, 2nd edn., McGraw-HiIl
Book Co. Inc., New York (1955).

Stanier, R. Y. and van Niel, C. B., Tbe concept oe a bacterium, Arch. Mikrobio/., 42,
437-66 (1962).
 2
Comparative Morphology
CELL STRUCTURE

In procaryotic algae (Cyanophyta) the photosynthetic, genetic and respira-

tory apparatuses are present in the cell .hut are not separated from each
other by abounding membrane of their own. Consequently, the cells lack

pOlyhedral body
~~~~§

outer layer

A
golgi body
(eye spot)

flagellum

B
Fig.2-1. Two basic types of cellular organization in algae. A, diagrammatic plan of a
procaryotic cell of blue-green algae. B, Heteromastix rotunda showing eucaryotic
organization (after Manton el al., 1965).
organized nuclei. chromatophores and mitochondria (Fig. 2-1 A). In
eucaryotie alsae, on the other hand, the photosynthetie, genetie and
12 A TEXTBOOK ON ALGAE

respiratory apparatuses are membrane-bounded and separated from each

other (Fig. 2-1 B).
CELL WALL

The zoospores, gametes, and mature cells of most of the algal flagellates are
devoid of a typical cell wall and are merely bounded by the cytoplasmic
membrane. In the majority of algae the cells possess a wall of non-Iiving
material outside the cytoplasmic membrane. The algal cell wall is composed
of pure or mixed carbohydrates or may be entirely silicified. Sometimes it
may contain proteins and mayaIso be impregnated with calcium carbonates,
iron, or chitin. It is usually made up of two layers, an outer amorphous
layer which is muciIaginous or pectic and dissolves readiIy in boiling water,
and an inner crystalline layer which is firm and relatively insoluble in
boiling water and contains microfibrils. According to their composition,
three categories of cell wall can be recognized in algae-cellulosic, silicified,
and mucopolymeric. The cellulosic walls, characteristic of green algae, are
fibriIIar and the microfibrils may be composed of only glucose molecules,
or of glucose plus other related sugars. The ac tu al composition of the
microfibrils varies so enormously that no generalization is possible. The
microfibrils are embedded in the amorphous layer of the cell wall. In
Xanthophyta the cell wall is mostly pectic and in Phaeophyta it is
characterized by the presence of additional compounds such as hemicellulose,
alginic acid, fucoidin and fucin. The members of the Rhodophyta and
Chlorophyta have cell walls made of cellulose and pectin. In diatoms
(Bacillariophyta) the cell wall is valved and composed of oc-quartz embedded
in a pectin-like matrix. Their silicified walls lack cellulose, hemicellulose or
polymers of other sugars. The mucopolymeric cell wall is characteristic of
blue-green algae and its principal components are glucosamine, amino acids,
diaminopimelic acid and muramic acid with some amounts ofsugar polymers.
In some algae the abutting walls of adjacent cells have pit connections
and are traversed by fine protoplasmic strands known as plasmodesmata
(cytoplasmic threads) wh ich interconnect the protoplasts.

FLAGELLA

Most algal phyla, except Rhodophyta and Cyanophyta. include species

which are either motile during their vegetative phase or reproduce by means
of motile asexual or sexual cells, the motility being due to flagella they
possess. Morphologically, there are two types of flagella in algae: (I) the
whiplash or acronematic type (Fig. 2-2 A) which are smooth, and (2) the
tinseI or pantonematic type (Fig. 2-2 B) which are covered with small fila-
mentous appendages. A flagellum grows from a basal body, the blepharo-
plast, located in the peripheral region ofthe cytoplasm. Under the electron
microscope it is seen to consist of a peripheral cylinder of 9 doublet fibrils
surrounding a central pair of singlet fibrils (Fig. 2-2 C). Such an arrange-
ment is known as the 9 + 2 arrangement and is characteristic of all flagellated
 COMPARATIVE MORPHOLOGY 13

cells except bacteria. The entire group of fibrils is enclosed in a double sheath
and the two central fibrils have a subsidiary (single) sheath of their own.

v//--mastigoneme

centrat fibr it

A c
Fig.2·2. A, upper.portion of an acronematic flagellum; B, upper portion of a panto-
nematic flagellum (after Manton, 1966); C, cross section of a flagellum as diagramma-
tized from an electron micrograph.
The motile stages of the Chlorophyta possess two or four anteriorly
inserted whiplash flagella of equallength. The Phaeophyta and Xantho-
phyta have one whiplash and one tinsei flagellum of unequallength, the
flagellar attachment being lateral in the brown algae and anterior in the
Xanthophyta. The Euglenophyta generally have two flagella, one of which
emerges from the anterior invagination of the cell and has mastigonemata
whereas the other is non-emergent, stops short within the reservoir and
lacks mastigonemata.

PLASTlOS AND CHROMATOPHORES

The plastids are double membrane organelles of two types, the colourless
or leucoplasts and the coloured or chromatophores. The usual practice is
to designate the chlorophyll-a and chlorophyll-b containing plastids as
chloroplasts; those which lack chlorophyll-b are termed chromatophores.
Typical chromatophores or chloroplasts are found in all algae except the
Cyanophyta in which the photosynthetic pigments are concentrated in the
peripheral zone of the protoplasm.
According to its location in a cell the chromatophore may be parietal or
axile. In the majority of algae only a single chromatophore is present in a
cell but the members of Conjugales, Siphon ales, Charales and Xantho-
phyceae generally contain more than one chromatophore per cell. The
morphology of the chromatophore vati es considerably in different algae
(Fig. 2-3 A-F) but the following main types can be recognized: (1) cup-
shaped, as in Chlamydomonas; (2) girdle-shaped, as in Ulothrix; (3) discoid,
14 A TEXTBOOK ON ALGAE

as in Vaucheria and Chara; (4) reticulate, as in Oedogonium; (5) spiral, as in

Spirogyra; and (6) stellate, as in Zygnema.
In some diatoms and the dinoflagellate Pyrocystis lunula, chloroplasts
can change their shape and location within the cell in response to light.
When exposed to light of either too high or too low intensity, the chloro-
plasts in Sl.4rirella gemma shrink and become rounded off, but when grown
in moderate-intensity light, they expand into normal shape and size. Light-
induced rotation of chloroplast, mediated by phytochrome, has been·
reported in tbe desmid "Mesotaenium.

Fig. 2-3. Shapes of chromatophore. A, Chlamydomonas; B, Ulothrix; C, Tribonema;

D, Oedogonium; E, Spirogyra; F, Zygnema.(After Fritsch, 1935.)

The electron microscopy of the algal chromatophore has revealed that

it is a double membrane structure composed of a photosynthetic lamellar
system traversing the colourless granular matrix, the stroma. The structural
unit of the lamellar system is a closed double membrane disc termed
thylakoid. A thylakoid membrane consists of a centrallipid layer covered
on both sides with pro tein particles, the lipid layer contains the fat-soluble
photosynthetic pigments. The water-soluble pigments (phycoerythrin and
phycocyanin) of Rhodophyta, Cyanophyta and Cryptophyta are located in
characteristic particles, termed phycobilisomes, which are found linearly
arranged along the surface of each thylakoid.
In Rhodophyta the thylakoids are widely separated and occur singly like
cyanophycean thylakoids. In all other phyla they are stacked together into
bands. The Phaeophyta generally contain a band of three thylakoids. In
Chlorophyta and Euglenophyta a band of 4-6 thylakoids predominates.
Except for Chlorophyta and Rhodophyta, the chromatophores of most
algae are enclosed in a characteristic sheath of endoplasmic reticulum lying
extern al to the double-membraned envelope of the plastid.
The chloroplasts of most algae have electron-translucent areas contain-
 COMPARATlVE MORPHOLOGY 15

ing DNA fibrils. In algae with a girdle band, e.g., Ochromonas, the plastid
DNA is localized in a peripheral circular nucleoid inside the chromatophore
but in algae without girdle bands, in general, the DNA areas are randomly
scattered in the chromatophore matrix. Another characteristic feature of
the chloroplast which is of some taxonomie and phylogenetic significance
concerns the site of location and storage of starch grains. In the green algae
starch grains are stored within the chloroplast; in dinoflagellates and red
algae, in the cytoplasm outside the chromatophores; but in the Cryptophyta
they are stored between the wall of the chloroplast and the sac of endo-
plasmic reticulum present around the plastid.
In all algae studied so far, plastid DNA has been found to have a lower
guanine + cytosine content, a lower melting point, and a lower buoyant
density than nuclear DNA. This is in sharp contrast to the situation in
higher plants in which the chloroplast DNA has higher values of these
parameters than nuclear DNA.
Chloroplasts or chromatophores are genetically semi-autonomous systems
containing histone-free DNA (deoxyribonucleic acid), messen ger RNA
(ribonucleic acid), transfer RNA and ribosomes of bacterial type. Besides,
new chloroplasts are always observed to arise frQm preexisting ones. These
features have made it highly probable that chloroplasts are subcellular
symbionts which had their origin in free living photosynthetic procaryotes
(see Raven 1970).

PYRENOIDS

These are organelles composed of densely packed proteinaceous fibrils and

are found lying within or on the surface of the chromatophore. Their
function is to accumulate starch in
green algae, or to concentrate some
similar food reserve in other algae.
A chromatophore may contain a
single pyrenoid as in Chlamydomonas
or many as in Oedogonium. The
pyrenoids have been demonstrated in ;~'.:J-~='1T"Pyrenoid
matrix
almost all algal phyla except Cyano-
phyta. They may arise de novo or
by the division of preexisting pyre-
noids. In Chlorophyta the pyrenoid
is located as a dense mass in the
chloroplast, and a layer of starch
plates encircles its proteinaceous
core; in most other algae the pyre-
noids do not seem to be associated Fig. 2-4. Diagrammatic sketch of electron
micrograph of pyrenoid of Chlamydo-
with reserve food. Except in Phaeo- monas.
phyta, pyrenoids are invariably
traversed by a variable number of chromatophore lamellae (Fig. 2-4).
16 A TEXTBOOK ON ALGAE

In many algae the pyrenoid is a transient structure wh ich may be formed

at certain stages but may disappear at other stages-the presence or absence
possibly depending on photosynthesis and the status of reserve foods in a
cello

NUCLEI
A membrane-bounded genetic apparatus consisting of typical chromosomes
and nuc1eoli is known as a nucleus and in this respect all algae except
Cyanophyta and dinoftagellates are similar. In Cyanophyta the chromatin
material consists of DNA fibrils which are DOt associated with basic proteins
(histones) so that DO organized chromosomes are found. The Duc1ear
membrane is also Iacking in the bIue-green algae. In dinoftagellates there
is a nuclear membrane but there are no organized chromosomes.
The nuc1eus is bound by two unit membranes of which the outer is
continuous with the membran es of the endoplasmic reticulum. The nuc1ear
envelope is traversed by numerous pores. In Xanthophyta, Chrysophyta,
Bacillariophyta and Phaeophyta the nuc1ear envelope is intimately asso-
ciated with a sac of the endoplasmic reticulum found around the chromato-
phores but no such plastid endoplasmic reticulum is met with in Chlorophyta
and Rhodophyta (Leedale, 1970). The interphase nucIei of most eucaryotic
algae contain uncoiled and expanded chromosomes but in Euglenophyta
and Dinophyta the chromosomes, which lack centromeres, remain con-
densed. According to Leedale (1970), persistence of nucIear wall during
mitosis, attachment of chromosomes to nuclear wall, lack of centromeres,
and absence of a compact spindIe are primitive features.
The cells of most algae are uninucIeate but those of Cladophorales,
Siphonales, Heterosiphonales aqd Charales are multinucIeate or coenocytic.
The position, size and shape of the nucIei is quite variable.
The nucIeoli and chromosomes remain suspended in the granular matrix
of the nucIeus. In general, there is a single nucIeolus per nucIeus but in
Conjugales there may be several nucleoli in a nucIeus.
In Euglenophyta the nucIear membrane and nucIeolus persist through-
out the division cycIe whereas in other algae the nucIeoli break up during
division and are reorganized after the completion of the nucIear division by
a nucleolar organizing chromosome. In some species of Spirogyra and
Zygnema the nuc1eolar organizing chromosomes are associated with satellite
chromosomes. The nuclear division may or may not be accompanied by
wall formation. In Conjugales the chromosomes have difl'used centromeres.
Recently, the characteristic structures called synaptonemal complexes
have been observed in the meiotic nuclei of certain red and brown algae.
Under the electron microscope, a synaptonemal complex appears to consist
of two lateral longitudinal rods wh ich give rise transversely to minute fibrils
extending inward. Each such complex is thought to correspond to a
bivalent. The existence of synaptonemal complexes is the most reliabie
evidence for meiosis not only in algae but also in higher plants.
 COMPARATlVE MORPHOLOGY 17

The nuelei of some euglenoids harbour eertain baeteria but the bio-
logical nature of such an association is not known.

GOLGIAPPARATUS
Structurally, the Golgi apparatus is an association of stacks of smooth
lamellae or cisternae in whieh each stack is referred to as a dictyosome and
consists of 2-20 cisternae. In cells having a single dict}losome, tbe Golgi
apparatus is equivalent to one dictyosome. Excepting Cyanopbyta dictyo-
somes occur in most other algae. Tbey may be located near the chromato-
phore, tbe flagellar base or tbe nucleus.
In other organisms tbe Golgi bodies bave been implicated in excretory
functions but their precise role in algae has not been conclusively established.
Dietyosomes are involved in the formation of tbe seales in some green algae
and Chrysopbyceae (including Haptophyceae and CoccoIitbopborids) and
also in tbe formation of tbe cellulosic wall component in Chrysophyceae.

SEPTA
The cross walls between adjacent cells of a filamentous or parenchymatous
tballus are known as septa, and each consists of a central core, tbe middle
lamella and a peripberal portion-the primary wall. Cell division usually
takes place by transverse furrowing of the cytoplasm midway betwecn tbe
two extremities of acelI. Witbin the furrow a transverse wall of pectie subs-
tance known as the middle lamella is first laid down, over which the
subsequent apposition layers (primary wall) consisting chiefty of cellulose,
are d e p o s i t e d . · .
Tbe septa may be complete or incomplete. The former are mainly of
five types: (1) plane, (2) replicate, (3) semireplicate, (4) colligate, and
(5) unduliseptate (Fig. 2-5 A-E). Tbe plane type has a homogeneous con-
tour as in Ulothrix. In some species of Spirogyra the middle lamella shows
circular infoldings: If eaeh of the adjoining eells has two infolds in opposite
directions then the septum is called replicate, and if there is only a single
infold in alternate positions, it is known as semireplicate. The colligate
septum is an H-shaped cross wall as seen in Microspora. In the unduIi-
septate type, the middle lamella is much expanded and has a wavy, undulat-
ing margin, as in Spirogyra undulisepta.
Tbe incomplete septum as found in most Rhodophyta and some Cyano-
phyta has a central opening known as a pit through which the eytoplasm of
adjacent cells remains connected (Fig. 2-5 F-H). In Sphaeroplea (Chloro-
phyta) tbe septa appear to be composed of radiating ingrowtbs of side walls
ß wbich faH to meet in the centre and form a central opening (Fig. 2-5 G).
~ During cell division in Oedogonium, a septum is laid down across the middle
~ of the cell and remains unconneeted with the side walls (i.e., free-floating)
t for some time.
N Except for the plane septa, the other types are thought to assist in thc
18 A TEXTBOOK ON ALGAE

LJ
n A
H HH ~rll
8 C D E

i~l
i
~~~s..~
1(--\I
F
I~
11
H
G

Fig. 2-5. Types of septa. A, plane; B, replieate; C, semireplieate; D. eolligate;

E, unduliseptate; F, Sphaeroplea, thiek septum showing finger-shaped proeesses;
G, Sphaeroplea, septum in surfaee view showing ecntral openi,ng; H, a pitted septum.
(Figs. A-E after Randhawa. 1959; F-H after Fritsch. 1935.)

fragmentation of the filaments. An alteration in the turgor pressure inside

the cellleads to the development of a shearing strain in the cross wall which
finally ruptures, resulting in fragmentation of the filament.

MISCELLANEOUS STRUCTURES AND

CELLULAR ORGANELLES
Stigma
The motHe, 1lagellated cells of algae have a pigmented organelle known as
the stigma or eye spot, which has been accepted as a light-sensitive photo-
receptor organ. However, certain mutant strains of Chlamydomonas lacking
eye spot still respond to light, this has cast some doubt on its photosensitive
nature. The stigma appears as a reddish streak or dot in the anterior,
median or posterior region of the cytoplasm. It is found in the coloured
(photosynthetic) as weIl as colourless motile cells. It is a collection of
separately membraned lipid drop lets arranged randomly in Euglena, or in
rows of two in CMamydomonas, with the space in between containing a
pair of thylakoids (Fig. 2-6 A). In the pigmented species it is often located
near the edge of the chloroplast or the base of the 1lagellum. In the sper-
matozoid of Fucus it lies outside the chromatophore and is unique in having
a pigment layer composed of a large number of chambers. The orange-red
pigments of the eye spot are present in the lipid drop lets.
Comparative studies have demonstrated that in Chlamydomonas and
other green algae thc eye spot is located on the chloroplast whereas in
euglenoids it is found in the cytoplasm outside the chloroplast.
 COMPARATlVE MORPHOLOGY 19

During cell division the eye spot may divide into two, each daughter cell
receiving an eye spot. Or, thc eye spot of the parent cell may go to one of
the daughter cells whereas in the other it arises de novo.

Vacuole
Except in Cyanophyta, the mature cells of algae characteristically possess
one or more vacuoles bound by distinct membranes. Such vacuoles play
an important role in osmotic relations and absorption or regulation of solutes
and water.

eye spot· ~cont,.actlle

Y1r'V \,'\ vacuole
B

A c o
Fig. 2-6. A, Chlamydomonas, stigma with two rows of lipid droplets and a pair of
thylakoids in between; B, Chlamydomonas, upper part of cell with contractile vacuoles
and stigma; C, upper part of a euglenoid cell showing a canal (cytopharynx). reservoir,
vacuolar apparatus and stigma (after Fritsch, 1935); D, mitochondrion with the inner
membrane infolded into cristae wh ich project into the lumen.

In motiIe algae, two types of vacuolar apparatus are recognized: (I) the
simple vacuolcs, known as contractile vacuoles (Fig. 2-6 B), which contract
periodically and expel their contents to the exterior as in Chlorophyta; and
(2) the complex vacuolar apparatus wh ich consists of a cytopharynx (canal),
areservoir, and a group of vacuoles of varying size (Fig. 2-6 C) as is
characteristic of thc Euglenophyta, Dinophyceae and some members of the
Chrysophyta. All these parts work in elose coordination: the smaller
vacuoles feed their contents into larger one wh ich then empties and releases
the contents into the reservoir. Periodical contraction ofthe reservoir then
expels the contents to the outside through cytopharynx.
The vacuolar system is generally considercd as an organ of osmoregula-
tion involved in pumping liquids out of the cytoplasm against an osmotic
gradient. In some holozoic algae the cytopharynx functions as a gullet (for
ingesting food partieIes) besides being an emptying canal for the vacuoles.
Sometimes food reserves such as laminarin and chrysolaminarin are also
stored in the vacuole.

M itochondrion
Except in Cyanophyta, the algal cells invariably contain mitochondria, the
20 A TEXTBOOK ON ALGAE

organelles of respiratory metabolism. Under the electron microscope they

appear to be bound by a two-layered membrane, the inner being infolded
into finger-like processes which project into the cavity (Fig. 2-6 D). The
size and number of mitochondria per cell vary widely and except for a very
few species (e.g., Fig. 2-1 B) most algae have more than one mitochondrion
per cell. Mitochondria are independently muItiplying cytoplasmic organelles
having DNA, RNA and ribosomes of baeterial type. Like chloroplasts,
they are also considered to have originated from free living heterotrophie
proearyotes (see Raven 1970).

Endoplasmic Reticulum
A system of tubules and vesicles traversing the cytoplasm and termed as
the endoplasmic reticulum is known to ex ist in algal eells exeept in the blue-
green algae. These tubules appear to have some particles (believed to be
ribosomes) disposed along their surface. The function of these partides is
to synthesize proteins or enzymes. Two different kinds of ribosomes,
differing in their physical properties and loeation, have been demonstrated
in Chlamydomonas. The 70S ribosomes, charaeteristic of proearyotic cells,
are found in the chloroplast whereas the 80S ribosomes, typical of euearyo-
tic cells, oceur in the cytoplasm. The existence of the procaryotic type of
ribosomes in chloroplasts of Chlamydomonas indicates that the plant chloro-
plast might be of the nature of an intracellular symbiont derived from
ancestral unicelluJar photosynthetic procaryotes (see Raven 1970).

Granules
Diverse kinds of granules such as polybedral bodies, structured granules,
cyanopbycin granules, cx:-granules and ß-granuJes are found within the blue-
green algal cello The exaet.function of most of these granular inclusions
is not clearly und~rstood.

Cuticle
Studies on the organization and chemistry of the cu tide of algae belonging
to Phaeophyta, Rhodophyta and Chlorophyta have shown that it eontains
mueh protein and consists of several alternate microfibrillar and amorphous
layers. In this respeet the algal cuticle differs from the cuticle of higher
plants which lacks the protein components (Haine and Craigie, 1969).

RANGE OF VEGETATIVE STRUCTURE AND HABIT

The organization of thallus in algae is basically of two types, the unicellular

or the muItieellular. The enormous range of forms that algae exhibit
arises from a modification or elaboration of these types. The cells of a
muItieellular thallus may be loosely held together by a muciIaginous matrix
or cemented together through middle lamellae. During its growth eyde an
alga passes through two phases, the vegetative or developmental, and repro-
ductive or multiplicative. In unicellular algae the growth in size constitutes
 COMPARATIVE MORPHOLOOY 21

the vegetative phase whereas increase in cell numbers by cell division

constitutes the reproductive phase. In multicellular thalli, both the vegetative
and the reproductive phases involve cell division and cell growth.

UNICELLULAR HABIT
All phyla except Phaeophyta include unicellular forms which may be motile
or non-motile. The motile type is of two kinds, the flagellated kind moving
by means of flagella (found in all phyla except Cyanophyta and Rhodo-
phyta), and the rhizopodial kind having the fine protoplasmic projections
(the rhizopodia) and showing an amoeboid movement (Xanthophyta and
certain other algae; Fig. 2-7 A). The cell envelope ofthe rbizopodial forms
is characteristically periplastic and rather soft, thus permitting extensive
changes in shape and size of the thallus. The ftagellated forms may be
periplastic, e.g., Euglena (Fig. 2-7 B), or may be provided with adefinite

flagellum

c .0 E

Fig.2-7. A, Chrysamoeba, motHe rhizopodial unicen; B, Euglena, motHe ftagellated

unicell; C, Phacotus, motile encapsuled unicell; D, Chlorella, non-motile, coccoid
unicell; E, Spirulina, spiral filamentous unicell. (Figs. A, C and E after Fritsch. 1935.)

cell wall, e.g., Chlamydomonas. External to periplast, some flagellates have

a special envelope which is provided with pores for tbe protrusion of the
flagella. The calcareous envelope is separated from the cell proper by a
space (Fig. 2-7 C). Such forms are known as the encapsulated forms. In
some the capsule may become calcareous.
Tbe non-motiIe or coccoid type includes forms of diverse shape and size.
They are provided with a rigid cell wall and are non-flagellated. Some
typical examples are Chlorella (Chlorophyta; Fig. 2-7 D) and Synechocystis
(Cyanophyta). In Spirulina (Cyanophyta) the cell is elongated into a helical
filament (Fig. 2-7 E).

MULTICELLULAR HABIT
Depending on the manner in which cells are produced and arranged during
thc vegetative phase, three principal types of habit are recognized-colonial.
filamentous and sipboneoDs.
22 A TEXTBOOK ON ALGAB

Colonial Habit
A colony is a group of separate cells generally simiJar in structure and
function and aggregated together by a muciJaginous envelope. On the basis
of morphology there are four main types of colonial organization-coenobial,
palmelloid, dendroid and rhizopodial. A coenobium has adefinite number
of cells arranged in a particular manner which is. determined at the juvenile
stage and does not increase during its subsequent growth even though the
cells enlarge. The coenobium may be motile or non-motile. In the motile
kind the cells are Bagellated, e.g., Volvox (Fig. 2-8 A). In the non-motile
type the cells are coccoid and more or less fused together, e.g., llydrodictyon
(Fig. 2-8 B).

l.
D
Fig. 2-8. A, Volvox, showing a flagellated motHe cocnobial colony (after Smith, 1955);
B, Hydrodictyon, non-motile colony (coenobium); C, Aphanorhece, palmelloid colony;
D, Chrysodendron, dendroid colony (after Fritsch, 1935).

Unlike the coenobium, the palmelloid, dendroid and rhizopodial types

of colony are not of constant shape and size and their cells divide leading
to increase in the size of the colony. In palmeIIoid colonies, the cells
remain embedded in a mucilaginous matrix of irregular shape and size (Fig.
2-8 C). The matrix is formed from the walls of individual daughter ceHs
wh ich gelatinize and mix together with the gelatinized walls of daughter
cells embedded in the common matrix. Some examples of palmeIIoid habit
are met with in Aphanothece (Cyanophyta) and Telraspora (Chlorophyta).
In the dendroid colonies the ceHs are united in a branching manner by
localized production of mucilage at the base of each cell (Fig. 2-8 D). The
whole colony looks Iike a tree in habit. Such an organization is found in
Ecbal/ocystis (Chlorophyta) and Chrysodendron (Chrysophyta).
The cells of rhizopodial colonies are united through rhizopodia, as in
Clzrysidiastrum (Chrysophyta).

FILAMENTOUS HABIT
A uniseriate row of ceHs joined end to end in a transverse plane through
middle lamellae constitutes a filament. Such ahabit arises as a result of
vegetative divisions being restricted to the transverse plane. In Cyanophyta
 COMPARATIVIl MORPHOLOGY 23

a filament includes both the trichome (i.e., the uniseriate row of cells) and
its sheath. Tbe filaments may be unbranched or branched. Tbe unbranched
filaments of Ulothrix (Chlorophyta) and Anabaena (Cyanophyta) do not
show any polarity but those of the Rivulariaceae (Cyanophyta) exhibit weil·
marked polarity (Fig. 2-9 A, B).

g~minate ps~lIdO­
branehes

reet system

~
';"
_lfl prostrate system

E F G
Fig.2-9. A, Ulothrlx, simple filamentoushabit; B, Calothrix, polarfilamentoushabit;
C, Scytonema, pseudobranching; D, Clodophora, true branching; E, Stigeoclonlum,
heterotrichous, branched filament; F, Voucher/a, branched siphoneous habit; G, U/va,
parenchymatous habit. (Figs. Band D after Fritsch, 1935.)

The branching of the filaments is oftwo kinds-false and true. In false

branching, which occurs in the Scytonemataceae (Cyanophyta), the trichome
generally fragments due to the degeneration of an intercaJary cell (or by the
formation of biconcave separation discs) after which one or both of its ends
adjacent to the dead cell grow out ofthe parent sheath, giving the semblance
of branching (Fig. 2-9 C). Sometimes false branching may arise through
24 A TEXTBOOlC ON ALG AE

loop-formatioD. In this method, a group ofintercalary cells ofthe trichome

becomes meristematic and for a time continues to divide at two localized
points, leading to the formation of a loop. The top (middle) cell of the
loop finaIJy degenerates setting free the two arms of the loop as a pair of
pseudobranches.
True branching results from repeated transverse divisions of the 'lateral
outgrowths produced by a few or many scattered ceIJs of the main filament.
The truly branched thalli are of four types: (1) simple branched filament,
e.g., Cladophora (Chlorophyta; Fig. 2-9 D); (2) heterotricbous in which the
thallus is differentiated into an erect and a prostrate system of branched
filaments (Fig. 2-9 E); (3) parenchymatous; and (4) pseudoparenchymatous
in which the thalli show uniaxial or multiaxial construction.
The heterotrichous habit is the most highly evolved type of filamentous
construction in algae. During its development a system of branched creep-
ing filaments is produced first and functions as the hold fast. These creeping
filaments give rise to a system of erect branched filaments. More or less
equally well-developed prostrate and erect systems are found in Stigeoc!o-
nium (Chlorophyta) and Ectocarpus (Phaeophyta). The more advanced
heterotrichous thalli show a progressive reduction or elimination of either
the prostrate system (e.g., Draparnaldiopsis) or the erect system (e.g.,
Coleochaete scutata).
The parenchymatous habit is a further variant of the filamentous cons-
truction and results from vegetative divisions taking place in more than one
plane followed by the failure of the division products to separate. Examples
are met with in U1va (Fig. 2-9 G) and Prasiola (Chlorophyta) and Porphyra
(Rhodophyta).
The pseudoparenchymatous habit results from a elose juxtaposition of
the branched filaments of a single or many axial filaments. If branches
from a single filament are involved, as in Batrachospermum, the thallus
construction is spoken of as uniaxial. If branches of many axial filaments
aggregate together, the thallus is multiaxial, e.g., Nema/ion.
The vegetative growth by cell division of filamentous and parenchyma-
tous thalli occurs in four ways-diffuse, intercalary, tricbotbaIlic and apicaI.
In simple filamentous forms like U10thrix and Nostoc, the growth of the
thallus is diffuse because each vegetative cell is potentially capable of growth
and division. A good example of intercalary growth is fumished by the
Laminariales (Phaeophyta) in wh ich the growth of the thallus is brought
about by the activity of a meristem located at the junction of the stipe and
the bl ade. The trichothaIIic growth is also intercalary but the meristem is
located at the base of a terminal hair, e.g., in Rivulariaceae, and CutJeriales.
The thalli of the Charophyceae, Dictyotales and certain other algae grow
by the activity of a single or a group of apical cells.

SIPHONEOUS HABIT
A siphoneous thaJlus is multinucleate and lacks septation except durinS the
 COMPARATIVE MORPHOLOGY 25

formation of reproductive organs. Tbe simplest form of such an organi-

zation is found in Protosiphon (Chlorophyta) and Botrydium (Xanthophyta),
both having a vesicular, uniceIIular thallus attached to the substratum by a
simple or branched rhizoid. In Vaucheria (Fig. 2-9 F), the siphoneous
filament is branched, advanced members of the Siphon ales (Chlorophyta)
show uniaxial and multiaxial forms.

TEST QUEST/ONS

1. Would you regard the cell waU as an essential part of an algal cell? Explain.

2. Which of the following parts of an algal cell is alive: cell wall, protoplasm, vacuole?

3. Differentiate between a chloroplast and a chromatophore and justify the suggestion

that variation in the external morphology of chromatophores is of IiUle significance
in the phyletic classification of algae.

4. Discuss the statement that electron microscopy has in no way alte red the primary
basis of c1assification of algae as previously emphasized by light microscopy.

5. What arguments can you offer in favour of the view that unicellular flagellates are
the most primitive forms of green algae?

6. Why is the motile colonial habit considered to have ended with a Volvox-Iike form?

7. On what grounds is the heterotrichous habit considered to represent the most highly
evolved type of filamentous organization in algae? What light does this habit
throw on the origin of land plants ?

8. Even though the basic structural unit of photosynthesis is the same in both algae
and higher plants, yet the chromatophore organization in the former is considered
more primitive than in the laUer. Why?

SELECTED REFERENCES

Haine, L. A. and Craigie, J. S., Studies on the algal cuticle, J. Phycol., 5, 89-102 (1969).

Leedale, G. F., Phylogenetic aspects of nuclear cytology in the algae. In Frederick,

J. F. and Klein, R. M. (eds.) Phylogenesis and morphogenesis in the algae, Ann.
N.Y. Acad. Sei., 175, 429-53 (1970).

Manton, I., Some possibly significant structural relations betwecn chloroplasts and
other cell components. In Goodwin, T. W. (ed.) The Biochemistry of Chloro-
plasts, Vol. I, 23--47, Academic Press, New York (1966).

Raven, P. H., A multiple origin for plastids and mitochondria, Science, Wash., 169,
641-46 (1970).
 3
Reproduction and Perennation
REPRODUCTION

It is a process by which a Iiving organism propagates and multiplies, and

increases the number of its individuals. Being devoid of the capacity for
active locomotion, most plants depend on dispersal of reproductive bodies
for movement from one favourable habitat to another. The algae generally
reproduce in three ways-vegetative, asexual and sexual.
Vegetative propagation does not involve rejuvenation ofthe protoplasts.
and the cell walls of its reproductive units are either entirely parental, or
partly parental and partly new. Asexual reproduction, however, involves
rejuvenation of the protoplasts, and the reproductive bodies are either naked
or provided with a newly synthesized wall of their own. Further, asexual
reproduction is accomplished by using specially differentiated cells that are
capable of directly multiplying the alga. Both the vegetative and asexual
reproduction are accessory means of propagation and are not concerned
with bringing together in one cell Iineage the genes from different cell
lineages.
In contrast to vegetative propagation and asexual reproduction, the
sexual reproduction involves alternation of syngamy (fusion of male and
female gametes) and karyogamy (fusion of gametic nuclei) on the one hand,
and meiosis on the other. This leads to the creation of new combinations
of genes by pooling together in one line of descent the genes derived from
two different parents, thus resulting in a reshuffiing ofthe gene material. The
recombinants so produced can then be subjected to natural selection. In
procaryotic organisms recombinants are formed by such processes as con-
jugation, transformation and transduction, none of which involves true
karyogamy and meiosis. This genetic recombination in bacteria and blue-
green algae accordingly falls short of the true and complete definition of
sexuality in the sense adopted here, even though it fulfils the samefunction
as true sexuality.

VEGETATIVE PROPAGATION

Vegetative propagation is very common in algae and takes place in a variety

of ways. In unicellular algae, flagellates, desmids and diatoms, a cell may
divide into two halves to produce new individuals. In multicellular forms
the thallus often breaks into small fragments each of which then grows to
become a new plant. Dictyosphaerium and Aphanothece multiply by splitting
 REPRODUCTION AND PERENNATION 27

of their colonies. Proliferation of fiJamentous thalli involves two methods-

fragrnentation, and hormogone formation. Tbe fragmentation of filaments
may be either due to rnechanical pressure exerted on them, or by dissolution
of the cross walls because of the developrnent of shearing strains by differ-
ences in turgor pressure between adjoining cells. In such algae as Sticho-
coccus, the tendency to fragment is so pronounced that filaments Ion ger
than a few cells each are seldom found.
Filamentous blue-green algae propagate vegetatively through a special-
ized process known as hormogone formation. The hormogones, which are
short fragments of a trichome, are produeed either as a result of fragmenta-
tion, or by the formation of biconeave, intercalary separation discs spaced
quite a few cells apart from eaeh other. The vegetative ceIJs Iying between
two separation discs eonstitute a hormogone. Devoid of a sheath and
endowed with the power of rnovement, the hormogones eonstitute an effi-
cient and quick method of vegetative propagation and dispersal of the
species during environmentally favourable growth periods.
Some aJgae reproduce by means of adventitious thalli as in Dietyota,
or by the formation of a few-celled branches, known as propaguJes, which
beeome detaehed from the parent and subsequentJy develop into new plants.
In Protosiphon, vegetative propagation takes plaee by budding of the vesicle.
In species of Trentepohlia eells of the prostrate system form a powdery
mass which is dispersed by wind and ean give eise to new plants.

ASEXUAL REPRODUCTION
Asexual reproduetion involves the formation of one or more of the foIJow-
ing types of spores: zoospores, apJanospores, autospores, hypnospores,
carpospores, tetraspores, endospores and exospores. These may be forrned
in any vegetative eell (Ulothrix), or in specially differentiated eells ealled
sporangia (Trentepohlia, Ectocarpus and Dietyota).
Tbe zoospore which probably constitutes the most important kind of
asexual spore, is a motHe body equipped with one or more flagella and
eommonly eontains a chromatophore, a nucleus and an eye spot. Aplano-
spores are non-flagellated, non-motHe and have adefinite eell wall distinct
from the parent wall. If the aplanospore, while still within the parent
wall, appears like the parent cell in all respeets exeept size, it is spoken of
as an autospore. Tbe endospores and exospores of Cyanophyta are
analogous to the aplanospores of Chlorophyta but not homologous beeause
the term aplanospore indicates a derivation from the motile zoospores
wh ich are not met with in blue-green algae.
The zoospores may be biflagellate (e.g., Eetoearpus), quadriflagellate
(e.g., macrozoospores of U/othrix), or multiflagellate (e.g., Oedogonium,
Vaueheria). During zoospore formation in Oedogonium, the entire proto-
plast assumes a pear shape and escapes from the cell and develops a ring of
ftagella around a colourless circular area towards its anterior end. In other
algae, e.g., C/adophora, the protoplast of a cell may give. rise to 8, 16, 32,
28 A TEXTBOOK ON ALGAE

or more zoospores by undergoing repeated divisions either after the comple-

tion of aH nuclear divisions (simultaneous division) or after each nuclear
division (successive division).
Aplanospores are produced in Ulothrix, Microspora and certain other
algae, and may be formed either from the entire protoplast of acelI, or a
single cell may give rise to many aplanospores. Autospore formation
occurs in many members of the Chlorococcales, e.g., Chlorella.
Endospores are produced by the division of the protoplast in three
planes, e.g., Pleurocapsa and Dermocarpa. In exospore formation, which
occurs in Chamaesiphon, the cell wall ruptures apically and the protoplast
thus exposed successively abstricts spherical spores.
The asexual spores produced by meiotic division in the unilocular
sporangium of Ectocarpus, in the tetrasporangium of Polysiphonia, and
during germination of some zygotes are known as meiospores or gonospores.
Such spores are unlike those produced by mitotic division (mitospores) in
that they are haploid and are concerned with the perpetuation of only the
haploid thalli. The mitospores may be either haploid or diploid and
accordingly perpetuate the haploid or the diploid generation.

SEXUAL REPRODUCTION
Typicalor conventional sexuality consisting of karyogamy and meiosis is
brought about by a variety of methods invariably involving the union of
the male and the fe male sexes. The reproductive units participating in the
sexual fusion are variously named according to the morphology, behaviour,
and the nature of the organs in which they are produced. The dictionary
meaning of the term gamete is "a germ cell carrying with it the implication
ofeither sex", and for sexual reproduction to occur, the gametes ofopposite
sexes must be present.
If the male and female gametes are ftagellated and motile, they are term-
ed zoogametes. Aplanogametes are non-ßagellated and may show amoeboid
movement. The antheridia and oogonia are the weil differentiated game-
tangia producing, respectively, the male and female gametes which differ
in morphology and behaviour. The gametes produced in antheridia are the
ßagellated, motile antherozoids (or spermatozoids), whereas the gamete
produced in the oogonium is the non-motile, non-ßagellated egg or ovum.
In Rhodophyta the oogonium having a swollen base (containing the egg)
and asiender projecting neck, the trichogyne, is called a carpogonium, and
the non-motile, non-f1agellate male gamete is termed spermatium, the struc-
ture producing it being known as a spermatangium.
True sexuality has been established for most members of the Chloro-
phyta, Phaeophyta and Rhodophyta and for some members of the Xantho-
phyta, Bacillariophyta, Pyrrophyta and Chrysophyta. In the Cyanophyta,
true sexuality in the sense of karyogamy and meiosis is lacking but a
parasexual phenomenon involving recombination of genes in a single line
of des cent from two genetically distinct parents has been reported in three
 REPRODUCTION AND PERENNA TION 29

different genera-Anacystis, Cylindrospermum and Anabaena.

In unicellular forms, sexual fusion may take pI ace between male and
female (or minus and plus) gametes produced either by division of proto-
plasts ofthe parent cells, or by fusion of the mature individuals themseIves.
The Iatter phenomenon, known as hoIogamy, occurs in a few species of
Ch/amydomonas. Besides, three other modes of sexual reproduction based
on the morphology and behaviour of the mating gametes are known. These
are isogamy, anisogamy and oogamy. In isogamy, the gametes of a fusing
pair are morphologically as weil as physiologically alike and are distinguish-
able genetically as plus or minus mating types. In anisogamy the gametes
differ in size and motility, the smaller, more active gametes being male and
the larger, less active being female. In oogamy, the gametes of the two
sexes are sharply differentiated, and with a few exceptions are produced in
specialized organs known as the antheridium and the oogonium. Both in
isogamy and anisogamy, the garnetes of opposite sexes mate outside the
parent cells except in the Conjugales in which mating is facilitated by the
formation of a conjugation tube and may occur either within the conjuga-
tion canal itself, e.g., Zygnema, or in the cell containing the fern ale (plus)
garnete, in which case the male gamete moves to the fern ale through the
conjugation tube, e.g., Spirogyra. In the majority of oogarnous green algae
the egg is not liberated from the oogonium and fertilization takes place
inside the oogonium.
Isogamy, anisogamy and oogamy are stages of a progressive series in
the differentiation of gametes and this series seems to have evolved indepen-
dently in different phyla. Thus, the Chlorophyta, Phaeophyta, Xanthophyta
and Bacillariophyta include forms ranging from isogamy through anisogamy
to oogamy. However, in diatoms isogamy is secondary and oogamy is
primitive. The members of Rhodophyta are characteristically oogamous.
In the Bacillariophyta, only those forms as show a progressive decrease in
cell size during vegetative multiplication undergo sexual reproduction; and
the formation of auxospores from the zygote serves as a device to offset
reduction in cell size resulting from continued vegetative propagation.
Sexually reproducing thalli may be homothallic (monoecious) showing
fusion between gametes derived from a single parent individual, or hetero-
thaIIic (dioecious) permitting fusion between gametes of different parentage.

ORIGIN OF SEX

The origin of sex is inherently related to the origin of gametes which are
thought to have been derived from asexual zoospores that had become too
small or too weak to produce healthy new plants by themselves (Fig. 3-1).
In forms such as Chlamydomonas and Ulothrix with isogamous sexual
reproduction, the gametes are very much Iike the zoospores except for the
difference in size. In the isogamous algae, both plus and minus gametes
resemble the zoospores whereas in the anisogamous or oogamous species,
only the motiIe gametes (antherozoids) show this resemblance. The zoo-
30 A TEXTBOOK ON ALGAE

spores and gametes differ fundamentally in tbe way they give rise to new
plants: the former germinate directly into new plants whereas the latter

Fig. 3-1. Hypothetical diagram iIIustrating how sex might have originated in such
forms as Ulothrix. Note the four types of swarmers (indicated by different kinds of
arrows) and their fate, and also the relative sizes of the germlings produced.

copulate in pairs to form tbe zygote whose division products ultimately

develop into new plants. How a mere variation in size can account for such
a fundamental difference in their behaviour is tbe basic question. Tbe
larger zoospores contain the minimum level of factors tbat help them grow
individually into new plants. Comparatively, tbe smaller zoospores are
not self-sufficient and therefore fail to grow. The chance fusion of such
 REPRODUCTION AND PERENNATION 31

zoospores in pairs is considered to restore thc factors to optimum level for

subsequent development. Tbe precise nature of tbe factors is not known
but it is Iikely tbat cbemical, bormonal or electrical components may be
involved.
Tbe difference in size of tbe larger and weaker zoospores wbicb can
potentially act as gametes, results from tbe number of divisions tbe parent
protoplast undergoes. If tbis number is smalI, tbe products are large and
bebave as zoospores, and if tbe number is large, tbe resulting cells are small
and bebave as gametes. Tbis view is supported by tbe observation tbat
under certain conditions if tbe gametes fail to copulate they can act Iike
zoospores. All these facts indicate tbat the gametes are derived from
zoospores and that sex in algae bas originated as a result of accidental
fusions between small-sized, weaker zoospores.
Tbe origin of gametes (and hence tbe sex) from zoospores is a hypo-
thesis developed mostly from observations made on materials collected
from nature and it has yet to receive conclusive experimental support. To
our knowledge, neither genetical nor physiological factors that might be
involved in regulating the zoospore-gamete balance sheet have been experi-
mentally verified.
The mechanism of sexuality has been investigated in detail mostly in
the genus Ch/amydomonas, especially in heterothallic and isogamous species
in which.sex is determined genotypically. In these species, gametogenesis
is induced by a variety of environmental factors such as light, temperature,
nitrogen content, and pH. The mating reaction between gametes proceeds
through a number of distinct phases: (I) morphological and physiological
differentiation leading to gamete formation, (2) cIumping, (3) pairing,
(4) cell fusion, and (5) nuclear fusion. Recent work on mutants of Ch/amy-
domonas lacking flagella has indicated that the mating type or sexual sub-
stances are located on the flagellar tips of tbe isogametes. Tbe mating
reaction between such gametes is initiated by the agglutination of tbeir
flagellar tips and then gradually proceeds downward tiII the bodies of tbe
two gamet es finally fuse.

LlFE CYCLES AND ALTERNATION OF GENERATIONS

The zygote is a diploid cell resulting from gametic fusiop- and is provided
with a thin or thick wall. Thin-walled zygotes often germinate directly,
without undergoing any resting per iod , as in most marine algae. Thick-
walled zygotes generally have their walls differentiated into three layers and
such zygotes (zygospores or oospores) normally remain dormant for a while
be fore germination. Botb karyogamy and meiosis may take place in the
zygote which on germination produces a haploid plant. If these processes
become separated in time and space, the zygote merely remains a place for
karyogamy and its germination gives rise to a diploid plant. It is during
sporogenesis by the diploid plant that meiosis occurs. The interpolation
of a diploid somatic phase between karyogamy and meiosis probably makes
32 A TEXTBOOK ON ALGAE

the individuals more stable, genetically and physiologically, with the conse-
quent attainment of greater adaptability and survival.
The growth and development of an alga proceeds through a number
of distinct morphological and cytological stages. The sequence of these
orderly changes is termed Iife cycle or life history. Accordingly, a Iife eycle
has two aspects, the somatie or morphological, and the cytological or
chromosomal. A generation is regarded as a somatie phase; the alterna-
tion of generations refers to a situation in which a plant has two somatie
phases: (I) the haploid or gametophytie, and (2) the diploid or sporophytie,
whieh regularly alternate during the Iife cycle. Both morphologieal and
cytological considerations have led to the recognition of four main types
of algal Iife eycle: haplontie, diplontic, isomorphie and heteromorphie.
The haplontie type includes algae in wh ich the somatic phase only is
haploid and tbe diploid stage is restricted to the zygote whereas in the
diplontie type the vegetative phase only is diploid and the haploid phase
is restrieted to the gametes. In between these two types are those Iife
cycles in which" a regular alternation of gametophytie and sporophytie
generations occurs. If the two alternating somatie phases of the life cycle
are morphologically alike, the life cycle is termed isomorphie and if the
two phases differ morphologieally, it is termed heteromorphie.
The haplobiontie haplontic life eycle is found in such algae as
Ch/amydomonas, Ulothrix, Oedogonium (Fig. 3-2) and Chara. The dip-
lontie type is characteristie of Ch/orochytrium, most Siphonales, Fucales
(Fig. 3-3) and Baeillariophyta. Most Nemalionales (Fig. 3-3) have a
haplobiontic life cycle in which the haploid gameophytic phase alternates
with a haploid asexual carposporophytie phase; the life eycle in this case is
biphasie with two morphologieally dissimilar, but cytologically similar,
generations and the only diploid stage in the life eycle is eonstituted by the
zygote itself. The isomorphie life cyele occurs in Ulvaceae, Stigeoc/onium,
Draparna/diopsis and Ectocarpus (Fig. 3-4) and the heteromorphie type is
charaeteristie of the Laminariales (Fig. 3-4). The Iife eycle of the.Florideae
is complieated by the interpolation of a parasitic earposporophyte between
the gametophytic and sporophytic generations. The carposporophyte is a
carpospore-producing sporophyte eonsisting of a cluster of sterile cells and
earposporangia which are borne terminallyon special filaments ealled
gonimoblasts. In more advanced members of the Florideae, e.g., Poly-
siphonia (Fig. 3-5), the life cycle is triphasie, involving a succession of three
generations: gametophytic, carposporophytie, and tetrasporophytie. The
carposporophyte and the tetrasporophyte are diploid and the latter is
morphologically similar to the gametophytie plant.
Aecording to Von Stotsch (personal communication), most probably
there are no .haplobiontic Nemalionales at all and even Batrachospermum
seems to lack zygotie meiosis.
Drew (1955), and Chapman and Chapman (l961) have reviewed the
algal life eyeles, introdueing eomplex terminologies. Aeeording to the
nuelear eyele a haplont is one in whieh thezYgote is diploid and meiosis
 REPRODUCTION AND PERENNATION 33

takes place during the germination of the zygote; a diplont is one in which
the plant is diploid and the gametes alone are haploid. A haplobiont
possesses only one kind of plant (a haploid or a diploid) whereas a diplo-

HAPLONT
(OEOOGONIUM)

0 ~
'000/
'./~O
MEIOSPORES'
(N) 0
A\\
\
SYNGAMY
J GAMETES
(NI

ZYGOTE
(2N)

Fig. 3-2. Haplobiontic Iife cycles.

HAPLONT
, (BATRACHOSPERM~UM)
~

CHAN~
~\)(N) " ßAMElES

o~
M0N05PORE
'OCARPOSPORE
"-~;.
~
" (N)
Sm3AMY
~ bZ'IGOTE
(N) CARPOSPOROPHYTE (2N)
(N)

Fig. 3-3. Types of Iife cycle.

biont possesses both the haploid and the diploid plants in its life cycle.
These workers favour the idea that an algal life cycIe should be quaIified
both cytologically and morphologically. Based on morphology, the life
cycle can be of three types: (I) the monomorphie, in which the gameto-
~ phytic and sporophytic generations are morphologically alike, (2) the
~ dimorphie, in wh ich the two generations are morphologically different, and
t (3) the trimorphie, in whieh a sueeession of three morphologieal types
...; oecurs. In Drew's terminology, the life eyele of Eclocarpus and C/adophora
34 A TEXTBOOK ON ALGAE

is monomorphic, dipIobaplont and that of the Nemalionales likc Batracho-

spermum trimorphie, haplont.

~r. GAMETES
~
q ')2MElOSPORES
~~(N)
• ,.<1'\

o ZYGOTE
(2N) ' ";4

oft-EIOSPORESISOMO~PHIC TYPE ~' HETEROMORPHIC TYPE

(ECTOCABPUS/ULVA) (N) (bAMINARIA)
- -
~..o (N) I!l.
ft- ZYGOTE ~
(2N) " ;.(.

Fig. 3.1. Isomorphie and heteromorphie alternation of generations.

TETRASPORE
(N) ~~o
R............
~ ... -o GAMETES
(N)
DPLOBIONTIC (ISOMORPHIe)
--SYNGAMY
( POLYSIPHONIA)

arV~~POSPOROPHYTE
O (2N)

CARPOSPORE
(2N)

Fig. 3-5. Isomorphie alternation oe generations.

PERENNATION
When the environmental eonditions beeome unfavourable, many algae form
resting stages for perennation, i.e., for tiding through adverse periods. In
many Chlorophyeeae, the only perennating stage in the life eycle is thc
thick-wallcd zygospore or oospore that commonly has a highly viseous
protoplast containing an orange or red oity pigment known as haemato-
cbrome. They ean tolerate unfavourable environmental eonditions without
any adverse effeets.
 REPRODUCTION AND PERENNATION 35

Many terrestrial or subaerial blue-green algae can perennate as such, i.e.,

without any apparent modification of the vegetative cell or filament. The
thick, stratified or gelatinous sheaths around their celIs or trichomes, tbe
gel-like consistency and viscosity of their protoplast, and the virtual universal
absence of vacuoles in their cells, are probably some features that enable

l ..
i. .\
i :

c
B
A

D
Fig. 3-6. A, Cylindrospermum, akinete and hcterocyst; B, Acelabularia, vertical section
through fertile disc showing gametangial rays and cysts (after Bold, 1967); C, same
showing germination of cyst; D, Wesliella, hormocysts and heterocysts.

them to endure and survive unfavourable conditions without much structuraI

modification.
Perennation in other algae involves tbe formation of specially differen-
36 A TEXTBOOK ON ALGAE

tiated resting cells. Some common examples are:

1. AkiDetes or "Spores". These are enlarged vegetative cells which
become thick-walled and store abundant reserve food material (Fig. 3-6 A).
With the advent of unfavourable conditions the ordinary vegetative cells
die whereas the akinetes successfully endure the harsh c1imate. When
favourable conditions recommence during the next spring or rainy season.
the akinete germinates and produces a vegetative filament, e.g., Pithophora.
Gloeotrichia and Cylindrospermum (Fig. 3-6 A). Akinete formation is wide-
spread in the nitrogen-fixing blue-green alga, Aulosira /ertilissima. which
occurs abundantly in tropical rice fields. Here, the vegetative filaments
become transformed into chains of akinetes (spores) when the stagnant water
in the fields begins to dry up, and it is in this sporulating condition that the
alga tides over dry and hot periods.
2. HypDospores or Cysts. Like akinetes, these are also thick-walled
but, as a rule, are formed by repeated divisions of the protoplast rather
than by the modification of a cell into a single akinete as in Acetabularia
(Fig. 3-6 B, C). Protosiphon botryoides and Sphaerella lacustris also furnish
good examples of cyst formation. During conditions of drought or light
of high intensity, their cytoplasm breaks up into a number of thick-walled
hypnospores which in Sphaerella may remain embedded in a common
gelatinous matrix.
In Fritschiella tuberosa and Botrydium cysts are formed in the sub-
terranean parts. When favourable conditions recur, the cysts either grow
direct into a new cell or give eise to zoospores.
3. Hormocysts. These are hormogonia which, instead of being liberated
from the parent filament, remain in situ and become thick-walled. They
mayaiso be regarded as many-celled akinetes. Typical hormocysts are met
with in the blue-green alga Westiella (Fig. 3-6 D).

TEST QUEST/ONS

1. Discuss the advantage of sexual reproduction when an alp can effectively multiply
by asexual or vegetative means.

2. Why do algae produce a variety of spores, e.g., zoospores, aplanospores, and hypno-
spores, when the function of asexual reproduction can be effectively discharged by
zoospores alone?

3. Distinguish between: (a) alcinete and hypnospore; (b) autospore and endo-
spore; (c) aplanospore and autospore; and (d) vegetative cell division and binary
fission.

4. How does the sexuality in eucaryotic alpe differ from that in bacteria ?

S. Why are isogamy, anisogamy and oogamy regarded as progressive stages in the evo-
lution of gametes in algae ?

6. What is the evolutionary significance ofthe algae with heteromorphie life cycle?
 REPRODUCTION AND PERENNATION 37

7. Occasionally, when gametes fai! to fuse, they may direct1y give rise to new plants.
What is its significance in relation to the origin of sex in algae ?

SELECTED REFERENCES

Allen, R. D., The moment of fertilization, Sclem. Am., 201, 124-34 (1959).
Chapman, D. J. and Chapman, V. J., Life-histories in the algae, Ann. Bot., N.S., 25,
547-61 (1961).
Drew, K. M., Life-histories in the algae with special reference to the Chlorophyta,
Phaeophyta and Rhodophyta, Biol. Rev., 30, 343-90 (1955).
 4
Economic Importance
With the phenomenal increase in urban population and industrial growth
during the past few decades, some of the major pr<?blems of a basic nature
confronting mankind are those of quantity and quality offood, feed, drink-
ing water, disposal ofsewage and industrial ~astes, and of proper conserva-
tion of soils and waters. Suitably controlled algal growth can provide
positive help in tackling these problems.

POSITIVE IMPORTANCE

EDIBLE FORMS

People of coastal countries, e.g., China and Japan, have Ions been using
seaweeds and certain other algae as a source of food. Some of the more
commonly used are Porphyra, Vlva, Alaria, Chlorella, Chondrus (Fig. 4-1 A-E),
Rhodymenia and Nostoc. These not only form an important ingredient of
soups but are also used for flavouring meat. Sometimes the blades and
stipes of seaweeds are eaten after frying, with or without salt.
Besides being rich in organic iodine, which serves as aprecursor of thy-
roxine, seaweeds contain significant amounts of other mineral elements
considered indispensable for a well-balanced diet. Vitamins B, C, folie acid,
and niacin are also found in them. They are, however, poor in protein con-
tent.
Among all the edible seaweeds the most important is the red alga
Porphyra tenera which is cultivated in shaIIow sheltered seacoasts on a com-
mercial scale in Japan. Chemical analyses of the processed alga by several
Japanese scientists established that it is very rich in proteins (3{}-35%) and
carbohydrates (4{}-45%). Fairly high concentrations of vitamins A, B, C
and niacin were also detected.
In the farming of P. tenera a talge number of bamboo networks, each
about 40 met res long and 1.25 metres wide, are placed horizontally so as to
be near the surface of water in the shallow sea. These frameworks are held
in place with the help of bamboos fixed verticaIIy in the seabed. Such
bamboo frames provide a favourable habitat for the germination of mono-
spores into young thalli. After about a month the entire bamboo networks
together with the young thalli are transferred to river estuaries along the sea
where the concentration of such nutrients as nitrogen, phosphorus and
others is quite high. These substances stimulate the growth of the trans-
plants. A number of Porphyra crops can be harvested successivcly between
 ECONOMIC IMPORTANCE 39

November and April. Harvesting is carried out by plucking mature thalli

by hand, washing and cutting them into pieces which are then resuspended
in water. The algal suspension is filtered through sieves made of bamboo

Fig. 4-1. Some common edible algae. A. Porphyra; B, Ulva; C. Alaria; D, Chlorella;
E. Chondrus.

frames, dried and then pressed into sheets for sale. These Porphyra sheets
are eaten with rice. Porphyra soup is highly prized in Europe and else-
where.
40 A TEXTBOOlC ON ALGAB

Tbe late Professor K. M. Drew-Baker of the University of Manchester,

on the basis of her researches in the fi,fties, conc1usive1y established that the
alga then known as Conchoce/is, found growing on oyster shells, was actually
a juvenile stage in the life cyc1e of Porphyra, the important "Nori" of the
Japanese seaweed industry. Her recognition of the Conchocelis stage in
shellshasenabled the Japanese to devise an artificialmethod ofspore cultiva-
tion. Tbe significance ofthis discovery was so great that it saved the Japa-
nese Porphyra industry from near collapse.
Many developing countries are facing grave food shortage. In order to
meet this challenge, suitable substitutes for conventional foods have to be
evolved. Added to this is the alarming rate ofpopulation increase for wh ich
additional accommodation has to be provided at the cost of agricultural
land. These facts necessitate a search for alternative crops that could be
grown a number of tiIiles in a year within minimum space at a cost weil
within the reach of the common man, and with a nutrition al value compar-
able to that of the conventional foods. Researches carried out in Germany,
Japan and the United States on the large scale production, chemical com-
position and nutrition al value of algae suggest that they are not only suitable
substitutes for conventional crops but also yield greater amount of food per
unitlandarea(Dawson,1965). Moreover, for their growth the algae require
simple inorganic nutrients, light and carbon dioxide (C02) wh ich they can
easily obtain when grown in sewage oxidation ponds. Even otherwise their
growth requirements can easily be met without much economic burden.
Although Porphyra has been in use as a supplementary diet in Japan for
a long time, its restriction to marine habitat, slow growth rate, and relatively
poor vitamin and protein contents make it a less favourable algal food than
Chlorella. The latter grows very fast under different conditions and there-
fore a much greater number of crops can be harvested. It is also richer in
protein and lipid content than Porphyra and contains a wider variety as weIl
as higher concentrations of vitamins. The nutritional value of Chlorella is
comparable to that of a mixture of soya beans and spinach. Simple growth
requirements, production of Httle waste material and high growth rate are the
features which ha ve encouraged the mass cultivation of Chlorella in Germany,
Japan and the United States.
Chlorella grows actively in culture when provided with sunlight (or
artificiallight), C02 and mineral nutrients. A provision has also to be made
for frequent or continuous agitation ofthe culture suspension so as to prevent
the cells from settling down, and to ensure a proper distribution ofnutrients,
C02 and light. Tbe open circulation syst~m of mass culture devised by
Tamiya (1960) in Japan consists of an open, shallow culture pond contain-
ing rotating pipes with numerous fine jets. These jets agitate the culture
suspension and keep it enriched with C02. At frequent intervals aliquots
of culture suspension are pumped out and the algal cells harvested by centrifu-
gation. The harvested cells are washed and dried in vacuum or extracted
with methanol to remove the pigments. Finally the dried algal mass i!\
ground in a mill and stored in powdered form. On an average an annua}
 ECONOMIC IMPORTANCB 41

yield of nearly 13 metric tons per acre can be expected if the cultivation
continues throughout the year.
Cultivation, harvesting and drying are the three major processes for
turning Chlorella into foodstuffs. Out of these, the harvesting technique
commonly employed, is stilI the most inefficient and expensive so that the
greatest hurdle in the large scale exploitation of Chlorella as food is its high
cost of production. Further, the indigestibility of the algal cell wall pre-
vents its direct use as food on an industrial scale.
Researches are also being conducted in the United States on Chlorella
and Synechococcus regarding their possible use in space ships and nuclear
submarines as oxygen regenerating and food and water recycling organisms.
In addition to their direct use as food, many algae provide meal to
human beings indirectly by virtue of their strategie position in the food
webs in natural habitats. In both fresh and marine waters, algae are in gest-
ed by lower animals (protozoa, insect larvae, copepods and rotifers) which
in turn are eaten by higher animals such as fish. Fish constitutes an impor-
tant source of protein.

MINERALS AND ELEMENTS

The brown seaweeds popularly known as kelps are a rich sourceof soda,
potash, iodine and alginic acid. Besides, they can also yield a good amount
of ammonia and tar or charcoal when carefully processed.
The weed is collected, air dried and then burnt in cylindrical kilns.
The resulting ash is used to recover soda and potash which are used in the
manufacture of soap, glassware and alum.
In Japan, the extraction of iodine from kelps represents 5-7% of the
world production. The genera commonly used for iodine production are
Laminaria, Fucus, Ecklonia and Eisenia. It is the most important compo-
nent of thyroid hormones and therefore has got a large application in
chemotherapy of thyroids. The high iodine yielding Phaeophyta are used
for the treatment of goitre as weIl.
Seaweeds are also rich in copper, iron, zinc, cobalt, vanadium,
molybdenum, manganese, boron and chromium. This has led to their use
as supplements to fodders and fertilizers.

ALGINATES
Alginates are the saIts of alginic acid found in the cell wall of Phaeophyta.
The genera most commonly used for alginate extraction are Fucus, LAmi-
naria, Macrocystis, Cystoseira, Lessonia and Ecklonia. Flame-proof fabrics
as weIl as plastic articIes are prepared from aJginates. Plastic materials of
alginates find many applications in medicine, e.g., in the preparation of
dental impressions. Alginic acid can effectively stop bleeding and is em-
ployed as a highly efficient gauze in internaIoperations. Because of their
non-toxicity and coUoidal properties, alginie acid derivatives have a number
42 A TEXTBOOIC ON ALGAB

of applications in preparation of commercial products for public consump-

tion, e.g., in the preparation of sou ps, sauces and creams of various kinds
and of antibiotic (aureomycin) capsules. Alginates are extensively uscd as
thickeners in the cosmetic, textile and pharmaceutieal industries, and as
emulsifying agents in the preparation of poJishes and paints.

AGAR-AGAR
A jelly-Jike substance, agar-agar (often called simply agar), is a complex
polysaccharide of great economic value. It is extracted in water from certain
species of red algae belonging to Gelidium, Graci/aria and other genera
whieh produce and store it along with cellulose in their cell walls.
The most extensive use of agar is aS'a base for culture media for algae,
fungi, bacteria and tissues. It is also used as a stabilizer or emulsifier in
food, cosmetics, leather and pharmaceutical industries. Agar also finds
good application in the canning of fishes, sizing of fabrics and in paper
industry. It is often given as a laxative and is sometimes prescribed for
treating a prolapsed stomaeh.

CARRAGEENIN
ChemieaIly, carrageenin is very much Jike agar and occurs as a cell wall
polysaccharide esterified with' sulphate. It is usually extracted from
Chondrus crispus (popularly known as 'Irish moss'), though sometimes
species of Gigartina are also employed. Carrageenin is often used as a
remedy for cough and as a pharmaceutieal emulsifier. It is also utilized in
the textile, Ieather, cosmetics and brewing industries.

FODDER
The brown algae Ascophyllum, Laminaria and Fucus are used as stock feed
for sheep and cattle in maritime districts. In Ireland and Scotland there
are established industries for processing them into a commercial feed. The
seaweed meal is very nutritious because of its high vitamin and mineral
content. The milk produced by cows that feed on such meals is richer in
fat content than by those fed on conventional fodder. Likewise, hens fed
with seaweed meal produce eggs rieh in iodine.
The primary producers in aquatic habitats, incIuding the sea, are
mainly algae and therefore algal metaboJism and productivity has a great
bearing on the natural flora and fauna of the aquatic ecosystem. Nitrogen-
fixing blue-green algae maintain and regulate the nitrogen budget of aquatic
habitats, and by excretion of fixed nitrogen into the surrounding water, aid
the growth of other microorganisms in places where nitrogen is a limiting
factor (Stewart, 1967). Planktonic algae form the major food of protozoans,
crustaceans and fishes (Fig. 4-2). Marine flagellates and other microalgae
are cultivated for rearing the larvae of marine fishes, especially shell fish.
 ECONOMIC IMPORTANCE 43

Practical methods for increasing algal feed are devised by placing bunches
of bamboo in fish ponds; this provides an enormous surface ror the growth
of algal epiphytes which serve as food for fish. Vitamin-rich fish oils are
in fact derived from diatomaceous feed. Efficient pisciculture involves the
joint cultivation of fishes and algae, the former feeding on algae and the

Fig. 4-2. Role and significance of algae in biological food webs, amI in converting
waste materials into foodstuffs.

latter using the waste products of fishes and other organisms for their own
nutrition and growth.

FERTILIZER
Seaweeds, being rich in potassium, phosphorus, trace elements and growth
substances, are extensively utilized as man ure by people of coastal districts.
The . weeds are either allowed to rot in the field or composted with other
organic materials. The resulting manure, when added to agricultural fields,
not only enriches them in mineral nutrients but also helps in soil bin ding,
in breaking down clays and in promoting good crumb formation. Such
eoralIine or lime-depositing algae as Lithothamnion and Lithophy//um are
used for timing the crop fields.
Tbe Irish eollect and use Fucus as man ure on a large seale. In India, a
comparative study of tbe yie~d of 'bhindi' (Abelmoschus esculentus) grown
separatelyon a mixture of cowdung and wood ash manure, and on seaweed
manure, indicated the superiority of tbe latter over tbe former (Tbivy.
44 A TEXTBOOlt ON ALGAB

J96O). The seaweed manure, however, is poorer in nitrogen and phosphorus

than the farmyard manure. Tbe blue-green algae, which are rich in nitrogen
and phosphorus, are better fertilizers than seaweeds. In tropical countries,
bottom mud of dried up ponds is regularly used as manure in crop fields;
the manurial value is mostly due to the high content of blue-green algae.
A suitably blended mixture of seaweed and cyanophycean manures (e.g.,
bloom of Microcystis) may serve as an ideal fertilizer and this can relieve
the acute shortage of fertilizers in developing countries.
Tbe fertility of natural habitats benefits from algal growth in many other
ways as weil. Many algae found in such habitats are believed to produce
considerable amounts of extracellular substances, including organic acids,
amino acids and polypeptides which chelate organic and inorganic ions and
maintain them in astate readily available to plants. The extracellular
products also serve as sources of carbon and nitrogen for other micro-
organisms that regulate the overall balance of soil nitrogen, sulphur, phos-
phorus and carbon.
In respect of nitrogen status of natural habitats, the nitrogen-fixing blue-
green algae deserve special mention. These algae grow luxuriantly in tropical
habitats, e.g., rice fields. Experiments conducted with Tolypothrix tenuis
in Japan and with Aulosirafertilissima in India have shown that the yield
of paddy is substantially increased following the inoculation of fields with
these algae. Algologists at the Central Rice Research Institute, Cuttack
(India) inoculated rice fields with four species of nitrogen-fixing blue-green
algae; the grain yield increased by nearly 30 per cent.
In the Sambhar Salt Lake in Rajasthan (India), cartloads of algal mats
consisting of Anabaenopsis and Spirulina are produced annually, from Sep-
tember to December, and are employed by local farmers as man ure.

SOlL RECLAMATION
Singh (J 96J) has demonstrated that blue-green algae can be used to help
reclaim saline and alkaline wastelands in various parts of India. These
algae in general prefer alkaline pH, waterlogging and high humidity coupled
with high temperature for their growth, and can successfully grow in
alkaline and saline wastelands during summer monsoons when all these
requirements are avaiIable. Reclamation experiments conducted on waste-
land enclosed by a 0.5 metre earth embankment so as to encourage
waterlogging during the rains, resulted in decrease in pH and increase in
nitrogen, phosphorus and organic matter content ofthe field thus converting
it, in due course, into a fertile cultivable land.

ANTIBIOTICS
Some algae produce antibacterial substances eft'ective against a number
of pathogenic bacteria. ChIorellin, obtained from Chlorella, was the first
such substance. Extracts of Cladophora, Lyngbya and certain other algae
 ECONOMIC IMPORTANCE 45

kill strains of Pseudomonas and Mycobacterium and exhibit antiviral activity.

However, the antibacterial or anti viral substances concerned have not yet
been isolated in a pure form.
Certain Cyanophyta and Characeae have been c1aimed to possess lar-
vicidal properties since very few mosquito larvae occur in waters supporting
the growth of such algae.

DIATOMITE
It constitutes the cell wall material of diatoms. Large dcposits of diato-
maceous earth have been discovered in marine and freshwater basins.
Diatomite is highly porous and insoluble and therefore is ideally suited as a
filter for oils and for clearing solvents. Most industrial filtration devices
are made of diatomite. It is also employed in wine and paper industries.
An effective insulator, it is light, and fireproof and finds application in the
preparation of high temperature furnaces.

SEWAGE DISPOSAL
Sewage consists of water-borne domestic and industrial waste which is fairly
rieh in dissolved or suspended organic and inorganic constituents, but very
poor in oxygen. It harbours a number of faecal and other anaerobic
bacteria, some of which may be pathogenic. The sewage has to be treated
before its disposal for two main reasons. Firstly, the anaerobic digestion
of sewage produces stinking and offensive odours. Secondly, the contami-
nation of potable water with sewage infested raw material is potentially a
rieh source for inorganic nutrients such as nitrogen, phosphorus, potassium
and sulphur. This aspect of sewage assumes considerable importance in
view of the fact that many developing countries are facing an acute shortage
of inorganic fertilizers on the one hand and an explosive population increase
on the other.
Sewage oxidation ponds have been created for bringing aboutits complete
oxidation into mineral components. Such ponds support luxuriant growth
of unicellular algae, e.g., Chlorella, Chlamydomonas, Scenedesmus and Eug-
lena. These help in the bacterial decomposition of sewage by providing
oxygen; in addition, they recover the mineral nutrients from sewage which
would otherwise have been lost in the efHuent.
Sewage algae have also been used as animal feed, as man ure, and for
ingestion by fish fingerlings in fish ponds. The main limitation preventing
a wider exploitation of sewage algae is the lack of a cheap and effective
harvesting technique.

NEGATIVE IMPORTANCE
TOXICITY AND PARASIT15M
Prymnesiumparvum, Gymnodinium veneficum and species of Microcystiscause
mortality in fish, and in domestic animals that drink water infested with
46 A TEXTBOOK ON ALGAE

these algae. Various species of Gon)'aulax produce endotoxins which accu-

mulate in the digestive glands of shellfishes feeding on them. If such shell-
fishes are consumed by other animals, including man, paralysis and even
death may result.
Bloom-forming blue-green algae such as Microcystis aeruginosa,
Anabaenaftos-aquae and Aphanizomenonftos-aquae have been irnplicated in
animal poisoning in temperate countries. Depending on the time interval
that elapses between injection of the toxic algae into mice and their resulting
death, these toxins have been classified into three categories: Very Fast
Death Factor (VFDF), Fast Death Factor (FDF) and Slow Death Factor
(SDF), killing mice within 10min, 2 hr and 48 hr respectively.
If some of the toxic planktonic algae happen to be ingested either along
with drinking water or during swimming, they may cause various disease
syndromes. For example, Anabaena and Microcystis cause gastric trouble;
G)'mnodiniuin brevis produces respiratory disorders, and Lyngbya and Chlo-
rella are responsible for certain skin infections.
Species of the parasitic green alga Cephaleuros (Chlorophyceae) cause
'red rust of tea' and infljct heavy economic losses by seriously affecting the
yield of tea.

EFFECTS ON AQUATIC ANIMALS

The excessive growth of certain algae (Microcystis aeruginosa) in a body of
water often results in severe depletion of oxygen in the habitat. This leads
to mass mortality of fish due to suffocation. Sometimes, high temperature
and bright sunlight result in massive disintegrati on of algal blooms, releasing
into the medium their noxious components. Choking of the mouth or gills
of fish by these algae is also partly responsible for their death.

FOULING OF MARINE VESSELS

Some seaweeds may grow on the metal hulls and woodwork of ships and
boats producing a fouling, corroding and destructive efTect. A thick growth
of weeds sometimes results in considerable increase in friction between hull
and water thereby accentuating wear and tear and shortening the Iife of the
vesseI.

IMPORTANCE IN MUNICIPAL WATER SUPPLIES

The problems associated directly or indirectly with aJgal growths in water
reservoirs and water supplies are: (I) loss of recreational (swimrning and
aesthetic) and fishing values of pools, ponds and Jakes due to excessive
growth of Microcystis, Spirogyra, CladopllOra and Pithophora; (2) imparting
ofabnormal tastes and odours by the metabolie or decomposition products
of nuisance organisms such as Synura, Synedra, Asterionella, Anabaena,
Microcystis and Dinobryon; (3) clogging of water filters by Oscillatoria,
 ECONOMIC IMPORTANCE 47

Spirogyra and certain diatoms, thereby shortening the filter runs leading to
serious econoinic losses; (4) colouration of raw and finished waters due to
the presence of planktonic algae, e.g., Chlorella, Chlamydomonas, Ellglena
and Oscillatoria; (5) spoilage of the quality of commercial products of food,
pharmaceutical and pulp industries due to the use of waters contaminated
with algal slimes; (6) production of toxic substances; (7) corrosion of con-
crete and metallic walls of pipes and boilers by carbonic, oxalic and silicic
acids excreted by certain algae, e.g., Anacystis and Chaetophora; (8) inter-
ference with purification of water; and (9) changes in pH, C02, bicarbonate
and oxygen contents of water.
In water treatment plants, the raw water is chlorinated in order to kill
algae and other microorganisms. However, even this water is sometimes
not completely free from algae since some of them are chlorine-resistant
and their presence may alter its quality and potability.
Tbe only useful purposes tbe algae ss:rve in water supplies are: (I)
maintenance of aerobic conditions by checking putrefaction of organic
substances, and (2) rcduction of total hardness of water by consuming
bicarbonates and the nearly insoluble carbonates.

CONTROL OF UNDESIRABLE SPECIES

BIOLOGICAL CONTROL
The virus, cyanophage LPP-l, and several other phycoviruses (e.g., LPP-2,
SM-I, N-I and AS-l) infect various blue-green algae and could conceivably
be employed to control their growth in surface water since they cause lysis
of the sensitive forms. Recently, the eucaryotic algae Chlorella and
Sirodotia have also been reported to be attacked by phycoviruses. Addition
of virus particles to cultures of susceptible algae results in considerable
increase in virus titres and rapid fall in algal cell numbers (see Safferman
1973).
A better method for algal control consists in introducing suitable
crustaceans or fish fingerlings into the water body. These animals feed on
algae either directly or indirectly, and may be harvested for food at maturity.

CHEMICAL CONTROL
The growth of algae in reservoirs and lakes is controlled by appropriate
applications of algicide cop per sulphate wh ich selectively kills the algae
within a dose range of 0.25-9.5 ppm. The effective dose varies with the
chemical character of water as weil as the concentration of algae. Chloro-
phenyl dimethyl urea, a photosynthetic inhibitor, has also been found effec-
tive in checking the growth of planktonic algae. Besides, there are other
promising algicides such as antibiotics, quinones, substituted hydrocarbons
and phenols. But they are not in common use because of their higher cost.
The quinone 2,3-dichloro-l,4-naphthoquinone is selectively toxic to
48 A TEXTBOOK ON ALGAE

many bloom forming cyanophytes as compared to other algae. This can

be effectively used to control water blooms caused by bIue-green algae.
A particularly suitable algicide in widespread use is Dichlorophen
("Panacide" of British Drug Houses Ltd.). It is inexpensive, potent and
non-toxic to animals and human beings, and can be used for preventing
algal growth in open air swimming pools.
The Metropolitan Water Board of London has found that artificial
circulation of water in reservoirs is an effective method of reducing growth
of planktonic algae. The basis underlying this method is not clear but the
breakdown of chemical stratification in water appears to be significant
(G. E. Fogg, personal communication).

TEST QUEST/ONS

1. Why is Chlorella preferred as a suitable substitute for conventional foods '1

2. What purpose do the bamboo net beds serve in Porphyra farming '1
3. The utilization of agar, alginate and carrageenin as food is restricted. Why'l
4. How do nitrogen-fixing blue-green algae serve to reclaim 'usar' lands '1
S. What factors determine a good manure '1 Suggest reasons for the superiority of
algal manure over farmyard manure.
6. Do you think that growth of algae in water reservoirs, and water treatment and
distribution systems, is to some extent helpful in improvina the pot ability ofwatcr '1
If so, give reasons.
7. Enumerate thc preventive measures that can be adopted to control the growth of
algae in water reservoirs and water supply systems.
8. What role can algae play in a food production unit for space travel so that most of
the food used by the space travellers is produced in the space vehicle itself rather
than being carried from earth '1

SELECTED REFERENCES

Dawson, R. C., Potential for increasing food production through microbiology, Bact.
Rev., 19, 251-66 (1965).
Safferman, R. S., Phycoviruses. In Carr, N. G. and Whitton, B. A. (eds.) The Biology
of Blue-green Algae, 214-37, Blackwell, Oxford (1973).
Singh, R. N., Role of Blue-green Algae in Nitrogen Economy of Indian Agriculture,
Indian Council of Agricultural Research, New Delhi (1961).
Stewart, W. D. P., Nitrogen fixing plants, Science, Wash., 158, 1426-32 (1967).
Tamiya, H., Role of algae as food. In Proc. Sympos. Algology, 379-89, Indian Coun-
eil of Agricultural Research, New Delhi (1960).
Thivy, F., Seaweed utilization in India. In Proc. Sympos. Algology, 345-65, Indian
Council of Agricultural Research, New Delhi (1960).
 PART II

STRUCTURE AND REPRODUCTION

OF
SELECTED ALGAE
 5
Cyanophyta
The Phylum Cyanophyta (Myxophyceae, blue-green algae) differs from
other algae in having a procaryotic cell organization, i.e., it lacks organized
double membraned nuclei, chromatophores and mitochondria, and possesses
characteristic photosynthetic pigments including biliproteins, myxoxanthin
and myxoxanthophyll in addition to chlorophyll-a and ß-carotene. No
flagellated cells are formed and movement of motile stages is brought about
by a characteristic gliding action. However, certain specialized cells known
as heterocysts are produced in many species. True sexuality, defined as
aIternating karyogamy and meiosis, does not occur but genetic recombina-
tion which fulfils the function of sex is known in some members.

OCCURRENCE AND ECOLOGICAL AMPLITUDE

The blue-green algae are perhaps the most widely distributed algal organisms,
especially in tropical countries. They are ubiquitous and cosmopolitan,
found in clean and poilu ted waters of lakes, ponds and reservoirs, in fresh
or salt waters, and in stagnant or flowing waters. They occur in the spray
zone of seacoasts as weil as in surface water of oceans, and are widespread
on or in tropical soils of diverse kinds. One of the most characteristic and
favourite habitats is the tropical paddy field. Other conspicuous places
include tree barks, rocks and stones, footpaths and plant pots.
The blue-green algae form a significant part of the free-floating popula-
tion of microscopic organisms wh ich constitute the plankton of eutrophie
(organically rich) lakes and oceans. They are also found in the benthos of
lakes and ponds and are largely responsible for the high fertilizing qualities of
bottom muds of dried ponds and tanks. In tropical countries the ponds that
are heavily polluted and exposed to high temperatures and intense sunlight,
provide optimum conditions for the growth of blue-green algae. Species of
Microcystis and certain other planktonic blue-greens multiply so prolifically
in such conditions that they impart bluish-green colour to the pond water
which appears as "pea soup". Such water blooms in tropical ponds are
more or less perennial but similar blooms, though ephemeral, are also form-
ed in temperate climates either sporadically or annuaIly.
Except for certain bacteria, the blue-green algae are the only organisms
which grow in adverse and harsh environments. While most of them thrive
in biologically normal habitats, so me can successfully end ure extremes of
temperature, light intensity, desiccation, and high or low concentrations of
dissolved nutrients. Species of Mastigocladus and Phormidium are impor-
52 A TEXTBOOK ON ALGAE

tant members of the flora of hot water springs and can tolerate temperatures
as high as 75°C. On the other hand, species of Phormidium are found in
the frigid arctic lakes and certain others,collected from the Antarctic, have
experimentally been shown to survive freezing. Tolerance to extremes of
light intensity is i1Iustrated by species found in caves which are virtually
dark, on bare rocks exposed to intense sunlight, or those growing in tropi-
cal deserts. Certain species of Nos/oe and Chlorogloea have remained viable
after prolonged storage in a calcium chloride desiccator. Likewise, some
blue-greens thrive in waters of highly polluted ponds or of non-polluted
mountain streams.
T4e Cyanophyceae are pioneer colonizers of bare rocks or virgin lands
and grow luxuriantly on alkaline or saline soils. Several species grow in or
on calcareous strata. Some have been found in oil field sump ponds and
in sam pIes of floating tarry scums. A few species of Nos/oe and Anabaena
grow as endopbytes in the roots of cycads, in Azolla and in An/hoeeros. A
number of blue-greens are the algal constituents of various lichens. Others
live symbiotically in diverse unicellular animals. In nature, they interact
with and affect the behaviour of a wide variety of diverse procaryotic and
eucaryotic organisms.

THALLUS ORGANIZATION AND RANGE OF FORMS

Unicellular, colonial (palmelloid), filamentous and heterotrichous forms are

(ommon but more elaborate macroscopic thalli are rare. The products of
cell division, instead of being liberated as distinct individuals, often remain
united in an assemblage of cells (a colony). Such colonies may be definite
(e.g., Merismopedia) or irregular and indefinite (e.g., Aphanolheee). The
actual shape of a colony is largely determincd by the number of planes in
which the cells divide.
A filamentous blue-green alga consists of a number of cells arranged
end to end. Simple, unbranched, uniseriate filaments are characteristic of
Lyngbya, Anabaena and many other genera of the Nostocales. Here, the
term filament includes the row of cells as weil as the surrounding gelatinous
sheath; if it consists of cells without a distinct sheath, it is designated tri-
chome. The mucilagc sheath is also found around unicellular and colonial
forms. Most filaments havc a single trichome each, but in some more than
one trichome are so disposed within the common sheath that the filament
appears to be branched. This phenomenon is known as pseudo or false
branching and is characteristic ofsuch genera as Sey/onema and Tolypo/hrix.
It is caused cither by the growth of free ends of some trichomes through
the sheath of the parent filament, or by thc degeneration of the top cell in
a loop, leading to tbe division of an originally single and continuous
trichome into two independent pseudobrancbes. In some genera, true
(lateral) branching occurs.
 CYA~OPHYTA 53

CELL STRUCTURE

The most striking difference in the cell structure of blue-green and other
algae is the lack of differentiation of parts in the former. There are no
organized nuclei, chromatophores, pyrenoids, mitochondria, or true vacuoles.
The only prominent structures seen within the blue-green algal cell under
the light microscope are the different kinds of granules whose number and
size vary in different genera and depend on physiological and nutrition al
factors. These are proteinaceous and are callcd cyanophycin granulcs.
Reserve food is also stored in the form of a carbohydrate known as
cyanophycean starch (=glycogen). Other storage products are trehalose
or sulpholipid.
In addition to the cyanophycin granules, the healthy cells ofmany plank-
tonic blue-green algae contain numerous small bodies of irregular shapes,
known as pseudovacuo)es or gas vacuo)es. These appear black under low
power and reddish under high power of a microscope. Under an electron
microscope the gas vacuoles appear to consist of packed arrays of cylindrical
vesiclcs, each with conical ends and bound by a single membrane. Fogg's
(1970) study on the biology of gas vacuoles proves that they contain meta-
bolie gases and therefore serve as a mechanism for buoyancy regulation.
The gas vacuoJes are produced in low light intensity and disappear in high
light intensity. In bright light there is greater production of sugars through
photosynthesis resulting in an increase in osmotic press ure ofthe cells which
causes the collapse of gas vacuoles. The ecological significance ofthis find-
ing is that the blue-green algae growing below the water surface receive less
light and form gas vacuoles, thus increasing their buoyancy. Such cells
then rise to the surface where they receive more light, develop higher cellular
osmotic press ure which leads to the collapse of gas vacuoles. This results in
reduction of buoyancy and consequently the cells sink to deeper layers of
water.
The core or central region of a blue-green aJgal cell appears somewhat
transparent and contains most of the genetic material. There are no nuclei
or nucleoli. Since DNA is not associated with protein material (histones or
protamines), no organized or firm chromosomes, characteristic of higher
plants and animals, are found in blue-green algae. Besides DNA, RNA is
present.
The cells possess a wall which may be differentiated into an inner and
an outer layer. Outside the cell wall there is a gelatinous sheath which is
either homogeneous and colourless or thick, stratified and pigmented. Such
a sheath is characteristic of those blue-green algae which grow in dry or
xerophytic habitats. A firm, pigmented sheath is indeed an asset to the
terrestrial or subaerial forms because it enables them to perennate. This is
due to the water-absorbing and water-retaining capacity ofthe sheath. The
cell wall contains muramic acid and other components not found in other
algae.
S4 A TEXTBOOK ON ALGAE

MICROMORPHOLOGY

Since the cells or filaments are spherical or cylindrical rather than flat, the
structure of deep-seated inner regions constituting the core of the sphere
cannot usually be obscrved by merely mounting the alga on a slide and
examining it under the microscope. Only the outcr or peripheral parts can be
observed in this way. For examining the inner or central regions, sections
must be cut. UItrathin sections of a typical cyanophycean cell studied
under an electron microscope at magnifications of about 25000 times reveal
manY.important details (pan kratz and Bowen, 1963) which are diagram-
matically represented in Fig. 2-1 A (sec also the Frontispiece), and are des-
cribed here.

SHEATH AND CELL WALL

The gelatinous sheath consists of three layers of microfibrils disposed reti-

culately within an amorphous matrix. The chemical nature of these micro-
fibrils is not known but recent studies indicate that cellulose is absent. In-
side the sheath is a double-Iayered cell wall whose inner layer contains
mucopeptide and muramic acid.
The cytoplasmic membrane is found inside thc cell wall and generally
consists of two electron opaque layers separated by a translucent layer.

LAMELLAE

The peripheral regions of the cell are traversed by a large number of photo-
synthetic lamellae or thylakoids which may be arranged regularly in two or
more parallel stacks, or disposed irregularly. A lamella appears as an
elongated, flattened sac and consists oftwo unit membranes, each of about
7S A thickness.

GRANULES

The various kinds of granules observed in different cyanophycean cells

include the cyanophycin granules, polyhedral bodies, ribosomes, a.-granules
and ß-granules. Their actual significance and function are not c1early under-
stood but someofthem, e.g., ribosomes, areconcerned with protein synthesis,
whereas others, e.g., cyanophycin granules, are thought to be of the nature
of reserve food. In planktonic forms pseudovacuoles are found; their
uItrastructure has been described earlier.

CENTRAL REGION OF THE CELL

The genetic material, consisting of DNA fibrils, is concentrated in the centre

of the cell and this region is also traversed by a few thylakoids.
 CYANOPHYTA SS

HETEROCYSTS

These are specialized cells which differ from ordinary vegetative cells in
having a thickened wall and one or two pores. Usually they are slightly larger
than the vegetative cells, and under the light microscope appear as empty,
homogeneous, double-walled and markedly distinctive cells. A recent study
of heterocyst development with the electron microscope has shown that the
general sequence of events during the differentiation of a vegetative cell
into heterocyst includes cell enlargement, synthesis of a many-Iayered,
rion-cellulosic polysaccharide wall, gradualloss of photosynthetic pigments,
decrease in granular incIusions, and reorientation of the photosynthetic
lamellae into a complex reticulate pattern (see Lang 1968). The heterocyst
lacks the normal lipid components of the photosynthetic lamellae present
in the vegetative cells or chloroplasts of algae and higher plants, but contains
two novel types of Iipids-glyeolipid and aeyl lipid-not found in vegeta-
tive cells. Besides, the mature heterocysts contain chlorophylJ-a but lack the
accessory pigment, phycocyanin, which is normally required for the opera-
tion of the Photosystem-II of photosynthesis or in Hili reaction.
Heterocysts are found in many but not all filamentous blue-green algae
belonging to the orders Nostocales and Stigonematales. They may occur
either terminally or in an intercalary position in the trichome.
Many views have been advanced about the function of the heterocyst.
An earlier belief that heterocysts are moribund or dead cells can be safely
dismissed because in exceptional cases they have germinated to produce new
filaments or endospores indicating their spore-like nature. The changes in
structure of the cell wall and formation of endospores within some hetero-
cysts, however, do not seem to support this belief. Their potentiality to
germinate under certain conditions, led some workers to regard them as
archaic reproductive bodies wh ich have now become almost functionless.
though sometimes they revert to their original reproductive function. Un-
'usually frequent germination of heterocysts in certain strains of Gloeotrichia
ghosei has been reported by Singh and Tiwari (1970).
The heterocyst has also been assigned a role in sporulation (Wolk, 1966)
and nitrogen fixation (Fay et al., 1968). Evidences suggesting its role in
the sporulation of Anabaena cylindrica are: (I) the vegetative cells adjacent
or proximal to a heterocyst sporulate earlier than those distal to it and
(2) the removal of the heterocysts from the filaments prevents sporulation.
According to Kulasooriya et al. (1972), the carbon: nitrogen balance is the
major factor which induces and controls the differentiation of a vegetative
.cell into a heterocyst.
Researches during the past three years have conclusively established that
the heterocyst is the site of nitrogen fixation in most blue-green algae.
Evidences in support of this conclusion are that: (I) virtually all the reported
nitrogen-fixing blue-green algae are heterocystous forms; (2) combined in-
organic nitrogen inhibits both heterocyst differentiation and formation of
the enzyme nitrogenase; (3) the nitrogen ase requires an an aerobic atmos-
56 A TEXTBOOK. ON ALGAE

phere for its nitrogen-fixing activity and the heterocyst has just such an
atmosphere; the lack of phycocyanin, the inability to photoassimilate C02,
and the high respiratory activity of the heterocysts measured in terms of
oxygen uptake and thc T.T.C. (Triphenyl Tetrazolium Chloride) reaction,
indicate that they have a highly reducing atmosphere; and (4) the localiza-
tion of active nitrogen ase within the heterocysts is indicated by the fact
that isolated heterocysts (free of vegetative cells) can reduce acetylene to
ethylene when supplied with a reducing agent (sodium dithionite) and an
energy source, ATP (adenosine triphosphate).
In vitro studies of Smith and Evans (1970) on Anabaena cylindrica
indicate that nitrogenase is also present in vegetative cells. The report that
Gloeocapsa, a colonial, non-heterocystous blue-green al ga also fixes nitrogen
seems to raise the possibility of nitrogen ase activity in vegetative cells of
non-heterocystous families as weil (see Wyatt and Silvey 1969).
Two other hypotheses concerning the functions of heterocyst are that:
(I) they help in splitting" a trichome into hormogonia; and (2) they are
enzyme receptacIes. '
The varied types of functions attributed to heterocysts suggest that
probably they function differently under diverse conditions. It is also Iikely
that their function involves interaction with adjacent cells. With the elec-
tron microscope connections between heterocysts and vegetative cells, and
between pairs of vegetative cells, have been demonstrated in some blue-
green algae.

COMPARATIVE ORGANIZATION OF VEGETATIVE CEllS.

HETEROCYSTS AND SPORES

Reeent cytological and biochemical studies reveal that these cell types differ
in structure, composition and function (Lang, 1968). Both heterocysts and
spores develop from vegetative cells. As compared to vegetative cells, the
wall of heterocyst includes an additionallayer lacking true cellulose, whereas
that of spore consists of several additional sheath layers with the outermost
1 or 2 tayers often being variously sculptured. The vegetative cells and
spores have nearly the same type of thylakoid arrangement. The heterocyst
development, however, is accompanied by extensive reorganization of
thylakoids in a reticular fashion at its two poles. In addition, heterocysts
contain osmiophiIic polar nodules. The number of various granules is
target in spores and smaller in heterocysts than in vegetative cells. Poly-
phosphate granules, which function both as an energy source and inorganic
phosphate reserves, are found in vegetative cells as weil as spores but not
in heterocysts. The lipid composition of the three ceU types differs (see
under Heterocysts). The vegetative cells contain a variety of photosynthetic
pigments, e.g., chlorophylI-a, phycocyanin, ß-carotene, myxoxanthophyll
and other carotenoids. Heterocysts have only chlorophyll-a and ß-carotene.
In spores no other pigments except carotenoids are found. The distribu-
tion of pigments in these cell types is highly significant in their metabolie
 CYANOPHYTA 57

activities.
Thc vegetative cells carry out the aerobic type of photosynthesis
charactcristic of other algae and generate both the rcducing power, NADPH
(reduccd Nieotinamide Adenine Dinucleotide Phosphate) and the energy
source (ATP). The heterocysts include only Photosystem-I and therefore
can only have the an aerobic type of photosynthesis as fouod in photo-
synthctic bactcria. The spores are photosynthetically defunct. The nitro-
genase activity is confined to heterocysts and not to spores.

PIGMENTS AND CHROMATIC ADAPTATION

The chief pigments are chlorophyll-a, ß-carotene, myxoxanthophyll, myxo-
xanthin (echinenone) and c-phycocyanin. These are not locatcd in any chro-
matophore but are found in photosynthetic lamellae (thylakoids). Smallcr
amounts of c-phycoerythrin, oscilloxanthin, zeaxanthin and lutein mayaIso
be found. However, their proportion, especially of phycocyanin (blue) and
phycoerythrin (red), varies greatly under different habitat conditions. Forms
whieh are exposed to bright sunlight are rieh in phycocyanin and appear
deep bluish-green; those growing in low light intensity generally have more
of phycoerythrin so that they appear reddish. The filamentous alga Tri-
chodesmium erythraeum which grows in the plankton of the Red Sea is an
extreme example where the phycoerythrin content is so high, and phycocya-
nin so low, that it appears distinctly reddish even though it is actually a
blue-green.
In contrast to other pigments wh ich are fat-soluble, the phycocyanin
and phycoerythrin, collectively known as biliproteins or phycobilins, can be
extracted in water after cell disruption and are proteinaceous.
In a few species of Cyanophyta, the pigmentation seems to be determined
by the quality of light in which they are grown. For instance, some species
of Oscillatoria can assurne a green colour in red light, reddish in green light,
and bluish-green in yellow light. Such a capacity to change colour com-
plementary to that ofthe light is known as chromatic adaptation or Gaidukov
phenomenon. This adaptation is of definite advantage since it enables the
al ga to maximally absorb the available light for photosynthesis.
Scheibe (1972) has isolated a new phytochrome-like photoreversible pig-
ment from Tolypotlzrix tenuis. The absorption maxima for the two photo-
reversible forms of this pigment lie in the green and red portions of the
spectrum instead of in red and far-red (red and far-red are characteristic of
the higher plant phytochrome). This new pigment may possibly control
in cyanophytes several morphogenetic phenomena, including chromatic
adaptation.

MOVEMENTS
Although motile ftagellate cells are absent, mature individuals of many
Oscillatoriaceae and hormogonia of most filamentous forms are capable of
58 A TEXTBOOK ON ALGAE

movement. A pendulum-Iike, osciIIatory, swinging movement of the tri-

chomes can be readily observed when Iiving material of Oscillatoria is
examined under a microseope. In different genera, the movement may be a
backward and a forward gliding, a slow waving of one end of the trichome,
or a spiral progression or retrogression. Some trichomes rotate around their
axes, simultaneously gliding forward or backward. The gliding movement
differs from swimming in that it occurs in contact with some solid or semi-
solid substratum, but without any visible organ or a visible change in the
shape of thc alga.
Thc precise mechanism of the gliding movement is obscure. Many
theorics have becn advanced but without conclusive experimental support.
Most workers consider that movcment is due to secrction of a mucilaginous
material through minute pores in the cell wall. That the gliding movement
is shown only by those species of Oscillatoria which produce mucilage,
supports this viewpoint. Another hypothesis claims that the movement is
due to rhythmic waves of alternate expansions and contractions along the
length of the trichome. A third view has recently been proposed by Halfen
and Castenholz (1970). According to them the gliding movement may be
the result of waves travelling in one direction along the cell surface acting
against so me solid substrate or an associated elastic sheath. Since there are
no flagella or muscular threads of any kind, the basic mechanism of how a
filament glides or oscillates back and forth is unknown. It may possibly be
associated, as in other movements, with phosphorylation and A TP.

NITROGEN FIXATION

Certain blue-green algae possess the capacity to convert (or fix) elementary
nitrogen found in the air to useful nitrogenous compounds which can be
assimilated. This property makes them truly autotrophie plants in the sense
that they can fix not only atmospheric CO2 but also gaseous nitrogen.
Like heterocysts, the nitrogen-fixing power is restricted to certain fila-
mentous species such as Au/osira, To/ypotlrrix, Anabaena, Cylindrospermum,
Nostoc and Mastigoc/adus. Such species contribute greatly to the fertility
of rice fields in tropical countries. In temperate regions also nitrogen
fixation seems to be mainly a cyanophycean process rather than bacterial.
A number of blue-green algae are found in symbiotic association with other
plants and the nitrogen fixed by them may be largely responsible for the
success of the partnership.
In the past decade significant advances have been made in elucidating the
biochemistry and mechanism of nitrogen fixation. By far the most important
of these researches has been carried out in the laboratory ofProfessor R. H.
Burris at the University of Wisconsin. Prior to 1960, studies on the
mechanism of this process had practically come to a standstiII because no
methods were known for extracting nitrogen-fixing enzymes from the
organisms. Its successful demonstration in 1960 in cell-free extracts from
nitrogen-fixing blue-green algae (and also bacteria) facilitated the biochemical
 CY ANOPHYTA 59

study. A second interesting discovery by the Wisconsin group is that of

acetylene reduction by nitrogen-fixing cell-free extracts. It was further
established that the reduction of acetylene to ethylene is brought about by
the enzyme nitrogenase whose primary function is to reduce nitrogen. The
importance of this reduction system is so great that it is now being envisaged
and e1aimed as an effective method of scanning blue-greens or other
organisms for nitrogen fixing capacity.
The nitrogen fixed by blue-green algae can be assimiJated by themselves
as weil as by other organisms. Substantial amounts of holuble nitrogenous
compounds are liberated from healthy cells of nitrogen-fixing bIlle-green
algae either into the culture medium or in their natural babitats. Besides,
wben the algae die, their nitrogenous materials decay and form ammonia
wb ich may tben be transformed into nitrates by nitrifying bacteria thus
enriehing the fertility of the habitat.

NUTRITION

Cyanophytes bave a greatly restricted range of their nutrition and in

this respect tbey differ sharply from most other proearyotes. Witb tbe
exception of some strains of Nostoc, To/ypothrix and Ch/orogloea, most
of them are obligate photoautotrophs and cannot grow in darkness even
in tbe presence of organic substrates. The biochemical basis of their
obligate pbototropby is still not e1ear._ Two bypotheses bave recently
been put forward to explain this phototrophy. One suggests that since the
key respiratory enzymes, cx-ketoglutarate dehydrogenase and nicotinamide
adenine nucleotide oxidase (NAD), are absent from blue-green algae, the
incomplete functioning of the tricarboxylic acid cyele might be responsible
for their phototrophic bebaviour (Hoare et al., 1970). Tbe second hypothesis
implicates tbe lack of enzyme repression control mechanism in the tricar-
boxylic acid cyele and amino acid metabolism as the principal causative
feature for the obligate phototrophy (Carr, 1970). The enzyme repression
control operates at the transcription (cistron) level, and aets by stopping
synthesis of biosynthetic enzymes in response to the exogenous availability,
or over production of their end products. The inhibition of enzyme syn-
thesis is termed repression and its release derepression.

REPRODUCTION, GENETICS AND LlFE CYCLE

In many unicellular and colonial forms, the chief method of reproduction is

by division of a parent cell into two daughter cells, or by fragmentation of
a colony into two or more colonies. Sometimes, in addition to cell division
or fission, non-motilc spores (endospores or exospores) are produced; each
of these gives rise to a new plant. The spores are commonly produced in
ordinary vegetative cells rat her than in differentiated sporangia. Unlike
eucaryotic algae, mitosis and meiosis are not involved in cell division or
sporogenesis.
60 A TEXTBOOK ON ALGAE

Filamentous forms belonging to the Nostoca]es and Stigonematales regu-

]arly multiply by the breaking of their trichomes into hormogonia which are
generally motile and ]ater mature into filaments. Some forms produce
spores or akinetes which function as reproductive or perennating cells.
True sexuality does not exist but a kind of parasexual phenomenon,
known as genetic recombination, first reported in Anacystis nidulans by
Kumar in ]962, has since been confirmed by other workers (Bazin, 1968).
Genetic transformation (a mode of gene transfer ]eading to genetic recombi-
nation) for antibiotic resistance characters has been demonstrated in
A. Nidulans (Shestakov and Khyen, 1970). Genetic recombination differs
from true sexuality in that it is not attended by syngamy or meiosis, and
yet the function of true sexuality (namely, to bring about gene recombina-
tion) is achieved. Besides Anacystis nidulans, genetic recombination has
been reported in Cylindrospermum majus (Singh and Sinha, 1965) and
Anabaena doliolum.
Lazaroff and Vishniac (I ?62) claim that under certain conditions different
filaments of Nostoc muscorum can actually fuse in a manner reminiscent of
heterokaryotic phenomenon in certain fungi. According to them the al ga
passes through a life cycle involving alternation between a heterocystous
and a sporogenous phase (Fig. 5-1). The former occurs in the presence of
light and the latter when the alga is grown heterotrophically in the dark.
It appears that the fusion of short non-heterocystous filaments from the
dark-grown cultures, wh ich takes place after exposure to light, initiates the
heterocystous phase of development.
Nothing is known about the genetic systems involved in recombination
in the Cyanophyta. However, it is Iikely that recombination is brought
about by conjugation between donor and recipient cells, as in bacteria.
Recent studies demonstrating the susceptibility of some blue-green algae to
a virus (known as cyanophage, phycovirus or blue-green algal virus) warrant
the possibility that gene recombination mayaIso be caused by transduction,
a process in which the virus (as prophage) acts as a vector of certain genes.
transferring them from donor to recipient cells.
. The blue-green algal virus, cyanophage LPP-I, was discovered by
Safferman and Morris in ] 963. It was so named as to indicate its host-
range (Lyngbya, Plectonema and Phormidium). Since then many more
viruses attacking diverse blue-green algae have been reported (Safferman,
1973), and can now be broadly c1assified into the following categories:
(I) LPP-I viruses; (2) LPP-2 viruses, which are generally similar to LPP-I
viruses morphologically but different serologically; (3) SM-I viruses (on
hosts Synechococcus and Microcystis), which are polyhedral and have much
shorter tails than those of LPP viruses; (4) N-I virus (on host Nostoc),
with general structure similar to LPP-I but with contractile tail; (5) AS-I
(on hosts Anacystis and SynecllOcoccUS), largest phycovirus described so far
and morphologically similar to LPP-I but with particle size (head diameter
90 nm and taillength about 245 nm) much targer than LPP-I, and contrac-
tile tail like N-]; and (6) miscellaneous phycoviruses, infecting certain
 CY ANOPHYTA 61

species of Cylindrospermum, Anabaenopsis, Raphidiopsis, or Anabaena.

Although the nucJeic acid and protein components of some ofthe above
phycoviruses have not been analyzed in detail, available information
(mostly on LPP-J) suggcsts that cyanophagcs contain double-stranded DNA.

CS:X:X::;O--+3 ~ ---.~

2~ ~4
A tenta1ive scheine
for the alternation
01 sporogenous and
heterocystous gene-
rations in Nostoc
muscorum
1

r
Heterocystous phase

1
OCcx::O cP 5
(1HamentfUslOn~ ~
() 7
,requlres light 'tS=
6 (
I ~
spore fbrmation requires light
ccb~~ ~....
f cf'
(,\ 111
°~
IV . (9::>
0 --+ V (9::0 --"'" I
Sporogenous phase
~
~O~~/
Fig. 5-1. Nostoc muscorum, hfe cyc1e and alternation of heterocystous and sporogenous
generations. (After Lazaroff and Vishniac, 1962.)

A study of the replication cycle of LPP-I in Plectonema boryanum

revealed the following stages: (1) attachment of virus to host cell wall hy
means of the tail; (2) injection of viral DNA into the algal cell; (3) replica-
tion of viral DNA in the nucleoplasm of host cell and hreakdown of host
cell DNA; (4) migration of replicating viral DNA from nucleoplasm to
photosynthetic thylakoids, Icading to slight displacement of the latter; and
(5) synthesis of protein coats and their enclosure around DNA particles,
leading to virus assemblage.
Rccent researchcs have also provided a strong indication of the existcncc
of tcmperate cyanophagcs similar in properties to the weIl known bacterio-
phages. Cannon et al. (1971) have discovered a new strain of LPP-l which
successfully lysogenized the host alga, P. boryanum. In this system, the
majority of the host cells were, of course, Iysed hut a few ce]]s resisted lysis
62 A TEXTBOOK ON ALGAE

and under.wcnt Iysogeny (i.e., they survived and dividcd in step with the
cyanophage DNA harboured internally).

TAXONOMY AND CLASSIFICATION

The taxonomy of the blue-green algae has long been in a highly confused
state. Enormous number of new species and genera have bcen created by
different workers on trivial grounds, and borderline forms are extremely
difficult to classify. Two biotic characteristics have contributed a great
deal to the present state of confusion. One is the way in wh ich various
morphological features vary with differences in ecological conditions. The
second is the variability of physiological and biochemical response observed
under different conditions. This may merely bc the physiological counter-
part of thc morphological variability or may represcnt gcnctically detcr-
mincd responses.
A few attempts have becn made to produce a rational and orderly system
of c1assification or taxonomic delimitation by reducing the number of genera
and spccies but these have either been drastic or confined to a study of only
one or two families rather than the entire phylum. A satisfactory solution
can be found only after a combined approach involving a thorough study of
the physiology, cytology and genetics of different taxa has been made.
This secms a distant goal.
In this book the c1assification system of Fritsch (1945) has been followed.
He divides the blue-green algae into five orders:
I. Chroococcales. Uniccllular or colonial (palmelloid) forms; rcproduc-
tion by ccll division and by endospore formation.
11. Chamaesiphona1cs. Uniccllular or colonial lithophytes or epi-
phytes exhibiting marked polarity; reproduction by endospores or
exospores.
111. Pleurocapsales. Heterotrichous filamentous forms lacking hetero-
cysts; reproduction by endospores; hormogonia absent.
IV. Nostocales. Non-heterotrichous filamentous forms often showing false
branching; heterocysts commonly present; reproduction mostly by
hormogonia and akinetes.
V. Stigonematales. Heterotrichous filamentous forms, mostly with hetero-
cysts, showing true branching and usually pit connections between
adjacent cells; multiplication mostly by hormogonia and rarely by
hormocysts and akinetes.

PHYLOGENETIC RELATIONSHIPS

The blue-greens are the most primitive of algal organisms. In their general
cytology, genetics and morphology as weil as procaryotic nature they seem
c10scly related to bactcria but differ from them in their photosynthetic
metabolism and lack of ftagellated cells. If the view that chloroplasts are
cellular symbionts is accepted, the relationship to be considered is not that
 CYANOPHYTA 63

between the procaryotic blue-green algae and the eucaryotic algae but
between the former and the chloroplasts of the latter. Thus blue-green
algae as a whole can be compared only with the chloroplasts of Rhodophyta
or Cryptophyta. Some affinity with red algae is indicated by the presence
of phycoerythrin and pit connections, and lack of flagellated structures.
Further, the blue-green and red algae have a common pattern of fatty acid
formation which differs from other plants in that the lipid content does not
increase as the thallus grows. In these algae, nitrogen starvation is also
not a factor in fatty acid accumulation (see Klein and Cronquist 1967).
Phycobilins link Cryptophyta to blue-green algae.
The resemblance between blue-green algae and bacteria appears stronger
and more fundamental than between bIue-green and red algae. The chemical
composition and ultrastructure oftheir cell wall and cytoplasm, the organiza-
tion of photosynthetic apparatus, and aspects of hydrogen, nitrogen and
sulphur metabolism link the blue-green algae with the rest of the Schizo-
mycophyta via the photosynthetic bacteria. Clearly, the bille-green algae
and the bacteria are two grollps of the same Kingdom, Procaryota (Klein
and Cronquist, 1967).

Order: CHROOCOCCAlES
Family: Chroococcaceae
Genus: Gloeocapsa

This is a terrestrial or subaerial alga generally found on rocks, walls or soils.

It is a palmelloid, colonial form. Each colony consists of a few cells (Fig.
5-2 A, B) which are more or less spherical and have conspicuous, coloured,
stratified and concentrically lamellate gelatinous sheaths around them. The

Fig. 5-2. A, Gloeocapsa magma; B, G. janlhil/u.

sheaths not only surround individuai cells but also the entire colony. In
most species the photosynthetic thylakoids are arranged in rows at right
angles to tbe plasma membrane, witb tbe outer tbylakoids being directly
connected with plasma membrane; but in G. a/picola they lie parallel to
longitudinal walls. Reproduction occurs by simple cell division, in three
64 A TEXTßOOK ON ALGAE

planes, at right angles to one another, so that more or less spherical daughter
cells are produced.

Order: NOSTOCALES
Family: Oscillatoriaceae
Genus: Oscillatoria

It is common and widespread in polluted waters, temporary rain\Vater pools,

drains and streams. O. rubescens forms dense deep-water blooms in tem-
pe rate lakes and ponds and its migration and colonization in deep water
lakes is often associated with high level of eutrophication. Also characteristic
of highly eutrophie waters are the populations of certain species having very
narrow trichomes (fess than 3 I-'m in diameter). Some species are marine.
In the living state many species exhibit a characteristic swinging osciIlatory
movement of their trichomes. In streams, so lang as the trichomes continue
to move, they are not attacked by protozoan predators but once movement
ceases, the trichomes are invaded and devoured by protozoa.
It is an unbranched trichomatous alga. The trichome consists of a
Ilumber of simple, unditferentiated celJs, commonly broader than long (Fig.
5-3 A, B). There are no constrictions between adjacent celIs, no recognizable
base or apex but the end cell in some
species may by round, conical or dilated
with its outer surface sometimes covered
by a thickened hood known as calyptra.
The end of the trichome is occasionally
bent on one side. No heterocysts or
spores are found in Oscillatoria.
The cells exhibit a typical cyano-
phycean structure, ar.d prominent
pseudovacuoles are formed in plank-
tonic forms. Reports of a true nucIeus
or chromosomes in certain species separation disc
require confirmation. Even if these
...... ,,:
are correct, it must be borne in mind
that no one species of this genus
possesses an entire nucIeus and mitotic A B
apparatus.
Whitton and Peat (1969) have Fig. 5-3. A, Oscillatoria curviceps;
B, O. brevis.
studied the developmental variability of
O. redekei grown in unialgal cuIture. They obscrved that gas vacuoles, tri-
chome sheath amI terminal cells having pointed ends were present at certain
stages but absent at others. Their presence or absence depended upon
physiological and environmental conditions. Furthermore, at certain stages
of its development, O. rcdekei could not be distinguished from certain other
spccies of Oscillatoria.
 CYANOPHYTA 65

Reproduction is brought about by the breaking up of trichome into

hormogonia as a result of the formation of biconcave separation discs (Fig.
5-3 B) at certain points along the trichome.

Family: Nostocaceae
Genus: Nostoc

It is a freshwater or terrestrial alga, abundantly found in paddy fields.

Some species grow on moist soil intermingled with mosses and lichens
whereas others may constitute the algal components of lichens or occur as
endophytes in higher plants. N. sphaericum forms a close symbiotic associa-
tion with the phycomycetous fungus Geosiphon pyrijorme. The capacity
for nitrogen fixation has been established in bacteria-free cultures of Nostoc
sp. isolated from lichens and it has been shown that in the lichen Peltigera
aplzthosa, the nitrogen fixed by Nostoc is utilized by the fungal partner.
An interesting feature concerning the association of Nostoc with the
angiosperm Gunnera is that tbe filaments are intracellular rather than inter-
cellular, are devoid of sheath, but are enveloped by a layer of host cell
cytoplasm.
It is a filamentous form and tbe filaments are often aggregated into
ball-like gelatinous colonies which vary a great deal in size and shape, so me
being as large as a hen's egg. The filaments are usually contorted and twist-
ed in various ways. Like Oscillatoria they are always unbranched but the
cells have prominent constrictions between them. Consequently they seem
to be arranged like beads in astring (Fig. 5-4 A, B). Such ahabit is ca lied

-
~ Fig. 5-4. A, Nos/oe commune; B. N. punc/ijorme.
'"~moniliform. Unlike Oscil/atoria, the filaments regularly form heterocysts
;. and occasionally also akinetes. The heterocysts are commonly two-pored
vi and intercalary but rarely one-pored and terminal.
66 A TEXTBOOK ON ALGAE

The sheaths of adjoining trichomes may coalesce to form a common

gelatinous matrix of the colony. Cell structure is typically cyanophycean.
Reproduction takes place by hormogonia and akinetes. Lazaroff and
Vishniac (1962) have also reported the occurrence of fusions and anastomoses
between different filaments.
Based on developmental patterns, the various species of Nostoc may be
broadly cIassified into three groups: (I) the commune type, in which the
colonies have a firm pellicIe; (2) the piscina/e type, in which firm pellicIe is
absent; and (3) the punctijanue type, in which filamentous organization is
lost and aseriate form results.
The essentiality of bacteria in the formation of normal colonies of
N. sphaericum has been established by Schwabe and Mollenhauer (1967).
When grown in the absence of bacteria, only minute colonies were formed
instead of the normal, macroscopic colonies formed in bacterized cultures.

Family: Scytonemataceae
Genus: Scytonema

It occurs in fresh and saltwaters, terrestrial and subaerial habitats, grass

lawns, on tree barks and on a wide variety of other substrata.
It differs from Oscillatoria and Nostoc in having branched filaments.
The branching is false, and both single and double pseudobranches may be
formed. Typically, however, two false branches are produced remote from
two intercalary heterocysts (Fig. 5-5 A). A filament or its branches may
have a few to many heterocysts though these are usuaJly always two-pored
and intercalary. The trichomes are not moniliform and the cells are more
or less quadratic (as lang as broad). Filaments have distinct mucilage

A B

Fig. 5-5. A, Scytonema ocellatum; B, S. insigne.

sheaths (Fig. 5-5 B) which are thick, stratified and variously pigmented in
terrestrial species.
False branches arise by the degeneration of a cell in the trichome follow-
 CYANOPHYTA 67

ed by outgrowth of one or both ends of the interrupted trichome through

the parent sh.eath as branches. Alternatively, they may arise by loop forma-
tion.
Reproduction occurs by hormogonia which are commonly liberated
through the extremities of the filaments or branches.

TEST QUEST/ONS
1. Mention the primitive eharaeters of Cyanophyta whieh resemble baeteria. Are blue-
greens more elosely re la ted to other algae, or to baeteria?
2. In what respeets does the ultrastrueture of a eyanophyeean eell differ from that of
other algae ?
3. "Heterocyst is a botanical enigma", diseuss this statement. Is it still an enigma?
4. A pond develops a thiek growth of algae sueh as Plectonema, Phormidium and
Lyngbya. Can you suggest any quick method of destroying sueh a growth ?
5. Can the blue-green algae be employed to replaee ehemieal fertilizers for inereasing
the fertility of soH? Give reasons for your answer.
6. Diseuss the signifieanee of blue-green algae in relation to publie health.
7. The blue-green algae have a wide eeologieal amplitude. What struetural or physio-
logieal eharaeteristies make this possible ?
8. Compare the genetic reeombination found in Cyanophyta with the sexuality of other
algae.
9. How would you distinguish between ordinary vaeuoles, eontraetile vaeuoles and gas
vaeuoles?

SELECTEO REFERENCES
Bazin, M. J., Sexuality in a blue-green alga: Genetie reeombination in Anacystis nidulans,
Nature, Lond., 218, 282-83 (1968).
Cannon, R. E., Shane, M. S. and Bush, V. N., Lysogeny of a blue-green alga Plectonema
boryanum, Virology, 45, 149-53 (1971).
Carr, N. G., Metabolie regulation and autotrophie behaviour in blue-green algae, 1st
Internat. Sympos. TaXcIJomy and Biology of Blue-green algae, Univ. Madras
(1970).
Fay, P., Stewart, W. D. P., Walsby, A. E. and Fogg, G. E., Is the heteroeyst the site of
nitrogen-fixation in blue-green algae? Nature, Lond., 220,810-12 (1968).
Fogg, G. E., Gas-vaeuoles and their eeologieal signifieance, Ist Internat. Sympos. Taxo-
nomy and Biology of Blue-green Algae, Univ. Madras (1970).
Halfen, L. N. and Castenholz, R. W., Gliding in a blue-green alga: a possible mechanism,
Nature, Lond., 225, 1163-65 (1970).
Hoare, D. S., Hoare, S. L. and Smith, A. J., Heterotrophie potentialities of the blue-
green algae, 1st Internat. Sympos. Taxonomy and Biology of Blue-green Algae,
Univ. Madras (1970).
68 A TEXTDOOK ON ALGAE

Kulasooriya, S. A., Lang, N. J. and Fay, P., The heteroeysts of blue-areen alpe.IU.
Differentiation and nitrogenase aetivity, Proc. Ro)'. Soc., 181 B, 199-209 (1972).
Kumar, H. D., Apparent genetic reeombination in a blue-green alga, Nature, Lond., 196,
1121-22 (1962).
Lang, N., The fine struetureoe blue-green algae, A. Rev. Microbiol., %2, 15-42 (1968).
Lazaroff, N. and Vishniae, W., The participation oe filament anastomosis in the deve-
lopmental eyde oe Nostoc muscorum, a blue-green alga, J. gen. Microbiol., 28,
203-10 (1962).
Pankratz, H. S. and Bowen, C. C., Cytology oe blue-green algae. I. The cells of Symploca
muscorum, Am. J. Bot., 50, 387-99 (1963).
Safferman, R. S., Phycoviruses. In Carr, N. G. and Whitton, B. A. (eds.) The Biology
of Blue-green Aigae, 214-37, Blackwell, Oxford (1973).
Safferman, R. S. and Morris, M. E., Algal virus: isolation, Science, Wash., 140, 679-80
(I963).
Sch~ibe,J., Photoreversible pigment: occurrence in a blue-green alga, Science, Wash.,
176, 1037-39 (1972).
Schwabe, G. H. and Mollenhauer, R., Über den Einfluss der Begleitbäkerien auf das
Lägerbild von Nostoc sphaericum, Nora Hedwigia, 13, 77-80 (1967).
Shestakov, S. V. and Khyen, N. T., Evidence for genetic transformation in a blue-green
alga Anacystis nidulans, Molec. gen. Genet., 107, 372-75 (1970).
Singh, R. N. and Sinha, R., Genetic reeombination in a blue-green alga, Cylindrosper-
mum majus, Na/lire, Lond., 207, 782 (1965).
Singh, R. N. and Tiwari, D. N., Frequent heterocyst germination in the blue-greenalga
. Gloeo/richia ghosei Singh, J. Phycol., 6, 172-76 (1970).
Smith, R. V. and Evans, M. C. W., Soluble nitrogenase from vegetative eells of the blue-
green alga Anabaena cyUndrica, Na/ure, Lond., 225, 1253-54 (1970).
Whitton, B. A. and Peat, A., On Oscil/atoria redekei Van Goor., Arch. Mikrobioi., 68,
362-76 (1969).
Wolk, P., Evidence of a role of heterocysts in tbe sporulationof a blue-green alga, Am.
J. Bot., 53, 260-62 (1966).
Wyatt, J. T. and Silvey, J. K. G., Nitrogen fixation by Gloeocapsa, Science, Wash., 165,
908-11 (1969).
 6
Chlorophyta
The Phylum Chlorophyta (green algae) incJudes two cJasses of eucaryotie
algae, the Chlorophyceae and the Charophyceae. These have, as in higher
plants, the same pigments (chlorophyll-a, chlorophyll-b, various carotenes
and xanthophylls) and in tbe same proportion. Tbe pigments are located
in definite chloroplasts. Tbe excess pbotosynthates are commonly stored
in the form of starch. Inner Jayer of the cell wall is completely or partly
cellulosic. Tbe flagella, wben present, are usually two or four in number,
equal in Jengtb and of wbiplasb type.

OCCURRENCE AND ECOLOGICAL AMPLITUDE

The Chlorophyta comprise tbree main types of forms, namely, free living,
attached and parasitic, and all or some of these types may be aquatic,
amphibious, terrestrial or subaerial. Of the aquatic forms, the Ulvaceae
and Siphonales are mostly marine wbereas Conjugales and Oedogoniales
are almost entirely freshwater.
Although the green algae are predominantly freshwater forms, quite a few
species grow on or witbin the surface of moist soil, tree barks, moist and
shaded rocks and on a variety of other terrestrial and subaerial habitats.
Species of Trentepohlia constitute some of tbc best known cxamples of sub-
aerial algae whereas those of Cladophora and some members of Chaeto-
phorales grow attached on sbells of molluscs in freshwater ponds and lakes.
Some species of Chlamydomonas and Characium ftourish on crustaceans,
turtles or other aquatie animals.
Chlorochytrium lemnae is aspace parasite (endophyte) in Lemna whereas
Cephaleuros and Rhodochytrium are true parasites. Species of Trebouxia,
Chlorella and Trentepohlia form symbiotie associations with fungi in various
lichens. Some unicellula~ green algae are regularly found within theepidermal
tissues of certain lower anintals.
In contrast to Cyanophyta, Chlorophyta are mostly cold water forms
although some strains of Chlorella are known to be thermophilie. Chla-
mydomonas spp. and various chlorococcaleans oceur in such abundance on
snow that they impart to it a reddish colour. This is due to the haemato-
chrome (mostly secondary carotenoids dissolved in oil) whieh they possess.

RANGE OF VEGETATIVE STRUCTURE

The thallus exhibits a wide variation in its habit and morphology, ranging
70 A TEXTBOOK ON ALGAE

from a motile or non-motile cell through colonial, filamentous, parenchy-

matous and siphoneous habits to the highly evolved heterotrichous filament.
The highest type of specialization seems to have been attained in such genera
as Draparnaldiopsis and Chara which have a highly differentiated, compli-
cated and organized thallus. However, no elaborate and bulky parenchy-
matous or pseudoparenchymatous forms, as are met with in the Laminariales
and Fucales of the Phaeophyta, are found in green algae. Simple, pseudo-
parenchymatous thalli are, however, known in the Siphonales.

CELL WALL

Except for a few primitive ftagellate forms, the vast majority of green algae
have a distinct, faitly rigid wall around their protoplast. The cell wall
consists of two or three layers of which the inner may either be completely
cellulosic (consisting of pure glucose residues), or a mixture of cellulosic and
non-cellulosic chains. Outside the inner cellulosic layer is a layer of pectose
or mucilaginous substance. The latter is water-soluble and hence is con-
tinuously regenerated in aquatic species. The gelatinous wall layer may be
impregnated with lime in some members of the Siphonales. In Cladophora
and Oedogonium a third chitinous layer may be laid down outside the
pectose layer.

FLAGELLA

The motile vegetative ceIls and most zoospores and gametes generally have
one, two, four, or rarely many, anteriorly inserted whiplash ftagella of equal
length. These are associated with a neuromotor apparatus which connects
them with a centrosome Iying within or just outside the nucleus.

PROTOPLAST AND CELL STRUCTURE

The green algae are typically eucaryotic, having definite nuclei, chloroplasts
and mitochondria. A central vacuole is generally found in mature cells,
with the cytoplasm confined to just inside the cell wall. In some species of
Spirogyra the vacuolar sap contains purpie pigments that mask the green
colour of the chloroplasts.
Most green algae have uninucleate cells but those of Chlorococcales,
Cladophorales, Siphonales and Sphaeroplea are multinucleate (coenocytic).
The nucleus has a double-Iayered porous nuc1ear membrane, one or more
nuc1eoli and centromeric chromosomes. Well-developed complex nucleoli
occur in Conjugales and in Acetabularia.
There is a great diversity in the number, morphology and arrange-
ment of their chloroplasts, and this has been interpreted to suggest the
manifestation of evolutionary tendencies among different orders of the
phylum.
A single, cup-shaped or bowl-shaped chloroplast per cell, with one or a
 CHLOROPHYTA 71

few pyrenoids, is characteristic of the Volvocales though rarely an axile

stellate (Chlamydomonas arachne) or reticulate (Sphaerel/a lacustris) chloro-
plast may be found.
Single parietal chloroplast, usually having a single pyrenoid, is found
in the Chlorococcales but Trebouxia has single, lobed, axile chloroplast with
one pyrenoid whereas Eremosphaera has many, smalI, discoid chloroplasts
devoid of pyrenoids.
A reticulate chloroplast with several pyrenoids is characteristic of Hydro-
dictyon, Cladophora and Oedogoniales, and a parietal ring-, collar- or
girdle-sbaped chloroplast containing a few pyrenoids is found in many
Ulotrichaceae. A parietal chloroplast having a variable number ofpyrenoids
is also found in most of tbe Cbaetopborales. In more advanced mem bers
of tbis order, e.g., Draparnaldia and Draparnaldiopsis, tbe main axes of
central cells have a diminutive chloroplast restricted to the middle of the
cell.
The Conjugales have three basic and common types of chloroplasts:
(I) a flat axile plate with several pyrenoids (Mesotaenium and Mougeotia);
(2) a pair ofaxile stellate chloroplasts, each with a single pyrenoid (Zygnema);
and (3) one or a few spiral, ribbon-shaped chloroplasts, each with many
pyrenoids (Spirogyra).
Many discoid chloroplasts with or without pyrenoids are found in
Siphon ales and in Charales.
The pyrenoid of green algae commonly consists of a proteinaceous
core wh ich is surrounded by minute plates of starch. It is often highly
refractile.
The cells of the primitive, motile green algae commonly possess a pair
of contractile vacuoles near the base of the flagella. The chloroplasts of
motile cells generally also possess an eye spot near the anterior end.
The mitochondria, Golgi bodies and endoplasmic reticulum characteristic
of eucaryotic cells are also found in the cells of Chlorophyta.
In Volvocales, Oedogoniales and Ulotrichales, the excess photosynthates
are stored in the form of starch. Some Chlorococcales accumulate both
starch and lipid, whereas most Siphonales and some Conjugales store
fat.

CELL DIVISION
In the simple forms belonging to Volvocales and Chlorococcales, cell division
is linked with reproduction but in most other green algae it may be both
vegetative and reproductive.
During nucIear division all the four stages, i.e., prophase, metaphase,
anaphaseand telophase are evident. There are wide variations in the number
and morphology of chromosomes but these seem to bear little, if any, taxono-
mie correlation or significance. In general, the nucIear cytology reveals two
important features: (I) most green algae, except Conjugales, have centric
chromosomes with localized centromeres in the median, sub-median or
72 A TEXTBOOK ON ALGAE

terminal position; the Conjugales, on the other hand, have diffused eentro-
meres; and (2) the wide range of variation in the ehromosome number of
different species is mostly due to aneuploidy although a few examples of
polyploidy have also been reported.
Following nucIear division, the eell divides by a transverse furrowing of
the eytoplasm, aecompanied by the seeretion of new eell wall material.
Cytokinesis ranges from cIeavage by invagination of the plasma membrane
(as in Chlamydomonas) to the development of a distinct cell plate (as in
Fritschiella), similar to that in higher plants.

REPRODUCTION

It may be brought about by any one or all of the three methods: (I) vegeta-
tive; (2) asexual; and (3) sexual.

VEGETATIVE

This takes place by splitting of colony or fragmentation of filament. The

latter results from some extern al mechanieal pressure, septum dissolution or
from dying out or eonsumption during sexual or asexual reproduetion of one
or a few intercalary cells of the fila.ment. In Vlothrix, Oedogonium or
Sphaeroplea, the number of filaments increases by oecasional fragmentation
ofonefilament into two, each ofwhieh then grows to the adult size. The best
known example is, however, eonstituted by Stichococcusin whieh the tendeney
to fragment is so strong and pronounced that as soon as the filament grows
to the stage offour or six eeIls, fragmentation oeeurs so that filaments Ion ger
than 4-6 eells are seldom found.

ASEXUAL

The commonest method is by repeated division of the protoplasts of certain

cells. The daughter protoplasts are either transformed into motile and
flagellate, commonly naked zoospores, or they round off into non-motHe
aplanospores provided with true cell walls. The number of zoospores or
aplanospores produced within a mother cell varies from one (Oedogonium),
1\\"0, four or more (in many algae) to as many as several thousand in Hydro-
dictyon. These asexual cells may be produeed either in ordinary, unmodified
cells (Vlothrix) or in specially modified eells known as zoosporangia (Trente-
pohlia).
In many forms, zoospores are produced during night and liberated in the
morning. Their formation is most active during favourable growth periods
when the alga is young and healthy. Formation ofzoospores may sometimes
be artificially induced by modifications of environmental eonditions such as
pH, temperature, light, and transference from flowing to still water.
Zoospores are Iiberated either by gelatinization of the parent cell wall or,
more commonly, through one or more pores formed in the lateral walls.
 CHLOROPHYTA 73

When Iiberated they are generally naked and swarm around in tbis state for
a few hours or a few days. They then come to rest and attach themselves
to some substrat um by their anterior ends, lose flagella, secrete a wall, and
become quiescent for some time before growing into new individuals.

SEXUAL
Most green algae, except so me Chlorococcales, reproduce sexually. In the
Conjugales the fusing gametes are amoeboid and non-flagellate whereas in
other orders at least the male gamete is always flagellated.
The Chlorophyta exhibit wide variations in the kind and formation of
their gametes. When the fusing gametes are morphologically identical and
indistinguishable they are known as isogametes; such isogamous reproduc-
tion is noticed, among others, in species of Chlamydomonas, Ulothrix and
Zygnema. When the fusing gametes are both motile and flagellate but
unequal in size, they are termed anisogamous as found in Chlamydomonas
braun;;, Aphanochaete, and others. Oogamous reproduction involves fusion
between a small flagellate and actively motile male gamete (known as the
antherozoid) and a large, non-motile, non-flagellate and passive female
garnete, the ovum or oosphere. The ovum is generally produced singly within
a specially differentiated and enlarged cell called the oogonium, e.g.,
Oedogoniutn and Coleoclzaete. Antherozoids are, however, many and are
produced within an antheridium; in Oedogonium, nevertheless, only two
antherozoids are produced per antheridium.
In Spirogyra, the fusing gametes, which are non-flagellate, are morpholo-
gically isogamous but physiologically anisogamous since one ofthem behaves
as male and the other as female. Sirogonium provides an isolated example
of morphological anisogamy in the order Conjugales since it forms a small
male and large female gametangium.
The advancement in sexual reproduction, from isogamy through aniso-
gamy to oogamy, seems to have been accompanied by a reduction in number
and inerease in size of the female gametes. The evolution from isogamy to
oogamy perhaps oecurred independently in different orders of the green
algae. Progressive evolutionary series ilIustrating the transition from isogamy
to oogamy are met with in the coenobial Volvoeales and in certain Chaeto-
phoraceae, e.g., Stigeoc!onium, Aphanoclzaete, Clzaetonema and Coleochaete.
With the exeeption of three species (Chlorogonium oogamum, Carteria
iyengarii and Chaetonema irregulare) in wh ich the ovum is released from the
oogonium prior to fertilization, in all others fertilization takes plaee while
the ovum is still within the mother cell.
Generally the two members of a pair of fusing gametes are derived from
two different individuals, such individuals being termed heterothallic. A
number of green algae are however homothallic; in these fusion ean take
plaee betweentwo gametes whieh have been derived from the same parent
individual.
The zygote forrned as a result of isogamous or anisogamous reproduction
74 A TEXTBOOK ON ALGAE

or eonjugation is ealled zygospore, whereas that produced as a resuIt of

oogamous fertilization is termed oospore. In most haploid Chlorophyta,
the diploid zygospore or oospore seeretes a thick wall and becomes trans-
formed into a resting cell. In the next favourable season it germinates to
produee one or more plants. On the other hand, in many marine Chloro-
phyta, some Cladophorales and certain diploid forms, the zygote germinates
direetly into a vegetative plant.
Oeeasionally, some gametes, without fusion, may directly give rise to new
plants, or may form resting cells known as parthenospores wh ich subse-
qu'ently produce new plants. Such parthenogenesis occurs in Ulothrix and
certain other algae.

PERENNA TION
Certain green algae produce special resting stages that enable the alga to
pass through adverse or unfavourable environmental eonditions. Such resting
cells may be vegetative (akinetes), asexual (hypnospores) or sexual (zygo-
spores or oospores).
Akinetes are prominently differentiated and enlarged vegetative cells
wh ich beeome thiek-walled and densely laden with food reserves to tide over
adverse periods. In these the spore wall fuses with that of the parent cell
and a few additional wall layers are deposited outside the whole strueture.
Charaeteristie interealary and terminal akinetes are found in species of
Pithophora. Hypnospores are modified aplanospores that secrete specially
thiekened eell walls around them, e.g., in Sphaerella and Protosiphon.
In many Chlorophyta the only mode of perennation is by the formation
of resting zygospores or oospores.
The ordinary vegetative cells of Pleurococcus, Trebouxia, Prasiola and
eertain other terrestrial or subaerial algae can perennate as such, i.e., without
the formation of any special stages.

lIFE CYCLE AND ALTERNATION OF GENERATIONS

Four main types of life cycle-haploid, diploid, isomorphie or homologous,
and heteromorphie-are met with in the green algae.
Ulothrix, Pandorina, Coleochaete and Oedogonium, for example, are
haploid, with the zygote representing the only diploid stage in the life eycle.
In t.hese, meiosis oecurs during the first division of the zygote nucleus
(Fig. 3-2).
In the diploid forms such as many Siphon ales and some Chlorocoecales,
the mature vegetative plant is diploid and meiosis oecurs during the forma-
tion of gametes whieh represent the only haploid stage. Tbe zygote germi-
nates direct into the diploid plant.
The isomorphie life cycle involves an alternation between a haploid
gametophyte and a diploid sporophyte, both of which are morphologically
quite indistinguishable from eaeh other. The haploid phase produces
 CHLOROPHYT A 75

gametes that fuse to form the zygote wh ich in turn germinates into a diploid
sporophytic plant and produces zoospores without undergoing any meiotic
division. Meiosis oceurs during the formation of zoospores and henee
they are haploid. These grow into haploid gametophytes (Fig. 3-4). This
kind of alternation of generations is found in the Ulvaeeae, many Clado-
phorales and certain Chaetophorales, e.g., Draparnaldiopsis and Fritschiella.
Urospora, in which the two free living alternating generations are mor-
phologically dissimilar, pro vi des an example ofthe heteromorphie life eycle.

CLASSIFICATION
The use of eleetron mieroseope has eonsiderably added to our understanding
ofthe micromorphology ofgreen algae. This study in eonjunetion with their
ehemieal and physiologieal features has led many phycologists to propose
different systems of classifieation of Chlorophyta. These systems vary main-
Iy in the degree of emphasis plaeed on a partieular eombination of phyletic
eharaeters.
Emphasizing the morphologieal and reproduetive features, Fritseh (1935)
proposed a widely aeeepted system of classifieation. He divided the green
algae into nine orders (Vo!voeales, Chloroeoeeales, Ulotriehales, Cladopho-
rales, Chaetophorales, Oedogoniales, Conjugales, Siphonales and Charales).
In this book we have followed the same system of cIassifieation but have rais-
ed Charales to the rank of a Class (Charophyeeae) eoordinate with Chloro-
phyceae. Thus, the Chlorophyta ineludes two cIasses of whieh the Chloro-
phyeeae has eight orders whereas the Charophyeeae has single order, the
Charales.
Two of the more widely followed systems of c\assifieation that may be
mentioned here are those of Bold (1967) and Round (1971).
Bold, following some earlier workers, has raised Charales to the rank of a
Phylum, the Charophyta. The Division Chlorophyeophyta includes only one
Class, the Chlorophyeeae, having ten orders (Volvoeales, Chlorosphaerales,
Chlorocoecales, UJotrichales, Ulvales, Cladophorales, Siphonales, DasycJa-
dales, Oedogoniales and Zygnematales).
Round (1971) has divided the green algae (Chlorophyta sensu FrUsch)
into three phyla eontaining six c\asses and 37 orders as folIows:

I. Phy!um CHLOROPHYTA
Class Chlorophyceae-Orders Chlamydomonadales, Volvocales,
Polyblepharidales, Tetrasporales, Chlorodendrales, Chlorosarci-
nales, Chloroeoeeales, Ulotriehales, Codiolales, Ulvales, Prasio-
lales, Cylindroeapsales, Microsporales, Chaetophorales, Trente-
pohliales, Pleuroeoeeales, and Ulvellales.
Class Oedogoniophyceae-Order Oedogoniales.
Class Zygnemaphyceae-Orders Mesotaeniales, Zygnematales,Gona-
tozygales, and Desmidiales.
Class Bryopsidophyeeae-Orders Cladophorales, Sphaeropleales,
76 A TEXTBOOK ON ALGAE

Acrosiphonales, DasycIadates, SiphonocIadales, Chtorochytriales,

Derbesiates, Codiates, Caulerpates, Dichotomosiphonales, and
Phyllosiphonales.
II. Phylum PRASINOPHYTA
Class Prasinophyceae-Orders Pyramimonadales, PrasinocIadales,
and Halosphaerales.
III. Phylum CHAROPHYTA
Class Charophyceae-Order Charales.

EVOLUTIONARY TRENDS WITHIN CHLOROPHYTA

White discussing the phyletic relationships ofany taxon the terms "primitive"
and "advanced" are frequently encountered. Their meaning is relative and
can be understood with reference to two hypothetical taxa A and B; B can
be considcrcd advanced over A if its life cycle includes a stage that resembles
A. Such a generalization holds true only for organisms that are phyletically
related.
In tracing the phyletic trends within Chlorophyta, the use ofthree charac-
ters has been emphasized: (1) thallus organization, (2) sexual reproduction
and (3) life cycle.
The motile or non-motile unicell, the motile or non-motile colony, and
thc colonial, the filamentous and the parenchymatous thalli constitute an
ascending series in which the unicellular habit is considered the' most primi-
tive and the filamentous or parenchymatous the most advanced. Similarly,
in the isogamous-anisogamous-oogamous series, isogamy is considered primi-
tive whercas oogamy represents an advanced type of sexuality.
The unicellular motile forms such as Chkimydomonas are generaIly accept-
ed as the most primitive among green aIgae from which evolution has pro-
grcssed along three different lines-volvocine, tetrasporine and chlorococcine
(coccoid). The volvocine line represents aseries in wh ich the products of
the vegetative cell division are flagellated and are retained in a common
mucilaginous envelope giving rise to a motile colony. This line ends abruptly
in Volvox. The tetrasporine line paralleIs the volvocine in all aspects except
that its vegetative phase is predominantly non-motile and the motile phase is
mainly confined to the reproductive stages. The chlorococcine line differs
from tetrasporine line in its in ability to divide vegetatively.
According to Fritsch (1935) the filamentous and parenchymatous forms
found in the Ulotrichales are derived from motile unicellular flagellat es of the
Chlamydomonas type; the cell division is in only one plane, i.e., transverse,
followed by the holding together of its products through the formation of
a common cross watl between the cells, thus leading to the development of
the fitamentous habit from the unicetlular ancestors. Cell division of the
unicellular ancestor in two or more planes seems to have given rise to the
parenchymatous habit of the present day Ulva and other similar algae.
Some members of the Tetrasporales which show a filamentous tendency
appear to have given rise to the UIotrichales~ The Chaetophorales probably
 CHLOROPHYTA 77

originated from the Ulotrichales. For the Charales,many authors have pro-
posed a chaetophoralean ancestry. There seems to be a general unanimity
on the derivation of the Siphon ales from the Chlorococcales. The origin
of Conjugales and Oedogoniales is more uncertain; however, Chapman
(1962) favours a ulotrichalean ancestry for the Conjugales, and Klein and
Cronquist (1967) likewise suggested a possible ulotrichalean ancestry for the
Oedogoniales.

Class: CHLOROPHYCEAE
Order: VOLVOCALES

The unicellular and colonial green algae with either a permanently motile,
or a sedentary vegetative phase that is readily revertible to motility, are alt
included in this order. On the basis of vegetative habit this order has been
divided into three suborders: (1) the Chlamydomonadineae, comprising
motile individuals, (2) the Tetrasporineae, consisting of palmelloid forms
and (3) the Chlorodendrineae, including dendroid species. The suborders
Tetrasporineae and Chlorodendrineae have many Chlamydomollas-like
characters, such as eye spot, contractile vacuoles and chloroplast.
The Chlamydomonadineae includes four families distinguishable by cell
wall morphology: the Chlamydomonadaceae has a cellulosic cell wall; the
Sphaerelhlceae has a thickened gelatinous wall provided with a number of
simple or branched processes and a chloroplast of variable morphology,
generally containing one or more pyrenoids; the Polyblepharidaceae includes
naked forms capable of e:xhibiting changes in the shape of the protoplast
and provided with two or more flagella; and the Phacotaceae characterized
by cells that are usually enveloped in a calcified sheath or capsule separated
from the cytoplasmic membrane by an intervening mucilaginous space.

Family:' Chlamydomonadaceae
Genus: Chlamydomonas

Occurrence
Most species of this alga are freshwater forms, or grQw in ditches, tanks,
ponds and takes. Some are marine whereas
others occur on moist, terrestrial habitats. Quite
a few grow in brackish water and in waters flagellum
polluted with sewage or industrial wastes. Chla-
mydomonas also occurs in airborne dust. C.
nivalis accumulates targe quantities of haemato- eye spo t
vacuole

chrome and is found on snow in alpine and nucleus

chloroplaSt
arctic regions, causing 'red snow'.
pyrenoid

Nutrition
Fig. 6-1. Chlamydomonas.
Most species are obligate phototrophs but C. structure as seen under light
dysosmos is a facultative heterotroph and can microscope.
78 A TEXTBOOK ON ALGAE

grow in the dark in presence of acetate as a carbon source.

Morphology
Light Microscopy. Chlamydom()nas is a motile unicellular alga. It is
generally oval but variations in its shape are also noticed. The cell is provid-
ed with a cellulosic wall and gives rise to two anteriorly inserted whiplash
flagella, each originating from a basal granule located in the anterior papillate
or non-papillate region of the cytoplasm (Fig. 6-1). Two, rarely more,
contractile vacuoles are found near the bases of the flagella and a promi-
nent cup or bowl-shaped chloroplast is present in each cell. The chloro-
plast contains at its posterior end a pyrenoid with astareh sheath, and
towards one side of its anterior end is found an eye spot. The single nucleus
remains suspended in the colourless portion of the cytoplasm.
Variability. Chlamydomonas exhibits great variability in cell shape
(Fig. 6-2), insertion of flagella, cell wall, chromatophore (Fig. 6-3 A-G) ,
pyrenoid, and in various other structural and reproductive features. A

Fig. 6-2. Chlamydomonas, some cell Fig. 6-3. Chlamydomonas, chloroplast

shapes. A, C. incerla; B, C. conica; morphology. A, C. ametaslalos; B, C.
C, C. gyroides; D, C. biconvexa; E, C. polydaclyla; C, C. corrosa; D, C. regu-
pseudocoslala; F, C. lunata. (After Ettl, laris; E, C. arachne; F, C. zygnemoides;
1965.) G, C. dijJusa. (After EttI, 1965.)

detailed study of such variability and the importance of such features in the
taxonomie delimitation of different species has been described by Ettl (1965).
Electron Microscopy. The flagellum shows the typical 9 + 2 arrange-
ment of the component fibrils. The cytoplasmic, mitochondrial, nuclear
and chloroplast membranes are all double unit structures. The chloroplast
contains bands composed of variable number of photosynthetic thylakoids
 CHLOROPHYTA 79

which are not organized into grana-like structures (Fig. 6-4). A colourless,
granular matrix separates the
two consecutive bands of a
thylakoid. The chloroplast
thylakoids are neither derived
from chloroplast envelope nor
from pyrenoid lamellae but .............,~ mitochondrion
originate by incorporation of vacuole -ffffHH-f-E)
fresh material into pre-existing 90lgl bodyiH7.HHt~..\'i. lamellae
membran es. The eye spot con- nucleus-tH-IH!f+ -r,~lH+-nucleolus

sists of two or three, more or

slarch~~~
less parallel rows of linearly sheath -fJ!j~'f-pyrenoid
arranged fat droplets. Also
present in the cytoplasm are
endoplasmic reticulum and Fig. 6-4. Chlamydomonas, ultrastructure.
(After Sager, 1965.)
dictyoso·mes (Golgi bodies).

Function
Motility is accomplished by the lashing action of flagella, the energy for
wh ich is derived from ATP (adenosine triphosphate). However, it has been
reported that the flagella of a paralyzed mutant of Chlamydomonas fail to
beat even if the alga is fed with A TP; this is thought to result from the
absence of the enzyme A TPase in the paralyzed mutant.
In C. reinhardii the flagellar apparatus can produce two different kinds
of motion, viz., a breast-stroke which propels the alga forward, and an un-
dulating beat which produces trailing, back ward motion.
The eye spot is apparently not an organ ofphototaxis since mutant strains
of the alga lackjng an eye spot are still phototactic. The removal of water
or other material from the cell is a function generally assigned to the con-
tractile vacuoles. The inability of mutants of Chlamydomo1/as, lacking
contractile vacuoles, to grow in solutions of high osmotic pressure suggests
that contractile vacuoles are involved in the osmotic regulation ofthe cello

Reproduction
Asexual. Ouring favourable environmental conditions, the al ga discards
its flagella and enters into a non-motile phase, its protoplast undergoes two,
three or four successive mitotic divisions (all longitudinal) resulting in the
production of 4, 8, or ] 6 uninucleate daughter protoplasts (Fig. 6-5 A).
Each of these acquires a pair of flagella and a wall before being released
from the mother cell through gelatinization or rupture of the mother wall.
The zoospores gradually develop into mature Chlamydomonas cells.
Under adverse conditions, the daughter protoplasts, instead of growing
into zoospores, continue to divide, each producing a group of2 to 4 aplano-
spores within its own newly formed ge]atinous wall. Meanwhile, the original
parent wall also gelatinizes and the entire groups of aplanospores remain
embedded within a common gelatinous matrix. Such a phase, which may
80 A TEXTBOOK ON ALGAE

be quite short-lived, is called a PaImella stage (Fig. 6-5 B). With the
recurrence of a favourable growing period, each aplanospore changes into
a zoospore which grows in size to become a Clzlamydomonas plant. In
C. nivalis, the aplanospores develop into hypnospores.

8
Fig. 6-5. A, diagram showing wosporc production; B, Palmella stage.

Sexual. It ranges from isogamy to oogamy (Figs. 6-6; 6-7 A, B) and thc
sexually reproducing alga may be homothaIlic (C. gynogama, C. media) or
heterothallic (e. reinhardii). Anisogamy and oogamy havc bccn describcd
on thc basis of observations with materials collectcd from natural habitats.

Fig.6-6. Isogamy.

Thcrcforc, virtually nothing is known about thc nature of genetic or environ-

mental factors controlling sexual reproduction in anisogamous or oogamous
species of Chlamydomonas. Much experimental work, however, has been
 CHLOROPHYTA 81

carried out on the isogamously reproducing heterothaIlic species.

In heterothallic C. moewusii adult individuals themselves copulate. In
other cases, gametes are generally formed by repeated divisions of the proto-
plast of a mother cell and are therefore smaller in size.
In C. debaryanum, the sexual reproduction is isogamous involving a
union between biflagellate gametes. C. braun;; shows anisogamy (Fig. 6-7 A).
The male zoogametes (microgametes) are sm aller in size and are produced
in greater numbers per cell than the female zoogametes (macrogametes).
As a rule, the male cell produces eight microgametes whereas the female
produces only four macrogametes. This difference between two kinds of
gametes is further advanced in the oogamous C. coccifera (Fig. 6-7 B) where
the female cell, instead of dividing to produce gametes, discards its flagella
and directly functions as an immobile female gamete, the egg. The male
gametes, on the other hand, are produced by repeated divisions of the proto-
plast of the cell into 16 smalI, biflagellate microgametes.

A B
Fig.6-7. Chlamydomonas. A, C. braunii. diagram showing anisogamy; B. C. cocci-
/era, ilIustrating oogamy.

Control of Sexuality. In C. reinhardii and C. moewusii (=C. eugametos),

two main kinds of factors-genetic and environmental-are known to affect
the onset and course of sexual reproduction.
Genetic Factors. A pair of sex-determining aIleIic genes designated +
and - exist in the diploid zygote. The complementary genes segregate in
+
the ratio of I : I during meiosis in the zygote, thus giving rise to half type
and half - type individuals (Fig. 6-8). Mating can occur only between
gametes of complementary sexes, i.e., +
and -.
The radiation-induced non-flagellate mutants of Chlamydomonas are
completely asexual suggesting that, for initiation of gametic union, the
N'
r-
presence of flagella is indispensable.
S!:: Environmental Factors. In general, nitrogen deficiency induces gameto-
~ genesis. Ammonium nitrogen is very specific in the inhibition of sexuality
~ (Fig. 6-8). Even the differentiated gametes lose their gametic nature in the
\Ci presence of ammonium nitrogen and behave as vegetative cells. The male
82 A TEXTBOOK ON ALGAE

and female cells differ in their response to light; male gametes ( - mating
type) are produced in thepresence of light and female gametes (+ mating
type) in its absence (Wiese, 1965). The presence of calcium is essential for

game~es .~
'O!.: (' sf----$~~gametes
\b
_
",':,

Jf '<.~~
~i" palrlng
""C:JJ:,, '. ~

~n ~'~
6-'
~ '"
0 ~
c'1
~medium
nltr~nnesis
amet

8/1
p,ametogeneSls , '...' ; I 3 free
~,,~l:;~n Iree ,: ,: ~~iJ, , zygospore 1\~
,..,- §/tQ ~ S~
~

f ]"-'<:
fll ("[ ) mature ~\;;.
~\~'"

,,2'"U.~ ljß
"l'lf \ ,',:' ~ zygospore ~~\~
&~§ ~~O'
6"'/& I ~~
W,"W'''''
_
• <"~
~
t
germinatiOn
~ •
vegetative cell

'" .. 0' '''0''0'"/

~j~
-zoospor<: +l-oospore

Fig. 6-8. Chlamydomonas reinhardii, diagram iIIustrating the experimental control of

sexual cycle.

mating. The intensity of light and temperature also affect the percentage of
mating gametes. With the increase in light intensity from 800 to 6000 lux,
there is a corresponding increase in the number of zygotes (Trainor, 1958).
Further, when grown under continuous illumination, the yield of zygotes in
C. chlamydogama was less than one half of that grown under aIternating
light and dark periods of 12 hr each. The number of zygotes at 27°C was
higher than at 2rC, even though the duration and the intensity oflig~t were
the same in both cases,
Zygote Formation. This indudes five distinct phases: (I) gametic
differentiation, (2) dumping, (3) pairing, (4) plasmogamy, and (5) karyo-
gamy.
The differentiation ofvegetative cells into gametes is conditioned by such
factors as nitrogen deficiency, high light intensity and high C02 concentra-
tion, Clumping (Fig. 6-9 A) involves the cIustering or aggregation of + and
- gametes thro.ugh agglutination and entanglement of their flagellar tips.
The initial attraction for the dumping of complementary gametes is probably
due to the extrusion from their flagellar tips of sexually compatible strands
(Brown, Johnson and. Bold, 1968). It is believed that this agglutination
results from the presence of complementary mating type substances (known
as isoagglutinins) of a glycoproteinaceous nature in the flagellar tips (Wiese,
 CHLOROPHYTA 83

1965). After clumping, the pairing phase starts in which the individual
gametes within a cIump pair off (Fig. 6-9 B) each pair consisting of a +
and a - gamete still united at their flagellar tips. The factors leading to
the formation of the gametic pairs are not known. Subsequent to the pair-
ing phase comes the gametic fusion during which the flagellar union pro-
gresses downward towards the papillate surface of the two gametes. The
gametic pair then establishes a connection or passage through the fusion of the
two papillae, and this is followed by the release of flagellar union, thus
making the gametic pair temporarily motile (Fig. 6-9 C). Immediatelyafter
this, the flagella are discarded and the cytoplasmic fusion is followed by the
fusion ofthe gametic chloroplasts (Brown, Johnson and Bold, 1968). Karyo-
gamy follows plasmogamy, the two nuclei coming to lie in the equatorial
region where they fuse.

A c E
Fig. 6-9. Ch/amydomonas, stages in sexual reproduction.

The zygote (Fig. 6-9 D) is normally a thick-walled structlire whieh germi-

nates after a certain resting period, the meiosis taking place during the first
nuclear division. The haploid zoospores (Fig. 6-9 E) resulting from zygote
germination give rise to Chlamydomonas plants.
Chromosomal and Non-chromosomal Genes. Cytological investigations
have failed to reveal the morphology or number of chromosomes in Chlamy-
domonas. The genetical system of the al ga is very interesting in view of its
having both chromosomal (e) and non-chromosomal (ne) genes affecting the
same phenotype (Sager, 1960). Thus, it has been shown that sensitivity,
resistance, or dependence on streptomycin are governed by c as weIl as nc
genes (Sager, 1962). The requirement for acetate in so me strains is also
known to be determined by nc genes. Recombination for both c and nc
genes has been demonstrated by Sager (1965). The nc genes are thought
to be present on the DNA of the plastids.
Life cycle of Chlamydomonas reinhardii. C. reinhardii is a convenient
experimental subject and some workers have recently grown it in synchronous
culture. Since most of the ceIIs in such cultures are at the same stage of
division or development at any given time, uniformly homogeneous material
can be obtained for the study of sexual process at the molecular level.
84 A TEXTBOOK ON ALGAE

There are two strains of C. reinhardii in respect of the number of zoo-

spores produced per zygote. The strain which produces 8 zoospores is refer-
red to as the octet strain and the one forming 4 zoospores as the quartet strain.
The heterothallic octet strain has a simple li fe cycJe with all those characteris-
tic features of the sexuality typical of higher plants. The synchronization
of mitotic cell division is achieved by simply growing the cells in alternating
light-dark periods. Thc products of mitosing cclls differentiate into zoo-
spores in the presence of a nitrogen source but into gametes in the absence
of nitrogen. When transferred to fresh medium containing combined
nitrogen, the gametes can dedifferentiate into zoospores. Karyogamy occurs
after 12 hr of gametic union and the zygotes undergo a maturation period
of a few days. The mature zygote germinates and divides meiotically on
transfer to a fresh cuIturc medium.
Three different kinds of DNA have bcen detected in C. reinhardii. These
are the Cl, ß and y which differ from each other in buoyant densities and
guaninc-cytosine contcnt. Thc Cl-DNA represents the chromosomal DNA,
the ß occurs in chloroplasts, and the source of y is still unknown. During
maturation of zygote, the y-DNA is replaced by a new type known as
'maturation DNA or M-DNA'. In addition to M-DNA, the zygote con-
tains a- and ß-DNA.
When the + and - gametes differing from each other in respect of two
contrasting pairs of genetic markers are crossed, 4 pairs ofzoospores, in wh ich
the two ceIls of each pair have identical genotype, emerge. This suggests
that crossing over takes place at the 4-stranded stage. This fact together
with the observation that only onc round ofDNA replication occurs between
gametic union and emcrgencc of 8 zoospores, suggests that gamet es contain
double the amount of DNA prescnt in the zoospores.

Genus: Volvox
Occurrence
It is a freshwater planktonic form occurring as green balls of pin-head size
in temporary and permanent pools and ponds. Thc spring and rainy seasons
are the usual periods of thcir active vegetative growth.

Morphology
The colonics (coenobia) generally have a spherical or ellipsoid al contour, and
are motile (Bonner, 1950); the movement is brought about by the joint action
of the flagella of individual cells. The coenobium consists of a mucilaginous
matrix within the pcriphery of wh ich lie all thc cells in a single layer (Fig.
6-10). The central portion ofthe colony is hollow and devoid of cells but is
filled with mucilaginous substance. Each individual cell has its own muci-
laginous sheath which may bc confluent with the sheaths of adjoining cells.
The coenobia of V. aureus are composed of 500-3000 cells whereas those of
V. globalor have up to about 20,000 cells. Other species may have
500-50,000 cells per coenobium. In their general morphology, the indivi-
 CHLOROPHYTA 85

dual cells of Voll'ox resemble CIrlamydomonas and Sphaerella in that a cell

has a pair of anteriorly inserted flagella and a cup-shaped or laminate chloro-
plast containing one or more pyrenoids. Two or more contractile vacuoles

Fig. 6-10. Volvox aureus, coenobium.

and an eye spot are found in the anterior region. Thc cells in the posterior
region ofthe coenobium are usually larger than those in the anterior region.
Most species have prominent cytoplasmic strands connecting the cells of a
colony but V. mononae and V. lertius lack such connecting strands. The
nucleus possesses cytologically distinct chromosomes, the haploid number
for V. globalor and V. aureus being 5.

Reproduction
In contrast to Chlamydomonas, wh ich discharges all its vital functions
(including reproduction) and normally does not die, the cells of a Volvox
colony show pronounced functional specialization.
Asexual. Most of the cells of a coenobium are vegetative but a few in
the posterior region are reproductive (gonidial) which enlarge and lose their
flagella.
Two to 50 asexual gonidia are produced in a coenobium. Before they
divide, the gonidia are slightly pushed into the interior of the colony. The
first two divisions are in planes perpendicular to each other (Fig. 6-11 A)
but subsequent divisions are longitudinal. Thc daughter protoplasts at
the 8-celled stage are in the form of a curved plate called the Plakea stage
(Fig. 6-11 B). At the 16-celled stage, cells are arranged within the peri-
phery of a hollow sphere with an opening toward the anterior end. The cell
division phase in the developing coenobium continues until it attains a
specific number of cells. At the end of this phase, the peripheral row of
almos t spherically arranged cells wh ich are still naked, have their anterior
ends pointing towards the centre of thc dcveloping coenobium. The
86 A TEXTBOOK ON ALGAE

coenobium then undergoes a compIete inversion, i.e., inside-out turning,

through the opening, with the consequence that the anterior ends of cells,
which were previously facing the centre of the sphere, now face outward.
Eaeh ceH of the daughter coenobium finaIly acquires a celI walI and a pair
of flagella at its anterior end. The daughter eoenobia are ultimately
released after the disintegration of the parent eoenobium.

Fig. 6-11. Volvox. A, part of colony showing asexual reproduction; B, Plakea stage;
C-D, stages in sexual reproduction. (After Smith, 1955.)

Sexual. This is oogamous. Species reprodueing sexually may be monoe-

eious (V. globator) or dioecious (V. aureus). Since the monoecious forms
are commonly protandrous, fertilization between antherozoids and eggs
from different individuals is effeeted. The gonidium-like male and female
ceIls are respeetively caIled antheridia and oogonia.
Eaeh antheridium undergoes repeated ceH divisions in a way similar to
that in the development of an asexual gonidium into a daughter eoenobium.
A mass of naked, biflageIlate fusiform antherozoids, ranging in number from
16 to 512 per antheridium, is ultimately produeed (Fig. 6-11 C).
Vegetative eells destined to become oogonia enlarge, lose their flageIla
and become rounded or flask-shaped, with mueh of the oogonial portion
projeeting into the interior of the eolony. The entire eontents of the
oogonium finally form a single non-flagellate egg (Fig. 6-11 D) with a beak-
like protrusion toward one side, through which the antherozoid enters the
oogonium.
During fertilization the entire mass of antherozoids is set free as a unit
eolony; this bundle remains intact until it approaehes an egg. The in-
dividual antherozoids are then set free and one of thern fuses with the eggt
 CHLOROPHYTA 87

leading to the production of the oospore (Fig. 6-1 I D). The oospore subse-
quently secretes a smooth or spiny waJl of three layers; it also accumulates
much haematochrome that colours the zygote orange red. The oospore
constitutes the perennating stage in the life history of Volvox.
During oospore germination, the two outer waJl layers gelatinize and the
inner layer forms avesiele in which the zygote protoplast migrates. The
zygote nucleus divides meioticaJly and of the resulting 4 nuclei, three degene-
rate but the fourth uninucIeate protoplast develops into a coenobium through
asexual reproduction.
The alga is haploid with its diploid phase restricted to the zygote.
A Volvox colony contains only two types of ceJls, the somatie and the
reproductive. The culture filtrate from male Volvox has been fouoo to
induce the formation of antheridia in the male strain and of oogonia in the
female strain. The nature of the filtrate component active in triggering the
process of sexual reproduction seems proteinaceous (Starr, 1968).

Order: CHLOROCOCCALES

This order includes uniceJlular and colonial forms in which the vegetative
thaJlus is non-motile and no vegetative cell divisions oceur. Vegetative cells
lack flagella, contractile vacuoles and eye spot. The eolonial members arise
as a consequence of the union of zoospores or autospores inside the parent
cell wall or soon after their liberation from the parent cell. The thallus
morphology, the features of asexual reproduction (zoosporie or non-zoo-
sporic) and, to some extent, the tendency of the celIs to become coenocytie
have formed the basis of c1assification of the order into families; the two
main series of forms in this order, namely, the azoosporie Chlorococcales
and the zoosporie Chlorococeales are iIlustrated he re with reference to the
representative genera Chlorella and Hydrodiclyon.

Family: ChlorelIaceae
Genus: Chlorella

Occurrence
It is a cosmopolitan, ubiquitous alga oecurring in freshwater, brackish water
and terrestrial habitats. C. liehina is an algal symbiont ofthe lichen Calicium
ehlorina. Some chlorelIas, e.g., Zooehlorella, grow as symbionts in Hydra or
other aquatie animals.

Morphology
Chlorella is a unicellular non-motile alga. The spherical, subspherical or
ellipsoidal cells are bound by a true celIulosic wall. Each cell has a bell-
shaped or cup-shaped parietal chloroplast with or without a pyrenoid '~;~.
4-1 D). There may be a hyaline cavity toward one side of the chloroplast
and in this cavity, or in the colourless central cytoplasm, are located the
single nucleus, the mitochondria and the Golgi bodies. The photosynthetie
88 A TEXTBOOK ON ALGAE

thylakoids lack grana-Iike organization. Toward the end of growth phase

and the beginning of reproductive phase, the cells become multinucIeate.
The fine structure of Chlorella cells has been found to be highty plastic
and polymorphie. Changes in environmental conditions such as light
quality, duration and intensity, temperature, and chemical composition of
the growth medium, greatly influence the micromorphology of the chloro-
plast, pyrenoid and cell wall.

Reproduction and Life Cycle

It reproduces excIusively by the formation of (normally four) asexual auto-
spores. Motile cells, zoospores or gametes are not produced.
Tamiya (see Morimura 1959) succeeded in synchronizing the population
of C. ellipsoidea by subjecting the cells to alternating light-dark cycIes. With
the technique of synchronous culture, four phases (Fig. 6-12) in the life cycIe
of Chlorella were identified: (1) the growth phase during which the auto-
spores grow in size at the expense of the photosynthetic products; (2) the
early ripening phase in wh ich the cells of the growth phase prepare them-
selves for cell division; (3) the post-ripening phase in which the cells divide
twice either in the dark or light; and (4) the division phase during wh ich the
parent wall gelatinizes or ruptures liberating the autospores which in the very
young stage are known as dark nascent cells (Dn ).
The growth and early ripening phases are both light- and temperature-
dependent, whereas the post-ripening and autospore liberation phases are
only temperature-dependent. As shown in Fig. 6-12, the growth and early

Fig. 6-12. Chlorella ellipsoidea, Iife cycle as studied by the technique of synchronous
culture. (After Tamiya, 1963.)

ripening phases are furtber subdivided into five stages distinguisbed on tbe
basis of photosynthetic activity, DNA contents and division performance in
the dark. Tbe Da (dark active) cells are uninucIeate, photosyntbetically
 CHLOROPHYTA 89

very active and are derived from D n ceIls. D-L (dark to light transitional)
cells are also uninucleate but are larger in size and photosynthetically less
active with comparatively higher DNA content than the Da cells. LI cells
are larger with more DNA than D-L cells. All these eells, i.e., Da, D-L,
and LI fail to divide when they are incubated in the dark. The L2 cells are
binucleate and are capable of incipient division in the dark. L3 cells always
become tetranucleate when kept in the dark or light and produce four
autospores. L4 cells are tetranucleate autospore mother cells which liberate
the autospores or Dn cells, independently of light. (L" L2, L3, L4 are light
cells.)
The cell division of Chlorella has a specific requirement for sulphur
because it cannot occur in its absence. Under photosynthetic conditions
it requires nitrogen in addition to a sulphur source, whereas in the dark the
eell division can proeeed normally with sulphur alone (Hase, 1962).

Importance
The alga grows very fast under a variety of conditions and its photosynthetic
pigments and reserve products are similar to those of higher plants. These
attributes have made it a favourite object of extensive study on the mechan-
ism of photosynthesis. Biochemical analyses of its eells have revealed that
they may be very rieh in proteins (about 50%), fats (about 20%), carbo-
hydrates (about 20%), amino acids, vitamins and minerals. In view oftbis,
many laboratory and pilot plant studies on the feasibility of growing Chlorella
in mass cultures for use as human food or animal feed, have been conducted
in Japan, Germany, the United States and Israel. However, the prohibitive
cost of growing, harvesting and processing has prevented its commercial
exploitation.
Another possible use is in the regulation of oxygen and CO2 supply in
nuclear submarines and space vehicles.

Family: Hydrodictyaceae
Genus: Hydrodictyon

Occurrence
Popularly known as 'water net', it grows in the plankton and benthos of
freshwater ponds and lakes. The eommon species, H. reticulatum, is found
free-floating in certain ponds in spring and summer. It is sometimes so
abundant that it alm ost covers ~he entire pond surface. Another species,
H. indicum Iyengar, is common in India.

Morphology
The alga is a macroscopic non-motile coenobium consisting of a network
of pentagons or hexagons with each corner of the polygon resulting from
the union of three large cylindrieal cells (Fig. 6-] 3 A). Each coenobium is
typically cylindrical, c10sed at both ends, and may consist of a few hundred
to several thousand cells in different species. The mature coenobium may
90 A TEXTBOOK ON ALGAE

sometime be as long as 30 cm. The young cells are generally uninucIeate

with a parietal, band-shaped chloroplast having a single pyrenoid. As
the cells grow and enlarge they become coenocytic and the band-shaped

dClughter
net

Fig.6-13. Hydrodictyon. A, part of coenobium showing polygonal meshes; B. formation

of new net inside a parent cell.

chloroplast be comes reticulate and forms many pyrenoids. Mature ceIls,

which may sometime measure 2-3 mm in length, contain a large central
vacuole which displaces the cytoplasm toward the periphery of the cell.

Reproduction·
Vegetative. This is sometimes effected by fragmentation of a net into
two or three nets.
Asexual. It is brought about by the production of a large number (up
to 20,000) of tiny biflagellate zoospores within any vegetative cell of the
thallus. These are not liberated but remain within the motber ceIl and for
a short while may exhibit some movement. The flagella are then with-
drawn and the zoospores elongate into tiny cylindrical cells which co me
together in groups of five or six and dispose themselves in such a manner
as to form a characteristic miniature net of Hydrodictyon (Fig. 6-13 B).
The parental cell wall ultimately softens, liberating the young Hydrodictyon
coenobium which later grows to adult size without undergoing any cell
division.
Sexual. It is isogamous and involves the fusion of biflagellate isogametes
(Fig. 6-14 A, B) produced in large numbers from ordinary vegetative cells.
U nlike the zoos po res, the gametes are liberated from the parent cell through a
hole in thc wall and gametic fusion (Fig. 6-14 C) takes place in water. Thc
zygote is thin-walled and green in colour. It undergoes almost immediate
 CHLOROPHYTA 91

germination by dividing meiotically and produces four haploid zoo-

spores (Fig. 6-14 D). Sometimes, however,
the zygote may perennate. The zoospores
escape from the zygote wall and develop
individually into non-motile polyhedral
cells (Fig. 6-14 E). With the advent of
a favourable growing season each zygote
or polyhedron divides to produce nume-
rous zoospores which arrange themselves
appropriately so as to form a daughter ABC
Hydrodictyon net. Sometimes, a polY-
hedron produces a disk net which
in turn produces zoospores. These then
, yg o,e8\
align to form a cylindrical net. The cells
of the net are observed to grow in size
only after they have established contacts
with adjacent cells.
\WV
'Oo'::~ ~
Order: ULOTRICHALES

This order incIudes unbranched filamen-

tous and parenchymatous forms showing
little structural or functional specialization
among vegetative cells except that the
lowest ceIl of the thallus may be modified
to serve as an organ of anchorage. Depend-
ing on ceIl structure, the order has been
cIassified into three suborders, the Ulotri-
chineae, the Prasiolineae and the Sphaero- Fig. 6-14. Hydrodictyon, stages in
pleineae. Ulotrichineae have uninucleate sexual reproduction. A-B, gametes;
C, zygospore; D, zygote with its
cells with a single, parietal, band- or girdle- germination product (zoospores); E,
shaped chloroplast and are segregated into a polyhedron.
four families on the basis of wall structure
and thallus organization; the Ulotrichaceae include simple filamentous forms
with ceHs having a one-piece simple cell wall and a girdle-shaped chloroplast;
the Microsporaceae have one filamentous genus in which the cell wall is
composed of overlapping H-shaped pieces; in Cylindrocapsaceae the cells
are elliptical with a thick, stratified wall and in old filaments some cells may
divide obliquely so that some parts of the filament become multiseriate; the
Ulvaceae contain forms with parenchymatous thalli and are retained in the
Ulotrichineae because Iike Ulothrix their young stages are generally fila-
mentous.
The Suborder Prasiolineae with a single Family Prasiolaceae includes only
one membranous or thalloid genus Prasiola whose cells contain an axile
stellate chloroplast.
The Sphaeropleineae is also a monogeneric Suborder reprcsented by
92 A TEXTBOOK ON ALOAE

Sphaeroplea, an unbranched filamentous alga made up of large coenocytic

cells eaeh provided with several band-shaped or discoid chloroplasts.
The prevailing modes of sexual reproduction and life cycle are: (I) iso-
gamous and haploid (Ulotrichaceae and Microspora); (2) isogamous and
isomorphie (Ulvaceae); and (3) oogamous and haploid (Cylindrocapsa and
Sphaeroplea). Prasiola exhibits an interesting type of life cycle in that the
thallus is either completely diploid, producing diploid zoospores; or is partly
diploid and partly haploid, with the gametes being produced from the cells
of haploid tissue (eole and Akintobi, 1963). In certain species of Prasiola
patches of gametophytic tissue develop upon the diploid sporophyte follow-
ing meiosis.

Family: UIotrichaceae
Genus: Ulothrix

Occurrence
The alga occurs in flowing waters, ponds, pools, lakes and in reservoirs where
the water is continuously renewed. It also grows on stones and pebbles lying
within the spray zone of waterfalls. When young, it generally grows attach-
ed to submerged objects. U. zonata is found in cold water streams, U. ftacca
is marine and U. implexa is a lithophyte in estuaries.

Morphology
Thc plant body is unbranched uniseriate filament composed ofshort, cylindri-
cal, quadrate or squarish cells (Fig. 6-15 A) of which the basal ceIl is colour-
less and modified to function as a holdfast. The growth is diffuse, every cell
(except the basal holdfast) being capable of division. The vegetative cells
are uninucleate, possess a cell wall and a girdle-, collar- or ring-shaped
chloroplast having one or more pyrenoids (Fig. 6-15 A, B). The ring-shaped
chloroplast may be closed or open at one end (Fig. 6-15 C, D).

Reproduction
Vegetative. A filament fragments into two or more pieces, each ofwhich
then grows into a new plant.
Asexual. Each cell except the basal holdfast produces up to. eight, rarely
16, zoospores and usually there exists an apical-basal gradient in their forma-
tion. The mother protoplast divides mitotically (Fig. 6-15 E), producing
naked and stigmatic zoospores which resemble morphologically the uni-
cellular Volvocales and which are liberated from the parent cell through a
pore in the lateral wall (Fig. 6-15 F). Depending upon the number of divi-
sions, the mother protoplast may produce quadriflagellate macrozoospores
(Fig. 6-15 G), quadriflagellate microzoospores (Fig. 6-15 H) or biflagellate
microzoospores (Fig. 6-15 I). The liberated zoospores swim about for some
time, anehor themselves to some solid object by their anterior ends, with-
draw flagella and secrete a cell wall. Each then divides into two cells, the
lov.·er forming the holdfast and the upper dividing and redividing to produce
 CHLOROPHYTA 93

vegetative cells of the filament (Fig. 6-15 J). The filaments formed from
macrozoospores are much stronger and healthier than those derived from
biflagellate microzoospores. Quadriflagellate microzoospores on germination
produce intermediate filaments.

~~Y~~d "/' nucleus

"" .
, ':'.'~.'

cytoplasmiC
C dlsc

OPY"""'"

E
A F J
Fig. 6-15. Ulolhrix. A, part of filament; B, three-dimensional view of a cell wlth
ring-shaped chloroplast; C, cross section through a cell with ring-shaped chloroplast;
D, a horse-shoe-shaped chloroplast; E, division of protoplast of a cell; F, liberation
of swarmers in a mucilaginous vesicle through a lateral pore; G, quadriftagellate
macrozoospore; H, quadriftagellate microzoospore; I, biftagellate swarmcr; J, germling
with holdfast.

Asexual reproduction also takes place by aplaoospores. The daughter

protoplasts iostead of forming zoospores, differentiate into non-motile thin-
wallcd aplanospores which may either be released from the parent cell or
remain enclosed within it during the unfavourable growth period. With the
onset of favourable periods, each aplanospore gives rise to a zoospore, or
germinates directIy ioto a filament.
Sexual. Almost all the experimeotally iovestigated species are sexually
heterothallic and gametic union (Fig. 6-16 A, B) always occurs betwecn
morphologically similar gametes
of opposite, + and - mating
types. Up to 32 or 64 gametes
per cell are formed and in the
mode of formation and morpho-
logy, they resemble the biflagel- A B C
late microzoospores. The zygote
(Fig. 6-16 C) is a thick-walled Fig, 6-16. Uloillrix, sexual reproduction.
resting structure and on germina- A-B, fusing gametes; C, quadriflagellate
zygote; D, germination of zygote and pro-
tion (Fig. 6-16 D) its nucleus duction of meiospores.
divides by meiosis during which
the + and - sex factors segregate giving rise to haploid zoospores or aplano-
94 A TEXTBOOK ON ALGAE

spores, half of whieh are of + type and the other half of - type. U p to 16
zoospores or aplanospores may be produced from a germinating zygospore,
though four is the usual number. Reduetion in ehromosome number during
germination of the zygote nucleus has been observed.
Tbe rare parthenogenetie development of agamete into a vegetative fila-
ment and also the occasional gamete-Iike behaviour of the biflagellate
Zoospores have also been recorded.

Order: CLADOPHORALES

This order includes fresbwater and marine filamentous a.1gae, mostly brancb-
ed, having large multinueleate cells whieh are thick-walled and possess
reticulate chloroplasts and many pyrenoids. Most members occur in rela-
tively clean, flowing water of streams. These algae generally do not occur
in absolutely quiet waters.
Reproduction takes place by fragmentation, by bi- or quadriflagellate
zoospores and by biflagellate isogametes. The Iife eycle involves an alter-
nation between isomorphie generations.

Family: Cladophoraceae
Genus: Cladophora

Occurrence
A large genus with ab out 150 speeies, Cladophora occurs in diverse fresb-
water, brackish water and marine habitats. Plants may be free floating or
attached on rocks, aquatic angiosperms or shells of molluscs. Some species
grow so prolifically during spring and summer as to form dense mats,
blankets or ropy aggregates, bloeking and clogging the streams. Other
species produce sponge-Iike balls in deep water. C. glomerata is a typical
freshwater species, often the most abundant filamentous alga found in
streams throughout the world. C. crispata becomes attaehed to the walls
ofmunicipal water reservoirs. Massive growth of various species of Clado-
phora in canals inipedes the flow of water through water distribution
ehannels.
Most species prefer hard waters having relatively high pR (about 7.5 to
9.5).

Morphology
Cladophoras are profusely branehed and, when young, are attached to the
substratum by septate rhizoidal downgrowths from the basal region of the
filaments or their lowermost cells (Fig. 6-17 A). The rhizoidal branches are
often perennial and help in perennation of the alga through unfavourable
conditions.
Cells are large, cylindrical, multinucleate, and have an elaborate chloro-
plast which may form a continuous reticulate network in younger cells but
may become parietal in older stages (Fig. 6-17 B, C). In specimens collected
 CHLOROPHYTA 95

from shaded habitats the chloroplast occurs as a den se reticulum whereas in

those from weIl exposed localities, the chloroplasts are cIearly separated from

Fig. 6-17. Cladophora jongionum Van den Hoek. A, habit; B, C, apical and inter-
calary cells with parietal chloroplasts.

each other (Whitton, 1970). Many pyrenoids occur in the chloroplast.

Cytoplasm is located centrally in the cell and contains many nucIei. Unlike
most uninucIeate plant and animal ceIls, in this al ga mitosis and cytokinesis
are quite independent of each other and this is why the cells are multi-
nucleate.
96 A TEXTBOOK ON ALGAE

The branching (see Fig. 6-18 A) of the thallus is usually dichotomous

and rarely trichotomous. Alternate (lateral) branches are also known. The
branches originate in a characteristic manner, just beneath the septum
between two cells. A bulge appears near the upper end of a cell (Fig. 6-18 B).
A portion of the cell contents of the parent cell passes into the bulge which
is then cut off by a septum. This septum is parallel to the long axis of the

n B

c
Fig. 6-18. A, Cladophora sp. showing branches; B-E, succcssive stages in evection.

filament and at right anglcs to the septum between the two parent cells. The
cell bulge is then moved upward and outward as a result of localized active
growth of the parent cell membrane just beneath the bulge (Fig. 6-18 C).
Consequently, the 90° angle between the new and old septa increases pro-
gressively (Fig. 6-18 D) until, in some cases, the two septa align in a
 CHLOROPHYTA 97

straight line (Fig. 6-18 E). By this stage the new bulge will have grown
into a new ceIl or even a sm all branch and the mode of branching will
appear dichotomous (Fig. 6-18 E) instead of lateral. Such a process of
branch formation has been styled evection and is characteristic of many
species. OccasionaIly, two such buds may be formed instead of one and
then the branches will appear trichotonious.
Unlike Ulothrix, the mode of growth in Cladophora is both apical and
intercalary. The relative extent of apical and intercalary growth in different
species, however, varies widely; in C. rivularis it is almost entirely intercalary.
The ceIl wall is thick, stratified and composed of layers of microfibrils
(mostly cellulosic) arranged at right angles to each other. The mierofibrils
of the lateral walls are often continued into those of the cross septa. The
microfibrillar layers frequently become so compaet with the amorphous
layers that the lamellar details of the outer wall become obscure.
The branches appear coarse and stringy to the touch and as there is little
secretion of mucilage, they harbour many diatoms and other microorganisms
as epiphytes.
Reproduction
Vegetative. A filament may fragment into two or more pieces, each of
wh ich may form a new plant. The smaller branches often become detaehed
from the parent lhallus and then grow into new plants.
Asexual. The protoplasts of apical or subapical cells break into uni-
nucleate segments whieh are transformed into a bi- or quadriflagellate
zoospore. Mature zoospores are released through pores formed in the
lateral walls of the cells. Zoospore formation oeeurs in ordinary, unmodi-
fied eells.
Following aperiod of motility and dispersal, the zoospores settle down
on some substratum and germinate to give rise to Cladophora plants whieh
are morphologically identical with the zoospore-producing parent thalli.
The reduction division in some speeies has been observed to occur at the
stage of zoospore formation.
Sexual. The apieal or subapical cells produce biflagellate isogametes.
Fusion of gametes from + and - strains oecurs outside the parent eells.
The zygote germinates after a rat her brief rest period to grow directly into
a diploid plant. The alternation of generations is isomorphie.
Fortnightly peak periods of gametogenesis and zygote formation have
been reported for this alga and it is thought that sueh periodicity may be
connected in some way with lunar tide cycles.
Many speeies have been shown to be polyploids. The number of
nucleoli and chromocentres has also been correlated with tbe degree of
~ polyploidy.
~ Seasonal Growth Cycle. Whitton (1970) has reviewed the seasonal growth
N-
<'l behaviour of C. glomerata in eutrophie waters of temperate rivers. Thc peren-
~,
7.! nating filaments attachcd to rocks start producillg new branches in spring.
r..: After a maximum of growth and branch formation in summer there is a
98 A TEXTBOOK ON ALGAE

slowing down of vegetative growth. A second period of rapid growth occurs

in early autumn. It is thought that such growth occurs from zoospores
produced in the preceding summer. During late autumn, the upright parts
of thalli become detached, leaving behind the basal fragments still attached
to rocks. Thick-walled akinetes are occasionally formed.
In standing water, on the other hand, thalli and akinetes survive the
winter On the bottom of the ponds under a layer of silt. Zoospore release
occurs during May-June.

Order: CHAETOPHORALES

The heterotrichous habit, i.e., the differ~ntiation of branching filaments into

erect and prostrate systems, is the most outstanding singular feature ofthis
order. These plants may further be characterized by the frequent produc-
tion of hairs (e.g., in Chaetophoraceae) or setae (e.g., Coleochaetaceae) or by
the formation of differentiated reproductive organs Iike sporangia and game-
tangia (e.g., Trentepohliaceae). The hairs have a single or a group of nar-
row elongated cells with weII-marked tapering apices whereas the setae or
bristles are the outgrowths of the wall with pointed ends. Certain members,
e.g., Trentepohliaceae, exhibit a marked terrestrial or subaerial tendency.
Fritschiella tuberosa (Chaetophoraceae) is exclusively terrestrial, its prostrate
system is buried in moist soil and serves as a means of perennation.
Except for Aphanochaete, Chaetonema and Coleochaete,other members
of the Chaetophorales show isogamous sexual reproduction.
According to Fritsch (1935), the heterotrichous habit represents the
highest type of vegetative organization in the green algae. He has postu-
lated that the Chaetophorales represent tbe surviving descendants of ancestral
forms from whicb higher land plants originated in the remote past. Of the
Chaetophorales, Fritschiella seems to stand nearest to the probable ancestral
forms because of its heterotrichous habit coupled with a terrestrial mode
of life.

Family: Chaetophoraceae
Genus: Fritschiella

Occurrence
Fritschiella tuberosa Iyengar is an interesting monotypic tropical al ga
widespread in India, Nepal, South Africa and many other countries. It
grows on moist soils of drying up rain water pools. It is eitber terrestrial
or amphibious but generally does not grow in free water.

Morphology
First described by the late M. O. P. Iyengar from South India, this
species was later studied in detail by R. N. Singh from Uttar Pradesh and
other places.
This alga is probably the most highly differentiated species among the
 CHLOROPHYTA 99
green algae. It is heterotrichous and filamentous and is endowed with the
potentiality of alm ost every conceivable kind of somatic advancement possi-
ble in a heterotrichous organism.
The thallus is diffcrentiated into four distinct regions (Fig. 6-19 A):
(I) the rhizoidal system, consisting of colourless, branched or unbranched,

primary erect
system

prostrate
system

c o
Fig. 6-19. Frilschiella. A, habit; B, funnel-shaped, perennating, tuberous body; C,
intercalary sporangium; D, terminal sporangium (after Patel and Patel, 1969).

multicellular rhizoids comparable structurally and functionally with the

rhizoids of bryophytes; the rhizoids not only anchor the plant to the sub-
stratum but also serve to absorb soil nutricnts; (2) the prostrate system,
composed of uni- or multiseriate (pseudoparenchymatous) clusters of rounded
100 A TEXTBOOK ON ALGAE

or irregular ceHs with dense contents (proteins, starch, haematochrome, ete.);

it is generaHy filamentous in the early stages of development in young plants;
sometimes its filaments become further differentiated into nodal and inter-
nodal regions; ceHs or filaments of the prostrate system give rise on their
lower side to rhizoids and on the upper to the primary erect system; they
are meant for food storage, perennation, and reprodllction; (3) the primary
ereet system, having uni- or biseriate ceHs very similar to the ceHs of the pros-
trate system; these are partly photosynthetie and partly condueting in fune-
tion; and (4) the secondary ereet (projeeting) system, eonsisting of freely
branched uniseriate filaments with green, elongate, uninucleate cells, eaeh
having a parietal chloroplast with a few pyrenoids; this system is probably
meant exclusively for photosynthesis.
The rhizoidal and prostrate systems are subterranean, being completely
em bedded beneath the surface of soil. The primary erect system is generally
partly embedded and partly projeeting above ground level. The secondary
erect or projeeting system is fully exposed to light and air.

Perennation
During adverse periods the ereet and rhizoidal systems eommonly degenerate
and disappear and the alga perennates by means of its prostrate system.
U nmodified cells of prostrate system tide through adverse periods as such
but the nodal regions of prostrate system, being capable of independent
existence, may get detached from the parent thallus through death and decay
of internodal regions, and thus not only serve the function of perennation
hut also vegetative propagation. Yet another mode of perennation is hy the
formation of speeially modified tuber-like bodies (Fig. 6-19 B) enclosed by
cell walls having a dark and thick cuticle. Formed in prostrate system,
these tubers are commonly either funnel-shaped or c1ub-shaped (Fig. 6-19 B).
Each such tuber may be up to 1 mm in length.

Reproduction
Asexual. It takes place by formation of bi- or quadriflagellate zoospores,
one or two of which are normally formed from a mother cell. Such zoo-
spores are structllrally similar to those of Ulothrix. Pate I and Pate I (1969)
have recorded differentiated intercalary and terminal sporangia (Fig. 6-19 C,
0) which are somewhat larger in size than the ordinary vegetative cells.
However, the nature and fate of the products of such sporangia are not
known.
Sexual. An interesting feature of F. tuberosa is the combination of a
highly differentiated vegetative thallus with an equally simple and primitive
mode of sexual reproduetion, viz., isogamy.
Biflagellate isogametes are formed from eeHs of prostrate system (Singh,
1954). After fertilization the zygote germinates to produee a diploid plant
(2n = 8) which is morphologically identieal with the parent, gametophytie
plants. Meiosis is believed to occur during formation of asexual spores by
the sporophyte. The haploid zoospores give rise to gametophytie plants
 CHLOROPHYTA 101

(n =4). Thus, the al ga exhibits an isomorphie alternation of generations.

Fritschiella has attraeted eonsiderable attention in connection with
speeulations on the algal aneestry of higher plants (see Singh 1954). It has
been strongly advocated that the ancestors of the early land plants were
green algae very similar to the present day Fritschiella. The reason why
such aneestors are sought among the green algae is that the basic physio-
logieaI, bioehemical and metabolie eharaeteristics of the high er plants (from
bryophytes onwards) can only be compared with those ofthe Chlorophyceae.
Within the Chlorophyceae, Fritschiella is considered as coming nearest to
the hypothetical ancestors on the basis of its foIlowing attributes:
1. terrestrial or amphibious habitat;
2. rhizoidal system strongly resembling the rhizoids of bryophytes both
structuraIly and functionaIly;
3. prostrate system being very much Iike the subterranean rhizomatous
structures of certain pteridophytes and angiosperms;
4. reproduction being mostly confined to prostrate system as in Antho-
cerotales in which reproduetive bodies are endogenous and embedded in
thallus;
5. tropical distribution (it is widely believed that the first transmigrants
were tropical forms); and
6. the high degree of thallus differentiation indicating the existence of a
strong potentiality for further elaboration in a chaetophoraceous alga in
the direction of evolution of an early algal transmigrant form.

Family: Coleochaetaceae
Gen us: Coleochaete

Occurrence
Coleochaete, the sole genus belonging to the Family Coleochaetaceae, has
some 10 species which grow epiphytically on the leaves and sterns of
freshwater higher plants such as Hydri/la and Potamogeton, or as endophytes
in Nitella.

Morphology
C. pulvinata is heterotrichous with the branched filaments of the erect system
organized into a more or less hemispherical cushion. In C. scutata (Fig. 6-20
A) the erect system is absent and the prostrate system is discoid showing
radial arrangement of its congregated branches thus giving rise to a pseudo-
parenchymatous disc which grows by means of apical (marginal) meristems.
The branching of thaIlus may result from lateral outgrowths or dichotomy
of the apical meristems. The ceIls are uninucIeate, and contain an irregular
parietal chloroplast with one or two pyrenoids. Coleochaete differs from
other members of the Chaetophorales in its chromosome number and
morphology.
Some or most of the cells of the thallus produce setae, each of which is
~omposed of a eytopl<~smic thread completely or partially encIQsed within a
102 A TEXTBOOK ON ALGAB

basal sheath. The setae are generally regarded as outgrowths ofthe plasma
membrane emerging through a pore in the cell wall. They readily break
off after some time, leaving only the basal sheaths.

Reproduction
Asexual. Single, ovoid, biflagellate zoospores are produced in ordinary
vegetative cells. They possess a lateral chloroplast but are devoid of an eye
spot. After abrief movement, the zoospores withdräw their flagella,osecrete
a wall and divide in a horizontal plane; the upper cell forms the seta and the
lower divides in one or two planes at right angles to each other to form the
primary initials which by their activity produce a discoid thallus (C. scutata).
In heterotrichous forms (C. pulvinata) the germinating zoospores at first
produce the branched prostrate system from wOhich the branches ofthe erect
system arise later.
Sexual. C. pulvinata is monoecious whereas C. scutata is dioecious.
The sexuality is of an advanced oogamous type and is accompanied by a
kind of fruit formation not found in any other green algae. The hetero-
trichous species bear the sex organs on the branches of the erect system.
In C. pulvinata antheridia are produced terminallyon branches, some-
times on the same branch that bears an oogonium. Antheridial initials
arise as colourless outgrowths from the terminal cells ofa branch (Fig. 6-20B)
and one antheridium gives rise to a single biflagellate oval or spherical, naked
gamete.
The antheridia of C. scutata are produced from some intercalary cells
lying within the periphery of the thallus. At tbe time of antheridial forma-
tion, an intercalary cell divides into a number of daughter cells each of which
finally differentiates into an antheridium. Frequently tbe antberozoids of
C. sculala are green rat her than colourless.
The oogonia of C. pulvinata are also produced terminally and singly but
their seemingly lateral position is due to growth and production of branches
from the cells beneath the oogonium (Fig. 6-20 B). Bach oogonium is a flask-
shaped structure wit~ a swollen base containing an egg and asIender neck
or trichogyne. This type of oogonium strongly resembles the carpogonium
ofthe Rhodophyta. Discoid species have typically papillate oogonia lacking
trichogynes.
During fertilization an antherozoid enters an egg through an apical open-
ing formed in the trichogyne or in the papilla ofthe oogonium. The resulting
oospore enlarges in size and certain branches from the neighbouring cells of a
fertilized oogonium grow out forming a rat her loose pseudoparenchymatous
envelope or jacket that completely encircles the oospore. The oospore plus
tbe envelope is ca lIed a spermocarp (Fig. 6-20 C) which is analogous to an
angiospermic fruit.
After perennation, when the oospore germinates, its nucleus divides
meiotically foIlowed by a few mitotic divisions, producing a 16- to 32-celled
structure (Fig. 6-20 D). Bach of these ceIl» farm" a biflagellate zoospore
 CHLOROPHYTA 103

(Fig. 6-20 E) which grows into a new plant. Coleochaete is haploid, with
oospore representing the only diploid stage in the life cycle.

gonium

sterile jacket B

~ ~ß~
~~<'l~G1 ~ f[
~~ ::d
c 0 ~\ E
Fig. 6-20. Coleochaete. A, C. scutata; B, C. pulvinata; C, spermocarp; D, libera-
tion of meiospores (zoospores) from germinating spermocarp; E, zoospores.

Order: OEDOGONIALES

The two main distinguishing features of this order are: (I) the mode of cell
division leading to the formation of characteristic caps on certain ceHs, and
(2) the production of multiflagellate zoospores or gametes.
The single Family Oedogoniaceae comprises the three genera Oedogonium,
Bulbochaete and Oedocladium which differ from one another in thallus
organization. Oedogonium is an unbranched, filamentous form growing
attached to the substratum when young but often becomes free-f1oating at
maturity. Bulbochaete is a profusely branched alga with unilateral branches
lying on alternate sides and almost every ceH bearing a characteristic attenuat-
ed hair with a bulbous base; the terminal eeJls are generally provided with a
pair of sueh hairs. The Oedocladium is terrestrial, and heterotriehous with a
prostrate system bearing numerous branehed, septate and eolourless rhizoids
on its lower side and branehes of the ereet system on its upper side.
104 A TEXTBOOK ON ALGAE

Family: Oedogoniaceae
Genus: Oedogonium

Occurrence
Oedogonium is a widespread freshwater alga that grows epiphyticaIly in
ponds, pools, and shaIlow tanks.

Morphology
The filaments are unbranched and consist of cylindrical ceIls except for the
basal ceIl which is modified into a holdfast. Each ceIl is provided with a
thick wall differentiated into an inner cellulosic, amiddIe pectic and an outer
chitinous layer. The terminal cells of the filaments are generally rounded or
acuminate and theintercalary cells often show an apical-basal polarity. Some
or most of the cells at the distal end exhibit parallel striations or annular
scars and are known as cap cells (Fig. 6-21).
Cells are uninucIeate and when mature
have a central vacuole containing cell sap; l J
and an elaborate, reticulate chloroplast
containing many pyrenoids (Fig. 6-21). A
sheath of starch granules surrounds each
pyrenoid .
. The cells of Oedogonium are similar to
nucleus
those of other green algae except that their
chloroplasts contain microtubules each made
up of two helically wound subunits. Such
elements have also been observed in the
chloroplasts of zoospores, zoospore germ-
lings and eggs (Hoffman, 1967). Thesemicro-
tubules are considered to provide structural
cap
support to the massive reticulate chloroplast
and perhaps also to facilitate its growth and
development.
Vegetative Ce/! Division. As the cell
enters the division phase, the nucIeus moves
from the lateral position to the centre.
So on a transverse ring of wall material
appears on the inner face of the lateral wall
just below the apical end of the cello The
nucIear division, the growth of the ring in
thickness and the formation of a groove
encIosing the growing ring occur concomit-
antly. An unattached floating septum is
formed between the two daughter nucIei.
( l
The middle and outer wall layers extern al Fig. 6-21. Oedogonium, part of
vegetative filament showing cell
to the groove then rupture, permitting ffee structurc llfid a cap cell witjl four
elongation of the ring wh ich forms a new c;aps,
 CHLOROPHYTA 105
piece of cell walllying hetween the cap and the sheath. Ultimately, the
floating septum moves upward and becomes fixed near the terminus of the
old cell wall. The new cell has the wall formed from the thickened ring
and the newly synthesized piece. The membranous striation ofthe ruptured
parental wall at the anterior region of the upper daughter cell is the cap and
the cell bearing it is known as a cap cell. The number of caps on a cell
indicates the number of cell divisions that have taken place (Fig. 6-21).
Arecent ultrastructural study (Fig. 6-22) of the mode of cell division in
Oedogonium borisianum (Hili and Michalis, 1968) has revealed that: (I) the
ring (Fig. 6-22 A) below the cap originates as a three-layered structure by a
method which excJudes the possibility of its origin from an infolding of the
innermost walilayer; (2) the ring grows in size with a gradual addition of
vesicular material from the cytoplasm; (3) the mother wall adjacent to the
fully-formed ring splits off; (4) the single, peripheral nucleus migrates to the
centre and karyokinesis occurs; this is followed by the formation in the

cap

·LJ ' .

G D
septum

o
o
~.li',,-'"111:.'~

A B c
no E
Fig. 6-22. Oedogonium borisianum, stages in cell division.

internuclear region of a row of microtuhules (Fig. 6-22 B) in a plane parallel

to that of the future cross wall; (5) concurrent with this development the
ring expands into a cylinder (Fig. 6-22 C) which becomes the cuticle of the
upper daughter cell. The septum, which for some time remains unattached
to the lateral wall, later migrates upward to the base of the cylinder (Fig.
6-22 D); (6) soon ~ new lateral wall is formed between the cuticle and plas-
106 A TEXTBOOK ON ALGAE

malemma of the upper daughter cell; (7) finally, the upward migrated
transverse septum grows into a mature cross wall which unites with the
newly formed wall on either side (Fig. 6-22 E). The mode of cross wall
formation rules out the possibility of cell division either by cytokinesis or
by annular furrowing of the cytoplasm.

Reproduction
Vegetative. Fragmentation is the usual method of propagation but
under certain conditions the alga mayaIso propagate by akinete formation.
Asexual. It occurs by the formation of multiflagellate zoospores pro-
duced singly within a cap cell. During their formation the cell contents
contract slightly and develop'a semicircular hyaIin~ area on one side of the
protoplast. A ring of basal granules appears at the base of the hyaline area
and from each comes out a single flagellum, thus forming a ring of flagella
around the base of the hyaline area. The basal granules remain connected
together by fibrous strands. After the development of the zoospore, the
cell wall near the cap region opens apart and the single zoospore moves out
ofthe cell in a mucilaginous vesicle which so on gets dissolved Iiberating the
zoospore (Fig. 6-23 A, B). The zoospore moves to and fro for so me time,

8 c
Fig.6-23. Oedogonium. A, formation and liberation of zoospore; B, one zoospore
with crown of flagella; C, oogamous reproduction in a homothallic macrandrous
species.

settles down on a suitable substratum, and attaches itself to the substratum

with its hyaline flagellar end. Thereafter the flagella are withdrawn and
the quiescent zoospore e10ngates considerably. The attached end of the
growing zoospore may differentiate into a smooth or lobed hold fast depend-
ing on the smoothness or roughncss of thc substratum respectivcly.
 CHLOROPHYTA 107

Sexual. Reproduction is oogamous and is of two kinds: (1) macran-

drous, and (2) nannandrous.
Macrandrous forms may be monoecious producing antheridia and
oogonia on the same plant (Fig. 6-23 C) or dioecious, producing antheridia
and oogonia on different individuals. In nannandrous forms the sexual
plants are dimorphic-the oogonia being formed in filaments of normal
size whereas antheridia are produced in filaments, known as dwarf males
(Fig. 6-24) or nannandria, which are extremeiy small and are always found
attached to oogonia proper or to the underlying cell.
Macrandrous Forms. The antheridial mother cell may be terminal or
intercalary and in either ca se it divides vegetatively into an upper antheri-
dial cell and a lower sister cell. The latter undergoes aseries of vegetative
divisions to produce a row of two to many antheridia. An antheridium
may give rise to a single multiflagellate antherozoid or it may undergo a
single transverse or vertical division followed by the transformation of each
of the two daughter ceIls into an antherozoid. Although quite similar to
the zoospores, the antherozoids are smaller and may have fewer flagella.
The oogonia are produced from cells which act as oogonial mother cells.
Each of these cells divides into an upper
cap cell functioning as oogonium pro-
per, and a lower supporting celL The
latter may again behave as an oogonial
mother ceIl, thus forming a row of two oogonium
dwarf
or more consecutive oogonia. Or, it may male
remain vegetative, in which case the
oogonia are solitary. The oogonium,
which is filled with reserve food, enlarges
to some extent and assumes a more or
less spherical shape. A colouriess region,
called the receptive spot, appears at one
point near the side wall and through it
an antherozoid finds its way into the egg.
Nannandrous Forms. Each dwarf
male or nannandrium is a few-celled
filament with a basal stalk cell and two
to three antheridial cells. Dwarf males
are derived from antherozoid-Iike zoo-
spores, known as androspores, which are
formed singly within antheridia-like cells
called androsporangia. A species that cap
bears both oogonia and androsporangia
is designated gynandrosporous, whereas
the one that bears them on different
Fig. 6-24. Oedogonium, a nannandrous
filaments is termed idioandrosporous. species.
The changes that take place during the
development of androsporangia and androspores are similar to those occur-
108 A TEXTBOOK ON ALGAE

ring in the production of antheridia and antherozoids. After their libera-

tion from androsporangia the androspores swim about for some time and
then only one or a few of them may settle on an oogonium or a suffultory
cell. The androspore then germinates into a dwarfmale. Each antheridium
of the dwarf male commonly produces two antherozoids Iying one above
the other.
Oogonial development in nannandrous species is similar to that in the
macrandrous species. In certain heterothallic, nannandrous forms, some
hormone is excreted by oogonial mother cells that attracts the androspores
to the oogonia. A gelatinous envelope is formed around the developing
oogonia and the attached dwarf male, and this serves to trap the spermato-
zoids so that the sperms remain in the immediate vicinity of the oogonium.
The oospore becomes thick-walled. After remaining dormant for some
time it germinates and its nucleus divides by meiosis to produce four proto-
plasts each of which forms a multiflagellate zoospore which grows into a new
haploid Oedogonium plant. In so me heterothallic species two of the four
zoospores develop into male filaments whereas the other two develop into
female filaments.

Order: CONJUGALES
This order includes both unicellular and simple, unbranched filamentous
forms which differ from ·other green algae in: (1) elaboration or complexity
of chloroplast, (2) absence of flagellated stages, and (3) occurrence of sexual
reproduction through conjugation involving the fusion of amoeboid gametes.
The cells commonly have a few, large and elaborate chloroplasts which
are in the form of: (1) a f1at axile plate, as in Mesotaenium and ."Jougeotia;
(2) a pair ofaxile stellate chloroplasts, as in Cylindrocystis and Zygnema;
and (3) one or more spiral, ribbon-like chloroplasts, as in Spirotaenia and
Spirogyra. The unicellular forms, popularly known as "desmids", contain
an axile (rarely parietal) chloroplast.
With rare exceptions, the chloroplasts contain large and well-defined
pyrenoids. The cells have a single, conspicuous, centrally located nucleus.
Many members have unusually a large number of rather small-sized
chromosomes.
The usual methods of reproduction are vegetative and sexual, though
rarcly asexual reproduction occurs by the formation of akinetes and
aplanospores.
Vegetative multiplication in the unicellular desmids is brought about by
cell division. In filamentous forms fragmentation is often attended by the
formation of special types of septa which may be replicate or semireplicate.
Sexually, the unicellular Conjugales reproduce by the formation of a
papillate bridge between two cells through which the contents of one, desig-
nated as male, pass into the other, termed as female. The contents of both
the conjugating cells may sometimes pass into a conjugation tube and gametic
fusion may take ~Iace within the tube rather than within one Qfthe two cells.
 CHLOROPHYTA 109

Family: Zygnemaceae
Genus: Spirogyra

Occurrence
A freshwater submerged or free-f1oating alga, it occurs in various water bodies
of both temporary and permanent nature. The filaments are slimy to touch
because of an outer mucilaginous wall layer.

Morphology
The plant body is an unbranched filament consisting of a uniseriate row of
cylindrical cells with the basal cell frequently developing into a branched, or
highly-Iobed, anchoring organ in those species which grow attached to the
substratum. The cell wall consists of two layers, an inner cellulosic and an
outer pectic layer. The septa between the cells may be plane, semireplicate
or replicate, änd as in higher plants, their formation is always followed by
the secretion of a cellulosic layer along both of its sides. A large vacuole
containing tanniferous vesicles lies in the centre of the cell. The cytoplasm
forms a thin layer Iining the cell wall, and the single nucleus situated in the
central part of the cell is connected with the peripheral cytoplasm through a
number of radiating cytoplasmic strands. One or more helical or spiral,
ribbon-shaped chloroplasts, with smooth or serrated margins and containing
sevcral pyrenoids, each ensheathed with starch grains, are present in the cello
According to Dawes (1965) Spirogyra shows that: (I) the inner layer of
the cell wall as weil as the wall of the conjugation tube consists of cellulose-I
microfibrils; (2) the chloroplast contains photosynthetic bands each consist-
ing of 4-12 thylakoids, numerous starch ensheathed pyrenoids and pyrcnoid-
likc bodies lacking starch sheath (karyoids or protopyrenoids); and (3) there
are other cellular organelles such as nucleus with a complex nucleolus,
ntlmerous Golgi bodies and mitochondria, and endoplasmic rcticulum.
Some species found in freshwater streams grow attached by means of
rhizoids occurring in the form of small or large protuberances, or rods or
rosettes. In a physiological study of rhizoid formation in Spirogyrafluvia-
tilis, Nagata (1973) has observed that in laboratory cultures, rhizoid forma-
tion was restricted to the terminal cells of a filament. Out of various stimull
inducing rhizoid formation, light proved to be the most important since no
rhizoids could be initiated in the dark. Red light was much more effective in
inducing rhizoid differentiation than green or blue light. The fact that the
effect of red light was completely reversed by subsequent exposure to far-red
light indicates the involvement of phytochrome in rhizoidal differentiation.

Nuclear Cytology
Among the polycentric (having several centromeres) chromosomes there are
invariably two or more chromosomes termed nucleolar organizing chromo-
somes (Fig. 6-25 A-E) wh ich contain a distinct nucleolus organizing region
(Fig. 6-25 A). During nuclear division, the nucleoli disappear (Fig. 6-25 B-E)
by metaphase and reappear by interphase. The interphase nucleolus is highly
110 A TEXTDOOK ON ALGAE

diffcrentiated containing a nucleolar organizing region in a highly coiled form.

This region remains coated with a deeply stainable substance known as

nucleolus organizing
region
chromosome

A
".~r--······" "'''''
o
E

Fig. 6-25. Spirogyra, stages in nucleolar organization. A, coiled nucJeolar organizing

region of the chrornosorne inside the nucJeolus at interphase; B, uncoiling and contrac-
tion of nucJeolar organizing chrornosorne at early prophase; C-D, dispersal of nucleolus
at rnid and late prophase; E, disappearance of nucleolus. (After Godward, 1966.)

organizing track. The complex nucIeolus resuIts from the fusion of a num-
ber ofnucleoli depending upon the number of nuclcolus organizing chromo-
somes.

Reproduction
Vegetative. It is the most common method ofreproduction and involves
fragmentation of filaIl\ents.
Sexual. Specics showing lateral conjugation are homothaIlic whereas
those showing scalariform conjugation are generally heterothallic. In scalari-
form conjugation (Fig. 6-26 A-D) the filaments of + and - strains associate
in pairs and in each pair the side walls of some or aIl ceIls facing each other
form papillate outgrowths. The latter extend towards each other and
finally fuse thus establishing a conjugation canal. The entire protoplasmic
contents of the - type ceIl go into the + type through the canal by amoe-
boid movement (Fig. 6-26 D). When the conjugation begins, the two
filaments are in contact with each other. As the papillae of the conjugation
canal are formed and grow to form the canal, the two mating filaments are
pushed away from each other. It has been observed that during the forma-
tion of the conjugation tube, one papillate outgrowth penetrates into the
other. The dissolution of the cross wall at the junction of the two papiIIae
is affected by the formation in the cytoplasm of a Iarge number of vesicIes
or contractile vacuoles which migrate to the junction and release substances
that dissolve the cross wall (Dawes, 1965).
After the completion of conjugation one of the two fusing filaments has
 CHLOROPHYTA 111

empty cells and the other contains dark, sphcrical or oval structures, thc
zygotes. This type of sexual rcproduction involving migration of the -
gamete into the + is physiologically anisogamous even though the two fus-
ing gamet es are morphologically isogamous. In so me species, the contents of
both - and + cells move into the conjugation tube and the fusion between
gametes takes place within the conjugation canal. Such species are morpho-
logically as weH as physiologically isogamous.

c
Fig. 6-26. Spirogyra. A-C, successive stages in scalariform conjugation; D, physio-
logical anisogamy and formation of zygospore in one of the two conjugating cells; E,
part of filament showing stages in lateral conjugation; F-G, early and late stages in
germination of zygospore.

In lateral conjugation (Fig. 6-26 E), the septum at the point of its attach-
ment with either side of the lateral wall dissolves, the lateral wall bulges out
slightly thus creating a passage between the adjacent cells. Soon, the con-
tents of one cell start exhibiting amoeboid movement and finally move into
112 A TEXTBOOK ON ALGAE

the next ceIl through a narrow lateral passage. If the filaments are examin-
ed at this stage, empty ceIls are seen in the male and zygote-containing ceIls
in the female.
Rarely, so me cells of a filament may act as male and others as female
with respect to another member of a filament pair. In this case, each of
the conjugating filaments contains male as weIl as female ceIls. It has also
been reported that three filaments A, Band C (instead of two) may con-
jugate, with the central filament B behaving as male toward A and as female
toward C.
The zygospore is always a three-layered, thick-waIIed structure. It
germinates after perennation and its nucleus divides by meiosis. Three of
the resulting four nuclei degenerate and the surviving uninucleate protoplast
undergoes repeated transverse divisions giving rise to a single Spirogyra fila-
ment (Fig. 6-26 F, G).

Genus: Zygnema

Occurrence
Commonly found free-floating in ponds and streams, Zygnema is a widely
distributed freshwater alga. Sometimes it may grow submerged, living
entirely under water, attached to stones, pebbles and other objects.

Morphology
Like Spirogyra, Zygnema is also filamentous. It differs from Spirogyra mainly
in plastid morphology. CeIls of Zygnema' contain a pair ofaxile stellate
chloroplasts (Fig. 6-27), each harbouring a single central pyrenoid with

stellate chloroplast
---.-J I~I:::;::::;:;;;:::;:::::::;~I

Fig. 6-27. Zygnema, ceJls showing pair ofaxile steJlate chloroplasts.

radiating starch grains. The cells are uninucJeate and the ' single nucleus
lies embedded in the middle of the cytoplasm that separates the two chloro-
plasts. The nucleus contains one nucleolus with one nucleolar organizing
chromosome.
 CHLOROPHYTA 113

Reproduction
Vegetative. It is brought about by the fragmentation of filament.
Asexual. In donal cultures, three species of Zygnema, i.e., Z. sterile,
Z. spontaneum and Z. sp. have recently been reported to lack sexual repro-
duction, and multiply solely by the formation of akinetes (Z. sterile) or
aplanospores (Z. spontaneum and Z. sp.). Filaments resulting from aplano-
spore germination in fresh culture medium are normal and unbranched but
become branched when grown in old culture medium.

Fig. 6-28. Zygnema. A-E, stages in sexual reproduction.

Sexual. Reproduction is always morphologically isogamous. Most
species are physiologically isogamous (Fig. 6-28 A-C) but some are an iso-
gamous (Fig. 6-28 D, E). Both scalariform and lateral conjugation are
known.
-~
~
0\

Class: CHAROPHYCEAE
~
oö The Class Charophyceae inc1udes a single Order Chaniles having six Iiving
114 A TEXTBOOK ON ALGAE

genera, all placed in the Family Characeae. Popularly known as stone-

worts, the Charophyceae constitutes a distinctive and isolated group of highly
differentiated and elaborately organized green thaHophytes that-because
of their uncertain affinities-have presented greater difficulties in classifica-
tion than any other group of plants. The class is characterized by such dis-
tinguishing features as: (1) enclosure of antheridia and oogonia by jackets of
sterile ceIls; (2) biftagellate and spiral-shaped nature of the antherozoids;
(3) non-formation of zoospores; and (4) germination of zygote into a
protonemal stage which gives rise to a mature plant.
The thallus of the Characeae is differentiated into nodes and internodes
with whorls of branches of limited growth being produced from each node.
Despite these distinctive features, the photosynthetic pigments, the food
reserves, the cell wall, the ftagellar morphology and the life cycle of these
algae are typically chlorophytean. Further, a thallus differentiated into
nodes and internodes and producing branches only from the nodal cells is
also found in Draparnaldiopsis. In view of these similarities, the Charo-
phyceae are classified here along with the Chlorophyceae in the Phylum
Chlorophyta.
Besides living genera, some fossil members of the Charales have been
recorded from as far back as the Devonian. All the living members are
submerged aquatics growing attached by means of muIticellular rhizoids to
the muddy or sandy bottom of shallow, brackish or freshwater lakes, ponds
or streams. Some species of Chara become heavily encrusted with calcium
carbonate.

Order: CHARALES
Family: Characeae
Genus: Chara

Morphology
Usually 6-10 inches tall, the main axis and branches of Chara are differen-
tiated into nodes and internodes (Fig. 6-29 A). Each node bears a whorl of
several branches and consists of a pair of central cells surrounded by a peri-
pheral group of 6-20 cells. The internode is always composed of a single
elongate cell which may be corticated. The corticating cells are also long and
are derived from both the upper and lower nodes. Branched multicellular
rhizoids are formed from the lowermost one or two nodes. The branches
of the first order, produced from the nodes of the main axis, are known as
primary laterals. These are also called laterals of limited growth because
of cessation of growth in these branches after they have formed a few nodes
and internodes. Single-celled branches called secondary laterals arise from
the nodes of primary laterals. Sometimes, from near the axils of the oldest
whorls of the primary laterals (pseudoaxillary position), branches of un-
limited growth come out with the same degree of differentiation as found in
the main axis.
 CHLOROPHYTA 115

Cell Structure
The nodal cells contain den se cytoplasm, a single nucleus and a few discoid
or elliptical chloroplasts but in the internodal cells there is a large central
vacuole, hundreds of nuclei and many discoid chloroplasts, the laUer two
occupying the peripheral cytoplasmic region". The chloroplast lacks pyrenoid.

Fig. 6-29. Chara. A, part oC vegetativo thallus showing habit; B, vertical scctiOD
through apex oC thallus.

On the basis of its streaming nature, the cytoplasm is divided into a

stationary exoplasm and a rotating endoplasm which continuaUy streams
up and down along the longitudinal plane. The streaming of the cytoplaSm
is thought to result from alternating contractioil and expansion ofthe protein
fibrils which remain fixed to the cell wall.

Growth
The main axis and primary laterals grow by means of a dome-shaped apical
cell (Fig. 6-29 B) which cuts off a transverse segment at its posterior end;
the resulting segment divides transversely into an upper nodal initial and a
lower intern odal initial. The laUer does not divide further but elongates
considerably. The first division of the nodal initial is in a· vertical pl,ane
producing two semicircular cells, but all subsequent divisions occur in
curved planes, each intersecting the preceding plane of division in such a
way that a group of 6-20 peripheral cells enclosing a pair of central cells is
produced.
116 A TEXTBOOK ON ALGAE

Each peripheral cell divides into an apical initial of the primary laterai
and a basal nodal cell and, therefore, the number of peripheral cells at the
node determines the number of primary laterals produced at that node. Each
basal nodal cell further divides into an upper and a lower cortical cell. Thus
an intern odal region becomes corticated with corticating cells, one balf of
which are derived from the upper node and tbe other half from the lower node.
The nodal initials of primary laterals also exhibit the same pattern but
the apical initials of secondary laterals located at the nodes of primary laterals
do not divide further and only elongate into single-celled, scale-like structures.

Reproduction
Vegetative. Reproduction in some cases takes place by means of amy-
lum stars which are stellate aggregates of cells densely filled with starcb grains
and produced from tbe lower nodes. Other means ofvegetative propagation
are by the formation of bulbils upon rhizoids and on lower nodes of ceIls, or
by the formation of protonemal outgrowths derived from tbe nodes.
Sexual. All species of Chara show a bighly advanced type of oogamy.
The antheridial filaments are borne witbin a structure called the globule
whereas the oogonium is present within the Ducule. Both are situated at
the base of secondary laterals (Fig. 6-30 A).
A mature globule is a hollow spherical body and consists of an outer,
amiddie and an inner group of cells termed respectively as shield celIs,
manubrial celIs and capitulum cells (Fig. 6-30 B). A capitulum generalIy
divides and gives rise to a group of two to four secondary head celIs or
capitula from each of which is produced a pair of antheridial or spermato-
genous filaments (Fig. 6-30 B). Each celI of the spermatogenous filament
forms aspiral biflagelIate sperm (Fig. 6-30 C, D). In an electron microscope
study of C. cora/lina, Moestrup (1970) has reported the presence of scales
on the flagelIa of its spermatozoids. Cells of spermatogenous filaments
generalIy divide synchronously.
The nucule is an oblong structure consisting of five spirally coiled tube
cells which form a sterile envelope around the oogonium and of five-Iobed
corona cells that project beyond tbe apex of the oogonium (Fig. 6-30 A).

Development of Sex Organs

Globule. It develops from an adaxial peripheral cell at the node of a
primary lateral. This cell divides into two, the lower forms the pedicel and
the upper acts as an antheridial initial (Fig. 6-31 A). The latter divides
thrice to give rise to an octant stage. Each cell of the octant undergoes two
periclinal divisions resulting in a radial row ofthree cells (Fig. 6-31 B, C) with
the outer cell functioning as the shield ceIl, the middle as the manubrium,
and the inner as the primary head cell. Subsequently, the shield cells expand
laterally and the manubria elongate verticaIly. By this time, the primary
head cell has al ready divided and the resulting secondary head cells bifurcate
verticalIy into a pair of cells, each developing into a spermatogenous filament.
In a mature globule, the large number of spermatogenous filaments radiate
 CHLOROPHYTA 117
in the hollow cavity.
Nucule. This is also produced from a peripheral cell close to the
antheridial initial. It divides into a row of three cells, of which the lower

c
A
manubrium

B o
Fig. 6-30. Chara. A, portion of a plant beaJ;ing nuculc and globule; B, structure of a
mature globule; C, spermatogenous filament; D, biflagellate spiral sperm.

forms the pedicel, the middle the sterile jacket or sheath cells, and the upper
the oogonial mother cell (Fig. 6-31 D). The middle cell divides repeatedly
to förm a group of five sheath cells enclosing a central cell. Meanwhile the
oogonial mother cell forms a lower stalk cell and an upper oogonium. The
five sheath cells undergo a single transverse division to form a lower tier of
five tube cells and an upper tier of five coronal cells (Fig. 6-31 E). Finally
the tube cells elongate and coil spirally around the oogonium thus pushing
up the coronal cells at the top of the oogonium. A large quantity of food
118 A TI!XTBOOlC ON ALGAB

reserves accumulates in the oogonium wh ich differentiates into an egg.

At the time a nucute matures, the spiral tube cells separate from one
another and the resulting space in between them provides an opening for
the entrance of sperms, one of which finds its way inside the oogonium and
fuses with the ovum.

oogonium
tube cell

~-'d'-.t"-nodeof :":H'!:'7:,.,.-"f!-n od e 0 f
oogonium oogonium

D
FiS. 6-31. Chara. A-C. development of slobule; D-E. development of nucule.

Tbe wall ofthe zygote thickens and after perennation the oospore germi-
nates; the nucleus migrates to the anterior region where it prohably
divides meiotically. A septum is then laid down forming a uninucleate upper
cell and adegenerative, trinucleate lower cell (Fig. 6-32 A). The upper
cell undergoes one mitotic division in a vertical plane. Of the two daughter
cells, one acts as the protonemal initial and the other as the rhizoidal initial.
These initials form the filamentous protonema and the colourless rhizoids
respectively (Fig. 6-32 B), hoth with nodes and internodes. The plant arises
from one of the appendages horne on the second node of the protonema.
Because of the suspected occurrence of meiosis during germination of
zygote (and not during gametogenesis), the Iife cycIe of Chara is helieved to
be of the haploid type.
 CHLOROPHYTA 119

While studying the influence of chemical substances on the regulation of

growth of protonema and adult thalli of Chara, Imahori and Iwasa (1965)
found that thiamine, pyridoxal, gibberrellic acid and kinetin differentially

o
o
o

A
primary protonema

primary rhiZOid

Fig. 6-32. Chara. A. longitudinal scction of gcrminating oosporc showing nuclcar

division; B, surfacc view of germinating oospore. (After Smith. 1955.)

promoted the growth of the protonema whereas vitamin-Bl2 and nicotin-

amide favoured the growth of the mature thalli. Various amino acids
administered individually or in mixtures increased the growth of both proto-
nema and of adult thalli.

Phylogany
This has long been a hotly debated subject and the available fossil records
are of little help in suggesting any clues to the origin ofthis Class. In respect
of photosynthetic pigments, food reserves, cell wall composition, flagellar
morphology of spermatozoids, and zygote germination, the Charophyceae
resemble Chlorophyceae. The differentiation ofthe main filament into nodal
and intern odal ceIIs, formation of corticating filaments, production oflateral
branches oflimited growth and participation ofthe laterals of limited growth
in reproduction are features that are shared also by the chaetophoralean
genus Draparnaldiopsis. These traits have been emphasized by many phyco-
logists to suggest a possible derivation of the Charophyceae from the
120 A TEXTBOOK ON ALGAB

Chaetophorales (Desikachary and Sundaralingam, 1962). But the apical

growth and the complex nature of the male and female sex organs point
against any possible origin of Charophyceae from the Chaetophorales.
Papenfuss (1955) on the basis ofparallelism in the development ofsex organs
has emphasized an affinity between bryophytes and Charophyceae. Bold
(1967) considers that morphologically the sex organs of Charophyceae are
in no way different from those of bryophytes.

TEST QUEST/ONS
1. Give reasons for the inclusion of such morphologically diverse forms as Chlamy-
domonas, Sphaeroplea, Vlva, Frilschiella and Chara in the Chlorophyta.
2. Name the organisms responsible for 'red snow' and 'red rust of tea'. How does
the mode of nutrition of an algal symbiont in a lichen differ from that of a para-
sitic alga such as Cephaleuros?
3. Compare the different types of chloroplasts met with in Chlorophyceae.
4. Why are Chlorella and Hydrodielyon included in the Order Chlorococcales and
not in Volvocales?
S. In what waysdoes the nuclear cytology of Chlorophyta differ from that of Eugleno-
phyta?
6. "Chlorophyta comprise a group of unicellular and colonial algae full of evolu-
tionary potentialities. .. Discuss this statement.
7. Can you offer some evidences to show that in Chlamydomonas: (a) the presence oe
flagella is essential for gametic union; (b) the contractiIe vacuoles are the organs
of osmoregulation; and (c) the eye spot is probably not the primary photoreceptor
organ in phototactic responses?
8. Explain the terms: complementary sex factors; flagellar agglutination; genetic
recombination; linkage groups and tetrad analysis.
9. What will happen if the + and - strains of Chlamydomonas moewusii are: (a)
grown in a medium lacking ammonium nitrogen; (b) kept in the dark; or (c) cul-
tured in a calcium deficient medium?
10. Mention the importance oe resistance and biochemical markers in the genetical
analysis oe any organism.
11. Devise an experiment to demonstrate the recombination for non-nuclear genes in
Chlamydomonas.
12. What is cell synchrony and what advantages does it offer in physiological and
biochemical investigations ?
13. How would you prove that Volvox is more than a mere aggregation of Chlamy-
domonas-like cells, and segregation of vegetative and reproductive functions in
Volvox occurs during its embryonie stage?
14. What could be the probable functions of light, temperature, sulphur and nitrogen
in the regulation of life cycle in Chlorella?
15. Why is Chlorella a favourite organism in research es on photosynthesis and as a
potential food source?
 CHLOROPHYTA J2J
16. Compare the asexual reproduetion of Volvox with that of Hydrodiclyon.
17. Diseuss the affinities and systematic position of Chara.
18. What is a nannandrium ? In what way does the sexual reproduction of macrandrous
Oedogonillm differ from that of nannandrous species ?
19. Elucidate the genetie signilicance of the degeneration of three out of four nuclei
produced during the germination of a zygote in Spirogyra.
20. Compare the different modes of perennation met with in green algae studied by
you.
21. Do YOll think that the protonemal stage of Chara is homologous with the prostrate
system of a heterotrichous alga ?
22. Distinguish between: (a) zygospore and oospore; and (b) oogonium and arche-
gonium.
23. Describe the cell structure of Spirogyra. List live substances wh ich Spirogyra mllst
obtain from water if it is to grow vigorously.
24. Draw a labelIed diagram of a mature vegetative cell of Spirogyra. Give at least
four features in wh ich this cell differs from: (I) a cell in the meristematic zone ofa
root apex; and (2) a typical parenchyma cell in the pith of an angiosperm.
25. Pleuroeoeeus commonly occurs on tree barks and other terrestrial substrata
whereas Euglena cannot grow in such habitats. What explanation can you offer
to account for this variation?

SELECTED REFERENCES

Bold, H. C., Morphology of Plants, 2nd edn., Harper and Row, .New York (1961).
Bonner, J. T., Volvox: a colony of cells, Seient. Am., 182, S2 (1950).
Brown, R. M. (Jr.), Johnson, O. P. S. C. and Bold, H. C., Electron and phase-contrast
microscopy of sexual reproduction in Chlamydomonas moe\llusii, J. Phyeol., 4,
100-120 (1968).
Cole, K. and Akintobi, S., The Iife-cycle of Prasiola meridiona/is, Can. J. Bot., 41, 661-
68 (1963).
Dawes, C. J., An ultrastructural study of Spirogyra, J. Phycol., 1, 121-27 (l96S).
Desikachary, T. V. and Sundaralingam, V. S., Affinities and interrelationships of the
Characeae, Phyeologia, 2, 9-16 (1962).
Ettl, H., Über systematische Probleme in der Gattung Chlamydomonas, Phyeologia, 5,
61-70 (l96S).
Hase, E., Cell Division. In Lewin, R. A. (ed.) Physiology and Biochemistry of Algae,
617-24, Academie Press, New York (1962).
Hili, J. C. and Michalis, L., An ultrastructural study of the vegetative cell division in
Oedogonium borisianum, J. Phyeol., 4, 261-71 (1968).
Hoffman, L. R., Observations on the fine structure of Oedogonium. 111. Mierotubular
elements in the chloroplasts of O. cardiacum, J. Phyeol., 3,212-21 (1967).
 122 A TEXTBOOK ON ALGAB

Imahori, K. and Iwasa, K., Pure culture and chemical regulation of the growth of
charophytes, Phyc%gia, 4, 127-34 (1965).
Moestrup, 0., The fine structure of mature spermatozoids of Chara cora//ina. with special
reference to microtubules and scales, Planta, 93, 295-308 (1970).
Morimura, Y., Synchronous culture of Chlorella. I. Kinetic analysis of the life cycle of
Chlorella ellipsoidea as affected by changes of temperature and light intensity,
PI. Cell Physio/., Tokyo, 1, 49-62 (1959).
Nagata, Y., Rhizoid differentiation in Sl'irogyra. PI. Cell Physiol., Tokyo, 14,531-54
(1973).
Papenfuss, G. F., Classification of the algae. In A Century of Progress in the Natural

..
Sciences, 115-224, Proc. Calif. Acad. Sci. (1955) .
Patel, R. J. and Patel, N. J., On Fritschiella tuberosa Iyengar from Vallabh Vidyanagar
(Gujarat), Jour. Gujarat Univ., 12, 242-45 (1969).
Round, F. E., The taxonomy of the Chlorophyta. H. Br. phycol. J., 6, 235-64 (1971).
Sager, R., Chlamydomonas. Science, Wash., 131, 1460-64 (1960).
Sager, R., Streptomycin as a mutagen for non-chromosomal genes, Proc. natn. Acad.
Sci., U.S.A., 43,2018-26 (1962).
Sager, R., On the evolution of genetic systems. In Bryson, B. and Vogel, H. J. (eds.)
Evolving Genes and Proteins, 591-600, Academic Press, New York (1965).
Singh. R. N., Fritschiella tuberosa and the algal ancestry of higher plants, Proc. vrn
Internat. Bot. Congr., 17, 1-4 (1954).
Starr, R. C., Cellular differentiation in Volvox, Proc. natn. Acad. Sci., U.S.A., 59,
1082-88 (1968).
Trainor, F. R., Control of sexuality in Chlamydomonas chlamydogama, Am. J. Bot., 45,
621-26 (1958).
Whitton, B. A., Biology of Cladophora in freshwaters, Water Research, 4, 457-76 (1970).
Wiese, L., On sexual agglutination and mating type substances (gamones) in isogamous
heterothallic chlamydomonads. I. Evidence of the identity of the gamones with
the surface components responsible for sexual flagellar contact, J. Phycol., 1, 46-
54 (1965).
 7
Xanthophyta
GENERAL CHARACTERS

Commonly known as yellow-green algae, the Xanthophyta incIude one

Class, Xanthophyceae, characterized by the following general features: (I) the
photosynthetic pigments consist of chlorophyll-a and -e, ß carotene and
xanthophylls; of these, ß carotene is usually present in fairly high concentra-
tions; (2) the food reserves are oil, lipid, and a glucose polymer termed
leueosin or chrysolaminarin; (3) the cell walls when present are generally
composed of two equal oroverlapping halves, e.g., Tribonema (Fig. 7-1)

..•••
and are composed mainly of pectic substances
with smaller amounts of cellulose; in some
members the cell walls may be silicified; and
(4) the flagellated cells possess two anteriorly
•, inserted flagella of unequallength, the longer

.....
being pantonematic and the shorter acrone-
matic.
•
• • • reell wall

~.
The Xanthophyceae are widely distributed
in aquatic, terrestrial and subaerial habitats

.'.
but .the maximum representation is in fresh-
((. water habitats.
i • • ' . ~ :-chromatcphcre Tbe Xanthophyceae exhibit well-marked

•••e'
parallelism with Chlorophyceae in thallus
structure which incIude motile, palmelloid,
dendroia, filamentous and sipboneous forms.
However, tbe yellow-green algae in general do
not reacb tbe level of evolutionary develop-
ment attained in tbe green algae. Tbus, tbe
Fig. 7-1. Tribonema, portion advanced, elaborate pseudoparenchymatous
of filament showing cell wall
made of overlapping H-pieces
and parenchymatous habits met with in the
and discoid chromatophores. Chlorophyta are not seen in the Xantho-
phyta.
The usual methods ofpropagation and reproduction are vegetative,
asexual and sexual. Sexuality is rather rare and has been established in
only three genera. In Tribonema fusion takes place between two biflagellat-
ed zoogametes one of which discards its flagella and becomes non-motile
during or shortly be fore fusion, thus indicating a sort of oogamy. In Botry-
dium, both isogamous and anisogamous forms of sexuality are known.
Vaucheria is oogamous.
124 A TEXTBOOlC ON ALOAE

The formation of flagellated struetures during the life eycle of most of

the Xanthophyceae together with the oeeurrenee ofpredominantly flagellated
unicells in primitive members 01' the class, suggests a flagellated ancestry for
the Xanthophyceae.
The Xanthophyceae resemble Chrysophyceae in their flagellation, am oe-
boid habit and food reserves; and Bacillariophyceae in food reserves and cell
wall composition. The class is unique in having a preponderance of pectie
substances in the cell wall and presence of chlorophyll-e in some members.
Round (1965) and Bold (1967) consider Xanthophyceae, Chrysophyceae and
BaciIIariophyceae to be cIosely related and accordingly include them in a
single Phylum, the Chrysophyta (or Chrysophycophyta).
The vegetative cells of certain members of Xanthophyta, now segregated
into a distinct Class caIlcd Eustigmatophyceae, contain an exceptional kind
of stalked pyrenoid which is polygonal, projects from the inner face of the
chloroplast, is enveloped by flat plates of some unidentified food reserve,
and is devoid of photosynthetie thylakoids (Hibberd and Leedale, 1970).
Another distinguishing feature of these algae is that zoospores have only
one, subterminally inserted, emergent flagellum having a flagellar sweIling at
its proximal end. This flagellum has stift" hairs.

Order: HETEROSIPHONALES
Family: Botrydiaeeae
Genus: Botrydium

The most primitive genus of siphoneous Xanthophyceae, Botrydium

strongly resembles the green alga Protosiphon
in its habit and habitat.

Occurrence
It is a terrestrial alga found on muddy or
damp soils near the banks of temporary or
permanent pools, ponds and streams. It
grows as pin-head vesicles which often form
a thick, yellowish-green coating over the
soil surface.

Morphology
The thallus is unicellular, coenocytic
and consists of a lower· profusely bran-
ched, colourless rhizoidal· portion and an
upper, globose or cylindrical vesiele whieh
may measure a few mm in size. The rhi-
zoids are subterranean and serve the func-
tion of thallus anchorage. The vesicle con-
tains a central vacuole and a peripheral
protoplasm with numerous minute nuclei Fig. 7-2. Botrydium, thallus.
 XANTHOPHYTA 125

and discoid chromatophores (Fig. 7-2). The chromatophores possess naked

pyrenoids and lie next to thc cytoplasmic membrane whereas the nuclei
occupy a position internal to the chromatophores. Rhizoids are also multi-
nucleate with nuclei Iying scattered in the vacuolated and non-vacuolated
cytoplasm.
Occasionally the thalli become heavily encrusted with calcium carbonate.
The cell wall is chießy composed of cellulose. The photosynthetic food
reserves are normally oil and leucosin.

Reproduction
Vegetative. It is rare and may be brought about by budding of mature
vesicles.
Asexual. It takes place by means of biflagellate zoospores, or by the
formation of aplanospores or hypnospores. High humidity and free water
both stimulate zoospore formation (Erben, 1962). The vesicular coenocytic
protoplast fragments into uninucleate parts, each developing into a pyriform
bißagellate zoospore. The flagella are anteriorly inserted, the longer being
pantonematic and the shorter acronematic.
Generally the zoospores differentiate directly into vegetative thalli.
Sometimes their behaviour is facultative and they mayaiso act as gametes.
Under certain conditions, uninucleate or multinucleate aplanospores are
formed instead of zoospores. During adverse environmental conditions,
aplanospores transform into uninucleate or multinucIeate hypnospores.
Both uninucIeate and multinucIeate aplanospores germinate directly into
new thalli. On the other hand, the uninucIeate hypnospores behave like
aplanospores in respect of germination but the multinucleate hypnospores
produce uninucleate aplanospores or zoospores which then give rise to new
thalli.
Sexual. Biflagellate zoogametes, morphologically identical with zoospor-
es, are formed in the same way as the zoospores. The fusing gametes may be
similar or dissimilar and accordingly the sexuality is either isogamous or
an isogamous. The zygote is believed to germinate meiotically. Normally
four haploid bißagellate zoospores are produced from a germinating zygote
and each of these gives rise to a new Botrydium plant. The life cycle in-
volves a haploid multicellular vegetative phase and a diploid unicellular
zygote.

Family: Vaucheriaceae
Genus: Vaucheria

Vaucheria represents the climax of-siphoneous habit among theXanthophyta.

Asexual reproduction in this alga occurs through the formation of synzoo-
spores (compound zoospores) whereas sexual reproduction is of a specialized
oogamous type. These reproductive features together with the branched,
filamentous, siphoneous thallus, form the basis ofits placement as aseparate
monogeneric family, the Vaucheriaceae. Previously Vaucheria was included
126 A TEXTBOOK ON ALGAE

in the Order Sipbonales of Chlorophyta but the revelation of features such

as thc absence of chlorophyll-b, prescnce of chlorophyll-e, excess of caro-
tenoids over chlorophylls, storage of food reserves in the form of oil, and
the unequallength of the paired flagella of the compound zoospores, led to
its removal from green algae and inclusion in Xanthophyta.

Occurrence
Species of Vaucheria occur widely in stagnant and flowing freshwaters, in
shaded terrestrial habitats, and on damp soils and walls. The alga may
occasionally exhibit a thick, deep green, felt-like growth on moist soils.
Some species are marine.

Morphology
The thallus is generally a sparingly branched, cylindrical tube lacking cross
walls or septa (Fig. 7-3 A) except during reproduction. In terrestrial
species, anchorage to the substratum is brought about by rhizoid-like branch~
es. The thallus contains an outer cellulosic cell wall, a central vacuole that
runs continuously from one end of the thallus to the other, and a conti-
nuous layer of protoplast with the peripheral region containing many
discoid chromatophores devoid of pyrenoids. Numerous minute nuclei lie
internal to chromatophores (Fig. 7-3 B) which exhibit phototactic response,
i.e., back and forth movement in response to weak, unilateral illumination.
Whcn the light intensity is strong, they show a lateral movement, away from

Fig.7-3. Vauclreria. A, branched sipho-

neous filament with rhizoid; B, trans-
verse section of the vegetative filament
showing relative position of chromato-
phores and nuclei.

A
the light source (see Haupt 1962). In other words, the phototactic response is
regulated in such a way as to permit maximum absorption ofphotosynthetic
light by chromatophores, simultaneously protecting ihem against excessive
photodamage.
 XANTHOPHYTA 127

The thallus grows in length by simple elongation of terminal portions of

the branches.

Reproduction
Asexual. A variety of environmental factors affect zoospore formation
(see Erben 1962). High humidity, transfer from running to still water, low
light intensity or darkness, and dilution of the growth medium causing a
reduction in the concentration of nutrients, hasten or induce sporulation.
Light-controlled, diurnal periodicity in the induction of sporulation has also
been reported in V. sessilis. It has further been observed that low light
intensity enhances sporulation.
At the time of zoospore formation, apices of branches become delimited
from tbe rest of tbe thallus by a septum and each of them develops into a
club-shaped sporangium (Fig. 7-4 A). SubsequentJy, nuclci and chromato-
phores exchange their position so that the nuclei which were originally

-f.!~~vacuole

c
Fig. 7-4. Vaucheria. A, club-shaped zoosporangium; B, transection of a compound or
synzoospore showing the relative position of chromatophores and nucJei; C, liberation
of synzoospore. (Figs. A, C after Smith, 1955.)

placed internal to the chromatophores, now come to lie extern al to the

chromatophores (Fig. 7-4 B). The sporangial protoplast tb~n shrinks slightly
and a pair of flagella of somewhat unequallength develops opposite each
nucleus (Fig. 7-4 B). Both flagella seem to be acronematic. Zoospores are
formed singly witbin the sporangia. At the time of liberation of zoospore
an apical pore is formed by gelatinization of the sporangial wall and the
zoospore gently glides out through this opening (Fig. 7-4 C). The naked
] 28 A TEXTBOOK ON ALGAE

zoospores move freely for some time, settle down on a suitable substratum,
discard their flagel/u und sccrete a new wall. The germinating zoospore
forms one to several germ tubes wh ich develop into Vaucheria plant. Subs-
tances responsible for the differentiation of polarity which leads to the
appearance of the germ tube, are believed to reside in the peripheral cyto-
plasm of the germinating zoospore.
The multiflage/late zoospore of Vaucheria is regarded as a compound
zoospore or synzoospore (Fig. 7-4 B) resulting from failure of the sporangial
protoplast to segment'into biftagellate uninucleate zoospores as is the case
in many zoosporic Xanthophyceae.
Terrestrial species commonly reproduce by the formation of non-motHe
aplanospores or hypnospores. Aplanospores are formed singly in club-
shaped aplanosporangia (Fig. 7-5 A), and are liberated through apical pores
(Fig. 7-5 B). Occasionally, aplanospore formation may even be induced in
aplanosporangium.

:..... : ..,.; aplanospore

.': ~.
i
L
~ : :. .... '
l\ -"
",

\I\J
f''' :''l
\ ....
k .
(8
,
. :':-
'·····...1· :~:.

A B
Fig. 7-5. Vaucheria piloboloides. A, c1ub-shaped aplanosporangium; B, liberation of
aplanospore.

aquatic species by their transference from running to still water or from

light to darkness. During the formation of hypnospores the entire thallus
segments transversely into a large number of coenocytic cells with each
cell giving rise to a hypnospore which either germinates directly into a new
thallus or divides into a number of small cysts. When the cyst germinates,
the protoplast comes out through a pore in the cyst wall, exhibits amoeboid
movement for a while and then grows into an adult thallus.
Sexual. Sexually reproducing species of Vaucheria are either homo-
thallic or heterothallic bearing well-differentiated antheridia and oogonia.
Sex organs may be sessile or stalked. In V. geminata, a homothallic species,
sex organs are borne on special reproductive branches, producing a central
curved antheridium surrounded by a peripheral group of 3-4 oogonia. Both
antheridiaand oogonia originate as protuberances(Fig. 7-6 A) which gradual-
 XANTHOPHYTA 129

ly grow in size and accumulate a large number ofnuclei andchromatophores.

The antheridia are generally tubular with their apices slightly curved or coiled
(Fig. 7-6 B). The oogonia are commonly spherical or subspherical. At
maturity the apical, curved portion of the antheridium becomes separated
from its subtending branch by a transverse septum (Fig. 7-6 C). Its coeno-
cytic protoplast divides into a large number of uninucleate, biBagelIate
(1 acronematic, 1 pantonematic) sperms (Fig. 7-6 C) wh:ch are usually
liberated from the antheridium during early morning hours.

Fig. 7-6. Vaucheria. A, B, stages in the formation of sex organs (after Smith, 1955);
C, release of antherowids.

As an oogonium matures, all the nuclei except one migrate from the
oogonium back into the main filament. Soon a transverse septum is laid
down between the oogonium and the filament or the branch bearing it.
Subsequently it develops a receptive regifin with an opening through which
a sperm enters the oogonium.
Homothallic species produce both kinds of sex organs in elose proxi-
rnity, but generally they do not mature simultaneously thus preventing self-
fertilization. The zygotes are thick-walled structures, rich in oil, and are
commonly detaehed from the parent plant along with the oogonial walls.
'" Occasionally the zygote is liberated from the oogonial wall while the latter is
~

--
~ still attached to the parent plant. There is no cytologieal proof for the occur-
~ renee of meiosis during zygote germination, and no definite knowledge
; about the fate of the zygote. However it is believed that zygotic germina-
~ tion in Vaucheria is meiotic, and it direetly gives rise to a new thallus. The
130 A TEXTBOOK ON ALGAE

life cycle of Vaucheria seems to resemble that of Botrydium but the cytological
proof is lacking.

TEST QUEST/ONS

1. Which practical methods will you adopt to distinguish between: (1) Protosiphon
and Botrydium; (2) Tribonema and Microspora; (3) Vaucheria and Dichotomosiphon;
and (4) Ophiocytium and Hydrodictyon 1
2. ExplalD with reasons whether the thalli of Botrydium and Vaucheria are multi-
cellular, acellular or unicellular.
3. Compare the zoospores of Vaucheria with those of Oedogonium.
4. What could be the reasons for the zoospores and sperms of Vaucheria to be res-
pectively multiflagellate and biflagellate 1
S. Describe the characteristics of xanthophytes generally and of Heterosiphonales
specifically. Name representative genera of the Heterosiphonales and discuss their
salient features and interrelationships.
6. Do you think that the frequent occurrence of two-piece silicified cell walls in some
members of the Xanthophyceae has any phylogenetic significance 1
7. Why is starch not found around the pyrenoids of Xanthophyceae 1
8. Enumerate the features that have been employed in the classification of Xantho-
phyceae.

SELECTED REFERENCES

Bold, H. C., Morphology of Plants, 2nd edn., Harper and. Row, New York (1967).
Erben, K., Sporulation. In Lewin, R. A. (ed.) Physiology and Biochemistry of Algae,
701-707, Academic Press, New York (1962).
Haupt, W., Intracellular movements. In Lewin, R. A. (ed.) Physiology and Biochemistry
of Algae, 567-71, Academic Press, New York (1962).
Hibberd, D. J. and Leedale, G. F., Eustigmatophyceae-a new algal class with unique
organization of the motile cell, Nature, Lond., 225, 758-60 (1970).
Round, F. E., The Biology of the Algae, Edward Arnold (publishers), London (1965).
 8
Bacillariophyta
Tbe Bacillariopbyta, popularly called diatoms, comprise a bomogeneous
assemblage of uniceJ1ular and colonial forms tbat differ generally from otber
algae in possessing bigbly sculptured and symmetrically ornamented cell
walls. Tbe main cbaracters tbat collectively differentiate tbe Bacillariopbyta
from otber pbyla are: (I) tbe diploid nature of vegetative cells; (2) tbe
presence of cbloropbyll-c togetber witb fucoxantbin, diatoxantbin and
diadinoxanthin in chromatophores, the usual brown colouration being due
to the preponderance of carotenoid pigments over chlorophylls; (3) the
siIicified nature of cell waUs wbicb consist of two bigbly perforated over-
lapping pieces sometimes bearing structures Iike spines and bristles; (4) the
storage of oil and chrysolaminarin and not starch in food reserves; and
(5) the reduction in size of cells occurring during vegetative multiplication
compensated by the production of spores known as auxospores.

OCCURRENCE

Cosmopolitan and ubiquitous in distribution, the diatoms form a major

component of the planktonic vegetation. They are very important as
primary producers in the food web of aquatic ecosystems. The most common
genera of freshwater habitats are Synedra, Melosira. Nallicula. Nitzschia
and Asterionella. Species of Cocconeis, Gomphonema and Eunotia grow
epiphytically on certain freshwater algae such as Cladophora and Oedogonium.
Triceratium and Hyalodiscus occur in the littoral and sublittoral zones as
epiphytes on seaweeds.

MORPHOLOGY

Tbe tballi are eitber unicellular or colonial. In respect of their sbape and
valve morpbology the unicellular diatoms bave been classified into two orders,
tbe pennate diatoms (Pennales) with isobilateral symmetry. e.g., Pinnularia
(Fig. 8-1 A) and tbe centric diatoms (Centrales) witb radial symmetry. e.g.,
Cyclotella (Fig. 8-1 B). Triceratium (Fig. 8-1 C) bas three planes of mirror
symmetry. Furtber classification of tbe Centrales is based on the presence
or absence of bristles or borns on tbe cell surface. Tbe Pennales are classi-
fied according to tbe presence or absence and number and morpbology of
tbe raphes on tbe valves.
Colonial diatoms may be organized into uniseriate filaments, as in some
species of Melosira. Or, tbere may be extensive brancbing ofthe mucilage
132 A TEXTBOOK ON ALGAE

with the stalks bearing at their free ends groups of cells organized into fan-
like structures as in Licmophora ftabellata. Sometimes a colony results

central
nodule

Fig. 8-l. A, Pinnularia, valve view; B, Cyclorella; C, Tricerarium.

from the linkage of the cells through bristles or horns present on the valve
surface as in Chaetoceros. Stellate colonies result from the union of cells
at their basal ends through localized production of mucilage as in Asterio-
nella.

CELL STRUCTURE

Cell wall (frustule) consists of two overlapping halves, the upper half is
known as epitheca and the lower hypotheca. Each theca is further divided
into two parts-the main surface and its incurved margins termed valve
and connecting band, respectively (Fig. 8-2 A, B). The epitheca and
hypotheca can be compared to a box, the lid corresponding to the former
and the main body to the latter. The two connecting bands represent in-
curved sides of the lid and the main body whereas the valve relates to the
top or bottom of the box. When fitted together, the connecting band of
epitheca overlaps that of hypotheca and the two bands remain united in the
overlapping region called girdle, bya cementing organic substance present
between them. Accordingly, a cell can be seen from two different aspects,
the falfe fie" and the girdle fie". Most diatoms appear rectangular in girdle
 BACILLARIOPHYTA 133

view but in valve view their shape is variable. The line connecting the
middle of the two valves constitutes the pervalvar axis, and the plane along

epltheca
1 hypotheca

I-connecting
band

Fig. 8-2. Plnnularla. A, transverse

section showing relative position of
epitheca, hypotheca, connecting
band, valve and raphe; B; girdle
view showing relative position of
epitheca and hypotheca.
conr.ccting band
B
which the cell divides (at right angles to the pervalvar axis) is called the
valvar plane (Fig. 8-3). The ornamentations are generally confined to the
valve portion of the silica wall.
apical plane
~------

valvar
plane

transapi;:al plane

Fig. 8-3. Diagram iIIustrating the three planes of symmetry in a pennate diatom. (After
Reimann et al., 1966.)

Tbe diatom cell walls are made up of crystalline silica (ot-quartz) and an
organic component of unknown nature. Silica is an absolute requirement
for the multiplication of diatoms and the cytoplasmic membrane is consider-
134 A TEXTBOOK ON ALGAE

ed instrumental in the formation of silicified wall (Lewin, J962). In Navicula

pelliculosa the wall consists of a silica sbell or layer
witb an outer coating of organic material termed
organic skin (Reimann et al., J966). The formation
of silica shell is intracellular and occurs by deposi-
tion of siIica inside the vesicles enclosed by a three-
layered membrane, the silicalemma (Fig. 8-4).
Apart from siIicalemma, there is a more or less
thick layer of acid polysaccharides lining the siIica
shell from inside. SiIicic acid uptake has been
studied in certain pigmented and colourless diatoms.
It involves some cell surface factors, the absence of
which impairs the process of silicic acid absorption.
The composition and origin of organic skin is still
unknown.

VALVE MORPHOLOGY
The fine lines or markings found on tbe surface of
diatom valves vary enormously in their microscopic
details, so much so that in the past they created
much confusion in the formulation of terminology.
However, according to Hendey (1959), four types
of secondary structures are present on diatom Fig. 8-4. Navlcu/o, graphie
representation oC silica-
valves: (1) tbe punctae, wbich are fine perforations lemma formation. (After
arranged in regular rows corresponding to the Reimann et 0/., 1966.)
markings or striae on tbe valve surface of silica
walls; (2) the areolae, which are cavity-like depressions, coarser and larger
than the punctae and are generally provided with sieve membranes; (3) the
canaJicuJi, which are tubular canals running through the valve surface; and
(4) the costae, which are specially thickened regions of the valve resulting
from heavy accumulation of siIica and represent the valvar ribs. The ribs
constitute the backbones of the cell wall. Hendey has also c1assified the
diatom walls into two types based on their nature: (1) the laminar wall, a
single silicified layer; and (2) the locular wall, basically made up of two
parallel wall layers with a number of loculi in between them.
In pennate diatoms the markings are arranged longitudinally, one on
either side of the eIIiptical or oblong valves, e.g., Pinnularia (Fig. 8-1 A)
whereas in centric diatoms they are distributed concentrically with reference
to a central point on the more or less circular valves, e.g., in Cyclotel/a
(Fig. 8-1 B).
In Pinnularia which is an oblong unicell, the valve view presents two
planes of iso bilateral symmetry-the apical and the transapical. The apical
axis joins the two poles of the valvc and the transapical axis passes along
the transverse axis of the valve. In the axial area of some pennate diatoms,
e.g., Navicula, there is a longitudinal slil termed raphe wh ich is interrupted
 BACILLARIOPHYTA 135

in the middle by a central nodule formed by internal thiekening ofthe valve

(Fig. 8-5). Eaeh pole of the valve also contains a nodule (polar nodule)
whieh is formed in the same manner as the central nodule (Fig. 8-5). The
centrie diatoms laek a raphe.

Fig. 8-5. Navicula, valve view iIlustrating details of cell wall structures. (After
Reimann et al., 1966.)

Navicula, Iike Pinnularia, is isobilaterally symmetrical with valve mark-

ings consisting offine pores bridged from inside by sieve membranes eontain-
ing numerous fine pores. The raphc situated in the eentral rib is also ealled
apical rib. The rows of pores situated on the lateral sides of apical rib
separated from one another by solid siliea ribs are termed transapical ribs.
Tbe pores within a row are separated by siliceous connections whieh link
the adjacent transapical ribs (Fig. 8-5). In valves whieh lack a raphe, the
axial area may give the appearance of a false raphe or pseudoraphe. Navi-
cula and Pinnularia possess two raphes eaeh, one on eaeh valvc, whereas in
Achnanthes only one of the two valves has a raphe. A pseudoraphe is
charaeteristie of Synedra and Fragilaria.

PROTOPLAST

The cytoplasm of thc diatoms is bounded by a membrane and surrounds a

central vacuole. The cells are uninucleate. In the Pennales the nucleus
in most species lies in a massive plasmatic bridge which separates the
vacuoles; in the Centrales it normally occupies a position within the peri-
pheral laycr of the cytoplasm. Nuclear division shows two interesting
features: (I) the spin die formation is intranuclear and the metaphase
chromosomes arrange themselves in an equatorial ring rat her than on an
equatorial plate, and (2) the spin die is an elongated eylinder with ftattened
ends, a feature common to Chrysophyta and Pyrrophyta. In Lithodesmium
undulatum certain plate-like struetures constituting a spin die precursor are
found along the nuclear membrane. During mitotic prophase in sperm-
producing ccIIs, a mitotic spindie consisting of long microtubules appears.
136 A TEXTBOOK ON ALOAE

between precursor plates and nuclear wall. Gradually the nuclear wall
degrades and a compact spindIe is formed. Subsequent stages in nuclear
division are similar to those in higher plants. The Centrales generally have
numerous discoid chromatophores and the Pennales have one or two large
plate-like frequently lobed chromatophores. Naked pyrenoids occur in
some diatoms, e.g., Nitzschia. Electron microscopy of the diatom cell has
demonstrated the presence of endoplasmic reticulum, GoI gi bodies and
mitochon dria.

LOCOMOTION
Pennate diatoms possessing a raphe exhibit a gliding movement whose
mechanism is not clearly understood. The protoplasm exhibits streaming
and, believably, as a result the cytoplasm comes in contact with the extern al
medium through the raphe. The friction caused by such contact brings
about the movement of the cello It has been suggested that the osmotic
pressure ofthe cell, which is equivalent to about 4-5 atmospheres, is sufficient
to force the entry of the cytoplasm into the complex canal systems of the
raphe thus causing. the streaming of the cytoplasm through the raphe. The
movement is generally jerky but at times it may be creeping and steady. The
gliding movement or cytoplasmic streaming has been explained by the
hypothesis that protein fibrils of the protoplasm attached to its outer surface
undergo alternate contraction and expansion. If the fibrils are on the inner
surface, their beating will cause the peripheral cytoplasm to stream and ifthey
are located in the outer surface, their beating will result in gliding movement.
Recent researches have established that cytoplasmic streaming, associated
with the raphe system, plays an important role in the gliding motion of
diatoms. The motiIe ceIIs possess a fibrillar system under the raphes, and
certain crystalline granules in the cytoplasm that produce some mucilagi-
nous substance which is secreted through the raphes, sticks to the substratum
and, during locomotion, is left behind as a traiI. Some capiIIarity mechanism
which causes apressure gradient along the raphe, based on the secretion
of the mucilaginous substance, possibly provides the motive force for locomo-
tion in gliding movement.
Tbe problem of floatation and sinking of planktonic diatoms has also arous-
ed considerable interest in recent years. One theory proposes that accumula-
tion of oi! and lipids in old nitrogen-deficient diatoms gives the necessary
buoyancy. This has been refuted on the basis that increase in thickness ofthe
silica wall paralleIs the accumulation of fats and this offsets the reduction in
specific gravity due to the fat. An alternative suggestion has been made
by Gross and Zeuthen (1948) who consider that by excluding metal ions
from their cell sap the diatoms decrease their specific gravity and hence
keep floating. Lund (1959), however, contradicts the presence of such a
mechanism in freshwater diatoms since their habitats are so low in ionic
concentration that loss of ions from the cell sap may not be of much help
in maintaining the buoyancy.
 BACILLARIOPHYTA 137

REPRODUCTION

Tbere are well-documented evidences for the occurrence of vegetative and

sexual reproduction in diatoms. Although the production of non-motile or
biflagellate zoospores for some members of the Centrales, e.g., Biddulphia,
has been elaimed, there is still some doubt as to whether the spores are
actually formed by the diatoms, or whether the supposed spores are in fact
only chytrids that parasitize the diatoms. If indeed they are spores, their
fate is unknown.

VEGETATIVE

The two tbecae of tbe parent cell sligbtly gape apart before division. Tbe
nucleus divides mitotically and karyokinesis is followed by cytokinesis of
the protoplast into two uninucleate parts along tbe valvar plane. Soon,
new valves and later connecting bands are deposited along the freshly
formed protoplasmic surfaces. Tbe parental hypotheca serves as epitheca
of one of the two daughter ceIls, wbereas the parental epitheca remains as
epitheca of the other daughter cell. Accordingly, the newly formed wall
pieces always serve as bypotheca. Such a division leads to the production
of two daughter cells of unequal size; the cell containing the parental
hypotheca.as epitheca is always smaller than its si ster cell by about twice
the thickness of the connecting band. Thus the progenies of a diatom
become progressively sm aller during successive cell divisions.

SEXUAL

In tbe Pennales an enlarged spore, termed auxospore, develops from the

zygote resulting from the union of two amoeboid garnetes. In the Centrales,
however, fusion of a ftagellated sperm occurs with a non-motile egg.
Consequently, sexuality in the former is generally isogamous and that in the
latter oogamous. Since the zygote enlarges to becorne an auxospore, the
production of auxospores is a compensatory measure för the vegetatively
reproducing diatoms to regain their normal size.
In the Pennales, two cells come to lie very elose with their girdle view
facing each other. A secretion I)f mucilage then envelops the conjugating
pair. The diploid nueleus of the copulating cells undergoes meiosis and
produces 3 or 4 haploid nuelei of which generally one or two form gametes
and the others degenerate. The gamet es move out of the parent cells and
gametic union takes place in the mucilaginous envelope (Fig. 8-6 A-G). If
each cell of the copulating pair gives rise to one garnete a single zygote
results, when two gametes are produced from each partner then two zygotes
are formed. Sometimes, physiological anisogamy occurs; in this case one
garnete is mobile and the other stationary, one zygote being forrned within
each mernber of the copulating pair.
In Centrales, the cell functioning as female slightly extends with its
138 A TEXTBOOK ON ALGAE

elongated nucleus to become an oogonium. Two meiotic divisions of the

oogonial nucleus take place and at each division one of the daughter nuclei
degenerates so that the protoplast contains a single haploid nucleus which

M N o p Q

Fig. 8-6. A-G, Cocconeis, stages in auxospore formation. A-C, meiosis and degenera-
tion of all nuclci cxcept one in each frustule; D, fusion of gametes; E, zygote; F-G,
dcvelopment of auxospore from zygote. H-Q, Afe/osira, auxospore formation in a
centric dia tom, H-L, formation of antherozoids; M-Q, stages in fertilization and
formation of auxospore. (After Smith, 1955.)

u!timately becomes the cgg nuc1eus. The cell acting as spermatogonium

divides meiotically to produce four uniflagellate spermatozoids. In Cyc!o-
tella tenuis/riata, a vegetative ceIl functions directly as a spcrmatogonium
 BACILLARIOPHYTA 139

but in Melo$ira varians a vegetative cell divides mitotically to producc 4-8

spermatogonia (Fig. 8-6 H-L).
At the time of fertilization, spermatozoids gather around an oogonium
and one of them fuses with the egg. The zygote escapes out of the female
cell, enlarges to a characteristic size and becomes an auxospore which secretes
a new wall and is now a diploid vegetative cell (Fig. 8-6 M-Q).
Schultz and Trainor (1968) have recorded uniftagellate and biftagellate
spermatozoids, and auxospores in cultures of Cyc/otella meneghiniana and
C. cryptica and found that spermatogenesis and auxospore formation are
stimulated with increasing concentrations of sodium. Both light and
temperature have been reported to regulate sexuality in some diatoms.

PHVLOGENV
Because of the siliceous nature of the diatom cell walls, weil preserved fossils
of Bacillariophyta are available. The Centrales have been reported from the
Jurassie and the Pennales from early Tertiary. Tbe fossil evidence, therefore.
suggests that the Centrales are more primitive from which the Pennales
rnight have originated. Tbe fact that most ofthe centric diatoms are marine
planktonic forms in contrast to the Pennales which are predominantly fresh-
water, also indicates a centric ancestry for the Pennales.
The presence of fucoxanthin and chlorophyll-a and chlorophyll-c links
the Bacillariophyta with Phaeophyta, and the characteristic food reserves
relate the diatoms to Xanthophyta and Chrysophyta.

TEST QUEST/ONS

1. What will happen when you attempt to grow diatoms in a nutrient medium with
or without silica ?
2. How can you establish that silicon uptake in diatoms is an energy requiring process ?
3. What reasons can you offer to account for the progressive reduction in the size of
vegetatively reproducing diatoms ?
4. Why do some algologists assign diatoms the rank of a Phylum whereas others group
them as one Class in the Phy)um Chrysophyta ?
5. Out of tbe Pennales and Centrales, which one do you consider more primitive and
why?

SELECTED REFERENCES

Gross, F. and Zeuthen, E., The buoyancy of plankton diatoms: a problem of eell physio-
logy, Proc. Roy. Soc., 1358, 382-89 (1948).
Hendey, N. 1., The strueture of the diatom eell wall as revealcd by the eleetron miero-
scope, J. Quekett Microsc. Club,5, 147-75 (1959).
140 A TEXTBOOIC ON ALGAE

Lewin, J. C., Silicification. In Lewin, R. A. (ed.) Physiology and Biochemistry of Algae,

445-53, Academic Press, New York (1962).
Lund, J. W. G., Buoyancy in relation to the ecology of the Creshwater phytoplankton,
Br. Phycol. Bull., 1, 1-17 (1959).
Reimann, B. E. P., Lewin, J. C. and Benjamin, V. E., Studies on the biochemistry and
fine structure of silica sbell formation in diatoms. 11. Tbe structure of tbe cell wall
of Navicula pelliculosa (Breb.) Hilse, J. Phycol., Z, 74-84 (1966).
Schultz, M. E. and Trainor, P. R., Production of male gametes and auxospores in tbe
centric diatoms Cyclolella meneghiniana and C. cryptica, J. Phycol., 4, 85-88
(1968).
 9
Phaeophyta
Tbe Phaeophyta (brown algae) are characterized by five chief distinguishing
features: (I) the photosynthetic pigments include chlorophyll-a and chloro-
phyll-c, ß carotene, fucoxanthin, violaxanthin, diatoxanthin and other
xanthophylls, and in general there is an excess of carotenoid over chlorophyll
pigments; fucoxanthin is present in sufficient quantity to mask the green
colour of chlorophylls and to impart its own brown colour to these algae;
(2) the excess photosynthate is commonly stored in the form of laminarin
and mannitol, rarely as fat droplets; (3) certain whitish granules, called
fucosan vesicles, are usually present in the cell; (4) the cell wall is composed
of cellulose, fucinic acid and alginic acid; and (5) the ftagellated structures have
a pair of laterally inserted unequal ftagella, of which the larger one is anterior
and pantonematic whereas the sm aller one is posterior and acronematic.

OCCURRENCE
With a few exceptions, e.g., Pleurocladia, Heribaudiella, Bodanella and a
few species of Ectocarpus which are freshwater forms, all other brown algae
are marine. They generally inhabit the colder waters but members of the
Encoeliaceae (Ectocarpales), Dictyotales and Sargassum are markedly warm-
water forms.
Depending on the topographical features of the land-sea boundary, the
littoral region is divided into three major zones: sublittoral-always sub-
merged with an approximate maximum depth of 100 metres; intertidal-
submerged during high tide and exposed during low tide; and supralittoral
-not very well-defined, and beyond which the highest tides or tide sprays
rarely reach. According to the nature of the shore, tides and algal associa-
tions, the intertidal belt is further divided into three minor zones: the upper,
middle and lower.
The sublittoral zone inc1udes thick forests of such algae as LAminaria,
with subftora of Dictyota, Cutleria, Alaria, Himamlzalia and Desmarestia.
In the intertidal belt the rockweeds (Fucaceae) grow in the upper zone,
Co!pomenia, lyengaria, Sphacelaria, Leathesia and others in the middle zone
and most of the Laminariales in the lower zone. Species of Ectocarpus form
the subftora of the supralittoral zone.

RANGE OF THALLUS ORGANIZATION

The thalli range in structure from heterotrichous filamentous types through

142 A TEXTBOOK ON ALGAE

pseudoparenchymatous uniaxial forms, to true parenchymatous forms.

Neither unicellular nor simple filamentous brown algae are known and the
simplest thallus organization in this Phylum is the branched, heterotrichous
filament.
The vegetative body of the Laminariales and Fucales shows well-marked
morphological and anatomical differences. In Laminaria the thallus is
morphologically distinguishable into hold fast, stipe and blade. A trans-
verse section of the stipe or blade shows the distinction of vegetative tissue
into three regions: an outer epidermis, amiddie zone of several-Iayered
cortex, and multicellular central medulla. Some cells of the medulla pro-
duce vertically elongated 'hyphae', known as trumpet hyphae, that lack
chloroplasts and discharge the function of water and nutrient conduction.
Nereocyslis and PelagophycllS (Laminariales) have certain cells that are simi-
lar to the sieve tubes of higher plants but are devoid of companion cells
(Parker and Huber, 1965).
The vegetative thallus usually grows by intercalary or apical meristems.
Trichothallic growth is characteristic of most Ectocarpales, Desmarestiales,
Cutleriales and Sporochnales. Intercalary growth characterizes the Lami-
nariales in which the meristem is located at the junction of the stipe and the
blade. The Sphacelariales, Dictyotales and Fucales show apical growth.

CELL STRUCTURE

The cell wall is made up of an outer mucilaginous layer containing alginic

and fucinic acids, and an inner cellulosic layer.
The cytoplasm generally contains many small vacuoles and a few refractile
bodies of an unknown nature termed as the fucosan vesieles'which are usually
present at the site of high metabolie activity. The chromatophores are
mostly parietal, frequently with more than one chromatophore per cell
except in Pylaiellafllh'escens which contains axile stellate chromatophores.
The Ectocarpales show wide variation in the shape of their chromatophores
which may be platc-like, ribbon-shaped or discoid. Projecting pyrenoids
laeking photosynthetic thylakoids are present in some members of the
Ectocarpales, Sphace!ariales and Dictyosiphonales. The chromatophore
lamellations in the Phaeophyta result from a rather elose aggregation of
generally three, rarely four, thylakoids that do not cohere into stacks or
bands but run parallel to each other (Evans, 1966). In the biflagellate cells,
the two basal granules, each with one flagellum, are laterally connected and
in addition the anterior flagellum and the nueleus are also joined together
through a fibrous structure comparable to a rhizoplast. Dietyota has
monoflagellate sperms with two basal granules of wh ich the posterior one
does not give rise to a flagellum (Manton, 1965). The phaeophycean
swarmers are commonly equipped with an eye spot and a chromatophore
devoid of pyrenoids.
The cells are uninucleate, with ODe or two nucIeoli. Some members,
e.g., Fucus, have in their resting nucleus many Feulgen-positive bodies called
 PHAEOPHYTA 143

chromocentres which arrange themselves Iinearly along the chromosome

during late prophase but their identity is lost with the progressive contrac-
tion of chromosomes during metaphase; this state is maintained until the
next interphase and prophase. The chromocentres are characteristic of
only the Phaeophyta, but nothing is known of their function. Another
feature is the presence of centrosomes at the poles of a dividing nucleus.
The chromosome organization has advanced to such an extent that mem-
bers of the Laminariales possess weIl differentiated X and Y sex chromo-
somes. Other subcellular organelles such as mitochondria, Golgi bodies, and
endoplasmic reticulum are also present. Synaptonemal complexes have
recenUy been reported in the meiotic nuclei of a few brown algae.

REPRODUCTION

VEGETATIVE
The commonest method is by fragmentation of a thallus into two or more
parts, each of which regenerates into a mature new plant. In the Sphace-
larialesspecial reproductive branches known as propagules are formed which
after detachment from the parent, give rise to ncw plants.

ASEXUAL
Except Tilopteridales, Dictyotales and Fucales, in all Phaeophyta zoospores
are produced in well-defined sporangia borne on the sporophyte. The
Ectocarpales and Sphacelariales produce unichambered or unilocular as
weil as multichambered or plurilocular sporangia. During the formation of
zoospores, the diploid nucleus of the unilocular sporangium divides by meio-
sis, followed by aseries of mitotic divisions of the rcsulting four haploid
nuclei. Finally the cytoplasm becomes segmented into a number of uni-
nucleate portions each of which subsequently acquires a pair of latcrally
inserted flagella. The zoospores produced from unilocular sporangia are
haploid and on germination give rise to haploid, gametophytic plants.
In plurilocular sporangia, the nuclei never divide meiotically and there-
fore the zoospores are always diploid and serve as an accessory means of
perpetuation of the sporophyte or the gametophyte. Other zoosporic
Phaeophyta bear only unilocular sporangia. In Dictyotales the sporophyte
forms tetrasporangia, and reduction division of the sporangial nucleus in
these results in the formation of four haploid, uninucleate aplanospores or
tetraspores. Tbe Tilopteridales form single, quadrinucleate aplanospores,
termed monospores, in each sporangium. Both tetraspores and monospores
give rise to gametophytic plants. The Fucales lack asexual reproduction.

SEXUAL
Gametopbytes bear only multicbambered organs, tbe plurilocular game-
144 A TEXTBOOK ON ALGAE

tangia, whieh form gametes that resemble zoospores in morphoIogy. The

Eetoearpales and Sphaeelariales are mostly isogamous but the Puneta-
riaeeae, Cutleriales and possibly Tilopteridales are distinctly anisogamous.
Several brown algae, e.g., the Fueales, Laminariales and Dictyotales, are
oogamous. In Dietyotales, the antheridia and oogonia oeeur in sori but in
Fueales they are produeed in eavity-like depressions (conceptacles) on a
fertile blade (receptacle).
A marked lunar periodieity in sexual reproduetion has been observed
in eertain marine brown algae. The ova are released around the first
quarter of the moon but there is no such rhythmieity for sporogenesis.

LlFE CYCLE AND ALTERNATION OF GENERATIONS

Three chief types of lire eyde-isomorphie, heteromorphie and diplontic-
are found in brown algae.
The Eetoearpales, Cutleriales, Tilopteridales, SphaeelariaIes and
Dictyotales show an isomorphie life cyde in whieh both the sporophytie
and gametophytie generations are morphologieally similar. The Chorda-
riales, Punetariales and Dietyosiphonales, incIuded in the Order Eetoearpales
by Fritseh (1945), as weH as the Sporoehnales, Desmarestiales and
Laminariales, aH exhibit heteromorphie alternation between a mieroseopie
filamentous gametophyte and an elaborate, often maeroscopie, sporophyte.
In Chordariales the zygote germinates into a filamentous strueture, desig-
nated as protonema if it generates the sporophyte as a lateral outgrowth,
and as plethysmothallus if it forms zoospores in pluriloeular sporangia for
its own propagation. In the Fueales the plants are diploid and laek an
alternation of generations. The only haploid stage in the lire eyc1e is eonfined
to the gametes. There is no independent, free living multieellular gameto-
phyte in the Fueales.

CLASSIFICATION

Phaeophyta indudes the single Class Phaeophyceae. On the basis of vege-

tative organization and sexual reproduetion, Fritsch (1945) divided the
Phaeophyceae into nine orders: Eetoearpales; Tilopteridales; Cutleriales;
Sporoehnales; Desmarestiales; Laminariales; Sphacelariales; Dietyotales; and
Fucales.
Most other phyeologists emphasize the three types of life eyele, and
divide the Phaeophyta into three cIasses (or subcIasses within the Class
Phaeophyeeae): Isogeneratae (incIuding isomorphie forms); Heterogeneratae
(eomprising heteromorphie forms); and CycIosporeae (constituting diploid
forms). Within the Isogeneratae and Heterogeneratae, genera have been
plaeed in different families, and families in different orders on the basis of
differenees in the nature of zoospores, sexuality, organization and growth of
thallus.
 PHAEOPHYTA 145

PHYLOGENY

The Class Phaeophyceae comprises a group of multicellular algae some of

which show a complex morphological and anatomical differentiation not
seen in other aIgaI phyJa. The hcterotrichous, uniaxial and multiaxial
types of thalli met with in the Chlorophyta, Phaeophyta and Rhodophyta
probably represent the parallel course of evolution that has taken place
independently in the three phyla. The Phaeophyceae resemble BaciIIario-
phyceae, Cryptophyceae and Pyrrophyta in having ehlorophyll-c; and
Chrysophyceae and BaeiIIariophyeeae in having xanthophylls, fucoxanthin
and neofueoxanthin. The three phyla Phaeophyta, Pyrrophyta and Chryso-
phyta (Class Baeillariophyeeae in particular) probably belong to the same
evolutionary line, showing predominanee of brownish earotenoids over the
yellow, and of earotenoid pigments over chlorophylls.
Oil and saturated fats are common food reserves in Xanthophyceae,
Chrysophyceae, BacilIariophyceae and Phaeophyceae. In addition, laminarin
is a distinetive reserve product of the Phaeophyta, and is ehemically similar
to leueosin. The BaeiIlariophyceae, however, differ from Phaeophyceae
in having eomplex cellular morphology, with a two-pieee silieified wall.
Both Xanthophyceae and Chrysophyceae resemble Phaeophyceae in
their flagellar morphology hut the presence of two-pieee silieified wall and
chlorophyll-e in certain members of the Xanthophyeeae are two features
unknown in Phaeophyeeae.
Evolution is eonsidered to have progressed along two different lines
within the Phaeophyceae: one leads to isomorphie alternation of generations
aeeompanied by advanee of sexuality from isogamy to oogamy and, in
vegetative organization, [rom simple to eomplex thalli; the other with
heteromorphie alternation of generations shows a similar progression in
reproduetion and vegetative organization. The diploid Fueales are assumed
to represent the evolutionary elimax in the heteromorphie series, with the
gametophyte reduced to the single-cellcd stage, i.c., the gamete.

Order: ECTOCARPALES

It has been differently delimited by various phyeologists. Fritsch (1945)

included in this order filamentous, pseudoparenehymatous and parenehyma-
tous types derived from a heterotriehous filament. Emphasis on the hetero-
triehous eharaeter led to the inclusion of [orms exhibiting isomorphie as
weIl as heteromorphie alternation of generations in the order. Most other
phycologists have, however, eonsidered the life eycle of greater importanee
than the heterotriehous habit and have eonsequently raised those famiIies
~ in the Order Eetoearpales sensu Fritseh that exhibit heteromorphie alter-
@. nation, to the rank of orders. This has led to the restrietion of the Eeto-
~ earpales to the single Family Ectocarpaceae charaeterized by simple, hetero-
~
'-' triehous members exhibiting isogamous sexuality and isomorphie alternation
g of generations.
146 A TEXTBOOK ON ALGAE

In general, the prostrate system of Ectocarpaceae grows by an apical

meristem but the development of the erect system is highly variable-Ecto-
carpus siliculosus shows diffuse growth, E. lucifugus apical growth, and E.
paradoxus trichothallic growth. Pylaiel/a, an alga related to Ectocarpus,
frequently shows longitudinal septation in its filaments as a result of which
the thallus tends to become parenchymatous. Streblonema lacks an erect
system.

Family: Ectocarpaceae
Genus: Ectocarpus

Occurrence
Worldwide in distribution, Ectocarpus grows in the littoral and supralittoral
regions. In India it is mostly found in the supralittoral zone along the
West and East Coasts.

Morphology
The thallus is made up of uniseriate filaments and is generally differentiated
into a much branched erect system (Figs. 9-1, 9-2 A) and a prostrate sys-
tem that is sparsely or profusely branched. Each branch generally arises
from just below a septum. The
main branches and their branch-
lets most often terminate in hairs
consisting of numerous elongated,
tapering, hyaline and much vacuo-
lated cells. The plants are at-
tached to the substratum by bran-
ched rhizoids produced from
lower cells of Iower branches and
in some cases these descending
rhizoids form a corticating layer
around the lower cells of the main
axis.
The cells are uninucleate and
have an irregular band-shaped,
or many discoid chromatophores
with smooth margin (Fig. 9-2 B).
The chromatophore contains a Fig. 9-1. Ectocarpus, photomicrograph
showing habit and plurilocular sporangia.
projecting pyrenoid from which
a new pyrenoid may develop by budding (Evans, 1966).
The growth and nutrition of Ectocarpus con/ervoides have been studied
in unialgal, bacteria-free cultures (Boa1ch, 1961). The alga, which is obli-
gately photoautotrophie, remains viable for over a year in light, but for
only about three months in dark. Potassium nitrate, potassium phosphate,
manganese chloride and a mixture of organic substances stimulate its growth
in natural seawater medium in light.
 PHAEOPHYTA 147

Reproduction
Asexual. The sporophyte produces unilocular as weil as plurilocular
sporangia (Figs. 9-1, 9-3 A) terminally and singly on sm all branchlets.
During the development of a unilocular sporangium, the terminal cell of a
branchlet enlarges considerably, assumes a globose or ellipsoid shape (Fig.
9-3 B) with numerous chromatophores. Meanwhile the diploid nucleus of
the unilocular sporangium undergoes a meiotic division and then many

A B

Fig. 9-2. A, Ectocarpus showing tufted habit; B, Ectocarpus siliculosus, cell structure.

mitotic divisions until 32-64 haploid nuclei are formed. After nuclear divi-
sions, the sporangial protoplast divides into a number of uninucleate seg-
ments each with a single chromatophore corresponding to the number of
haploid nuclei. Each segment is finally transformed into a haploid, bifla-
geHate, pyriform zoospore in which the anterior longer flagellum is panto-
nematic and the posterior shorter flagellum is acronematic. After the pro-
toplast of a sporangium has been used up in the formation of zoospores, a
second sporangium may sometime regenerate from within the wall of the
first sporangium.
The plurilocular sporangium (Fig. 9-3 C) also develops from a terminal
cell which enlarges and accumulates numerous chromatophores. Subse-
quently it undergoes aseries of transverse and vertical mitotic divisions
producing several hundred small cubical cells arranged in definite tiers.
Bach cubical cell is diploid, uninucleate and forms a single biflagellate zoo-
spore identical to the zoospores produced from a unilocular sporangium
except that it is diploid. The haploid zoospores from uniIocular sporangia
are liberated en masse through a terminal pore but those from pluriIocular
sporangia generally come out one by one, rather in an orderly fashion,
148 A TEXTBOOK ON ALGAB

through a terminal (Fig. 9-4) or lateral pore. After moving freely for some
time the zoospores settle on some solid substratum and with their anterior
end still attached to some object, grow into new plants. The haploid zoo-
spores from unilocular sporangia form garnetophytes or sexual plants where-
as the diploid zoospores from a plurilocular sporangium produce sporo-
phytes or asexual plants.

B
Fig. 9-3. Ectocarpus. A, E. siliculosus with unilocular and plurilocular sporangia;
B, E. landsburgii with unilocular sporangia; C, E. /asciculatus with plurilocular
sporangia.

Sexual. The plurilocular gametangia arise in a manner similar to

plurilocular sporangia of the diploid sporophyte. In E. siliculosus, which
is heterothallic~ the loculi of all gametangia are of eq ual size. Consequently,
the zoogametes produced singly from each locule are also of equal size and
are morphologically identical with the zoospores. In such situations the
sexuality is isogamous though sometimes it may be accompanied by
physiological anisogamy since the male garnete may be more active and
 PHAEOPHYTA 149
motile than the passive and sluggishly motile fernale garnete. E. secundus
forms two kinds of gametangia and gametes, the megagarnetangium with
Iarger loeuli and Iarger gametes, and the mierogametangium with smaller

Fig. 9-4. Eclocarpus showing liberation of zoospores.

Ioeuli and smaIler gametes. Thus, E. secumlus may be morpbologically
anisogamous (Fig. 9-5 A-C).
In E. si/iculosus, several male garnetes cluster around a single femaIe
garnete with their Ionger flageIla attaehed to its surface (Fig. 9-5 D). The
fernale garnetes are known to produee a volatile sexual attraetant (cis-I-
[eycloheptadiene-2', 5'-yl]-butene) which eauses clumping of the spermato-
zoids (MuIIer et al., 1971). This is believed to be the first authentie report
of ehemical characterization of a sex hormone in plants. Sueh a sexual
union between zoogametes is unlike that observed in Chlamydomonas where
several male and female gametes cluster together through agglutination of
their flageIlar ends. Ultimately one of the male garnetes fuses with tbe
female (Fig. 9-5 E-G). The zygote (Fig. 9-5 H) always germinates directly
into a new plant and during its germination no reduction division oceurs.
The resulting plant is therefore diploid and represents the asexual sporophyte.
If agamete faiIs to unite with another gamete, it may germinate direetly into
a new plant.

Alternation of Generations
Sexual plants of Ectocarpus are dioecious and form only plurilocular game-
tangia. The zygotes upon germination do not divide meiotically. Hence tbe
150 A TEXTBOOK ON ALGAE

plants produced from germinating zygotes are alt diploid, and bear both
unilocular and plurilocular sporangia which always form diploid zoospores.
Since meiosis does not oeeur during their formation, they invariably produce

~ .
, .'
fernale ~
~ ; ga~mete
... " •
. 0

:v
. ( ..; " B
~ -1male
~ gamete ~
zygote
C

o
~ H
Fig. 9-S. Ectocarpus, sexual reproduction. A-C, E. secundus, stages in heterogamy;
D, E. silicu/osus, c\ump formation between one female gamete and many male gametes
E-G, stages in fusion of male and female gametes; H, zygote. (After Fritsch, 1945.)

diploid sporophytie plants and serve as an accessory means of asexual

reproduetion. But during the formation of zoospores in uniloeular spor-
angia, the first nucIear division is always meiotie and therefore such zoo-
spores are haploid and germinate into haploid gametophytic plants.
The gametophytie and sporophytie generations of Ectocarpus are mor-
phologically- alike and indistinguishable and, therefore, its life eycIe is
isomorphie. However, recent researehes have indicated that the life history
may be mueh more flexible and variable than hitherto assumed. MuIler's
(1967) work on E. siliculosus has established that its life cycIe is somewhat
heteromorphic-diplohaplontie rather than strictly isomorphic. Germination
of unfused gametes gives rise to haploid sporophytes whieh bear both uni-
loeular and pluriloeular sporangia. Similarly, diploid gametophytes and
tetraploid sporophytes mayaIso arise under certain conditions.
Order: FUCALES

Tbe Fucales are characterized by the absence of alternation between multi-

cellular sporophytic and free Iiving multicellular gametophytie generations
in the life cycle. The plants are all sporophytic or diploid. The gameto-
phytic or haploid phase is restrieted to the unicellular sperms and..eggs pro-
duced within unicelled reproductive organs termed antheridia and oogonia.
 PHAEOPHYTA 151

The number of eggs produced inside an oogonium ranges from one,

two, four to eight in various genera, but that of antherozoids within an an-
theridium is constant, being nearly 64 for all genera. The antherozoids of
the Fucales have a shorter anterior and a longer posterior flagellum whereas
in those of the other brown algae the arrangement is just the reverse. The
sexual reproduction i& oogamous and at the time of fertilization eggs and
antherozoids are both extruded from the fertile cavities or the conceptacles.

Family: Fucaceae
Gen us: Fucus

Occurrence
This alga grows attached to rocky substrata in the intertidal belt ofthe litto-
ral zone. Species of Fucus are predominantly cold water forms and grow
luxuriantly in the North Atlantic and Pacific shores.

Morphology
The plants, commonly less than 0.5 metre in length (the length depends on
the degree of exposure), have complex vegetative organization. Externally

Fig. 9-6. Fucus vesiculosus, thallus morphology.

each plant is differentiated into a basal discoid hold fast that an chors it to
the substratum, a short stipe and a flattened, dichotomously branched
frond or blade (Fig. 9-6). The thallus is generally dark brown, and slimy
152 A TEXTBOOK ON ALGAB

to the touch. The stipe is continued in the form of midrib in the frond of
many species, though it never reaches the frond apex. F. vesiculoslis con-
tains a number of air bladders in pairs and these give buoyancy to the sub-
merged plants. The mature thallus is in some respects similar to animal
cartilage, the cells being embedded in a biphasic matrix composed of rigid
fibres of alginic acid and cellulose.
Tbe swollen tips of fertile branches harbour a large number of smalI,
somewhat raised, flask-shaped cavities, the conceptacles (Fig. 9-7 A) which
contain the antheridia and oogonia and open to the exterior through an
ostiole. Lateral to the midrib are formed flat, wing-like expansions of the
thallus with sterile conceptacles called cryptoblasts or cryptostomata.
Internally, the thallus is differentiated into· three distinct regions-an
outer single-layered epidermis called meristoderm, a central many-layered
cortex, and an inner several-Iayered medulla. The cells of the medullary
region are loosely arranged, so me of them forming an anastomosing net-
work of hyphae. These first-formed hyphae are smooth-walled and are
called primary hyphae which may develop secondary thickening at matu-
rity. The region of the cortex cIose to medulla has much elongated muci-
laginous cells but tbe outer cortical cells and those of the epidermis are pro-
vided with chromatophores.
ostiole
I
--~~~~ ~~~

apical cell A

Fig. 9-7. A, Fucus serratus, transverse seetion through male receptacle; B, F. vesi-
culosus, vertical section through the apex.

The growth is initiated by an apical cell (Fig. 9-7 B) situated in the

hollow depression at the branch apex. The apical meristem is Jike a trun-
catcu pyramid with three cutting faces-two lateral and one basal. The
lateral segments derived from the meristem initially divide periclinally and
of the resulting two ceIls, the outer one, by repeated divisions in various
planes, organizes the epidermis and the cortex whereas the inner one, in as-
sociation with the basal segment cut off by the apical meristem, forms tbe
medulla. The epidermal layer as weil as the inner cells of cortex retain
meristematic potentiality, the former by cell divisi.Qns in vertical plane iu-
 PHAEOPHYTA 153

creases the girth and the latter gives rise to secondary hyphae. Thus, in
addition to primary growth, the alga also exhibits secondary growth.
In respect of functional differentiation, the epidermis and chromatophore
containing cortical region are photosynthetic, the inner cortical region stores
food reserves and the meduUary cells are conductive in nature. The
primary and secondary hyphae serve a mechanical function.

Reproduction
Vegetative. A thallus may fragment or dissociate into many parts,
each of which grows into a new plant.
Sexual. Plants are monoecious (F. spiralis) or dioecious (F. vesiculosus).
They may bear both antheridia and oogonia in the same or in different con-
ceptacles on the same plant in monoecious species, and in separate concep-
tacles on different plants in the dioecious species.
A conceptacle develops from a superficial cell whose rate of growth and
cell division slow down in comparison to those of neighbouring cells. Con-

tongue celll-':-~

A B

c o
Fig. 9-8. A-D. Fucus serratus, early stages in the development of conceptacIe.

sequently this apical cell comes to lie at the base of a flask-shaped cavity
(Fig. 9-8 A). This development is followed by transverse division of the
apical initial into two celIs, a lower basal cell and an upper tongue cell
154 A TEXTBOOK ON ALGAB

(Fig. 9-8 B). The latter may develop into a hair or degenerate. The basal
celI, by aseries of vertical divisions, gives rise to a layer of chromatophore-
containing celIs that line the f100r of the cavity. These cells undergo one or
two transverse divisions and form 2-3 layers (Fig. 9-8 C, D) of which the
superficial layer is always fertile giving rise to antheridia or oogonia, or
both.
Superficial cells wh ich start functioning as oogonial mother cells divide
into a tower stalk ceII and an upper oogonial ceII (Fig. 9-9 A). The first

c
A
Fig.9-9. Fucus vesiculosus. A, oogonium; B-C, formation and liberation of ova.

two divisions ofthe oogonial nucleus are always meiotic, invariably followed
by a mitotic division of aII the four haploid nuclei. The 8-nucleate oogonial
protoplast cleaves into 8 uninucleate eggs (Fig. 9-9 B, C) enclosed inside a
three-Iayered oogonial wall. The remaining celIs of the superficial layer
which fail to act as oogonial mother cells develop into multicellular hairs.
Some of the peripheral cells near the ostiole also form un branched hair-like
appendages, wh ich co me out of the ostiole in the form of a tuft.
·The antheridia are formed on lower branches (Figs. 9-10, 9-11 A) of the
hairs produced from superficial cells lining the cavity. Like oogonial nucleus,
the antheridial nucleus also divides meiotically, followed by four consecutive
mitotic divisions of the four haploid nuclei. Then after the completion of
nuclear division the antheridial protoplast fragments into 64 uninucleate,
phototactic antherozoids bearing two lateral ftagella of unequal length and
containing a large nucleus that occupies most of the cell cytoplasm thus
restricting the vestigial chromatophore, the well-developed eye spot and
other cellular organelles to a sm all area within the sperm (Fig. 9-11 B).
The antheridial wall is only two-Iayered.
During antherozoid development the eye spot originates de novo, as the
vegetative cells do not have any eye spots. In a detailed study of the eye spot
of F. vesiculosus, Bouck (1970) observed the following stages of develop-
 PHAEOPHYTA 155

ment: (1) appearance of electron-opaque granules along the chromatophore;

(2) increase in size of the granules; and (3) shifting of chromatophore, atong

Fig.9-10. Fucus, photomicrograph of transverse section through a male. conceptacle

showing antheridia.

with the eye spot, from centre of the cell towards its periphery, followed by
fusion with the plasma membrane. Four distinct microtubules and a few
fibrils arise near the base of the posterior flagellum between the chroma-
tophore and the plasma membrane. It is believed that the granules of the
eye spot arise from these microtubules and fibrils. Bouck (1970) considers
that light is reduced in intensity by a shading device (the eye spot) and
brought into focus by a focussing device (the flagellar base), and the photo-
receptor is probably sandwiched between these two structures, and bound
to the plasma membrane.
The internat strueture of the sterile eonceptacles (eryptoblasts) is similar
to that of the fertile conceptacIes except that they do not bear sex organs.
The hairs show either basal or diffuse growth, and are unbranehed. The
cavity contains some mueilage.

Fig. 9-11. Fucus. A, branched filament bearing antheridia; B, structure of antherozoid.

The wall layers of antheridia and oogonia are hydrophilie. They imbibe
water at the time of liberation of antherozoids or eggs as a result of which
156 A TEXTBOOK ON ALGAB

the outer waU layer sweIIs up and bursts pushing out the antherozoids or
the eggs through the hairs and ostiole to the exterior. The remaining wall
layers also become dissolved in the surrounding water and liberate the
antherozoids or eggs. In cultures, the cIustering of antherozoids around
a single egg has been seen (Fig. 9-12 A) and it results from a chemical
stimulus provided by a hydrocarbon (probably n-hexane) present in the eggs
of Fucus. UItimately, one of the antherozoids finds its way into the egg,
leading to the formation of the zygote.

D E

Fig. 9-12. Fucus, stages in thallus development. A-C,F. furcatus (after Smith, 1955);
A, fertilization of egg; B, formation of rhizoidal initial; C, transverse section through
two-day-old embryo; D-E, subscquent stages in embryo dcvelopment.

The FUCIlS egg can show parthenogenetic development and in the

laboratory parthenogcnesis is induccd by treating the eggs with dilute solu-
 PHAEOPHYTA 157

tions of acetic or butyric acid. During zygote germination (Fig. 9-12 B-E)
or parthenogenetic development of the egg, the point at which the primary
rhizoid develops is away from light indicating the existence of a light-
controlled mechanism which regulates the development of a polarity gradient.
Apart from visible light, uItraviolet light, hydrogen-ion concentration, tem-
perature and auxin gradients, electrical field, mechanical deformation and
the point of entry of the spermatozoid, all seem to control the development
of polarity leading to the formation of rhizoids. A C02-pH gradient seems to
control the auxin concentration and the region rich in auxin activity
differentiates the rhizoid.
After the point of rhizoid formation has been determined, the zygote
divides mitotically in a plane transverse to the axis defined by the rhizoid
(Fig. 9-12 B). The upper cell of the two-celled embryo divides verticalIy
whereas the lower divides transversely, thus forming a 4-celled embryo
(Fig. 9-12 C). The upper two cells, together with the median celI, divide
and redivide transversely and periclinally giving rise to a tissue differentiated
into medulla and cortex (Fig. 9-12 D). The lowermost celI of the quadrant
from wh ich the primary rhizoid originated also forms secondary rhizoids.
With the growth of the embryo, there differentiates an apical cell in
a small depression which forms a filamentous hair (Fig. 9-12 E). Sub-
sequently, the cells adjacent to the hair also grow into hairs and form a
tuft on the upper surface. This development is followed by degeneration
of all the cells of the hair except the basal cell of the first-formed hair.
This basal cell finally functions as an apical initial of the Fucus thalIus.

Family: Sargassaceae
Genus: Sargassum

Sargassum, like Fucus, is diploid and lacks an alternation of generations.

The two genera mainly differ in their external morphology as weil as in the
number of eggs produced in an oogonium (eight in Fucus but one in
Sargassum).

Occurrence
The alga, popularly called gulfweed, grows abundantly in tropical oceans in
the Southern hemisphere. It may be free-floating or attached to rocks.
The chief centres of its growth are the Australian and Caribbean coasts,
Gulf of Mexico and the Sargasso Sea of south central Atlantic. It is also
abundant on the West and East Coasts of India.

Morphology
Thalli of Sargassum are monopodially branched and the members of Sargassa-
ceae are chiefty identified by the peculiar features of their lateral branch
systems (Fig. 9-13 A, B). The plant body is usually differentiated into a
holdfast, a short stipe and much branched long laterals which give rise at
regular intervals to leafy laterals bearing cryptoblasts. From near the axil
158 A TEXTBOOK ON ALGAB

of the leafy laterals, called primary branch, comes out a second order of
branches, the first formed one or two branches of which transform into air
bladders and the remaining branches serve as receptacles (Fig. 9-13 C, D)
bearing both conceptacles and cryptoblasts. The 'leaves' have smooth or
serrate margins and are often provided with a prominent midrib.

Fig. 9-13. A, Sargassum longi[olium, part of fertile plant; B, Sargassum swartzii, part
of plant; C-D, S. swartzii, portions with receptacle. (Figs. B-D after Chauhan and
Thivy, 1964.)
A transverse section through a branch, leaf or stipe shows well-marked
differentiation into three regions, the meristoderm, cortex and medulla. In
some species, which are free-floating, the medulla may be almost absent.
Unlike Fucus, the cortex of Sargassum neither contains mucilaginous cells
nor hyphae. The air bladder consists of an epidermis, cortex and a central
hollow cavity filled with air.
As in Fucus, growth and organization of the thallus is achieved through
the activity of an apical cell with three cutting faces.
 PHAEOPHYTA 159
Reproduction
Vegetative. S. natans and S. hystrix, which grow free-floating in the
Sargasso Sea, are known to multiply exclusively by vegetative means. It
is interesting that free-floating algae of salt marshes, e.g., Pelvetia and
Bostrychia, also multiply vegetatively.
Sexual. Species are both dioecious and monoecious but, in contrast to
Fucus, Sargassum forms special branches that bear conceptacles which
may be fertile (Fig. 9-14 A) or sterile (Fig. 9-14 B). The conceptacles and
antheridia develop in the same manner as in Fucus; oogonial development

conceptacle

c
Fig. 9·14. A, Sargassum swartzii, transverse section through rachide showing cornpressed
nature (after Chauhan and Thivy, 1964); B, S. filipendula, section of youog cryptoblast;
C, S.filipendula, sectioo of young conceptacle; D, S. Iinijolium, section through a youog
fern ale cooceptacle showiog ernbedded oogonia; E, S.filipendula, gerrnIing at rhizoidal
stage.

(Fig. 9-14 C, D) also follows the same course up to the stage of formation
of 8 nuclei, but 7 out of the 8 nuclei degenerate and the remaining nucleus
along with the oogonial cytoplasm forms a single egg.
 160 A TEXTBOOIC ON ALGAB

After maturation of the oogonium, the outer wall tayer ruptures and thc
middle develops into a mucilaginous stalk wh ich pushes the entire oogonium
through the ostiole to the exterior while still attached to the mother plant.
Fertilization occurs inside the oogonium and even the resulting zygote starts
germinating within the oogonial wall. The remaining features of sexual
reproduction and development of embryo into a germling (Fig. 9-14 E) are
similar to those of Fucus.

TEST QUEST/ONS
1. What explanations can you offer ror the occurrence of brown algae in distinct zones
along the littoral and sublittoral regions?
2. Both Chrysophyta and Phaeophyta are similar in respect of their pigmentation, food
reserves and ftagellar morphology, yet they are treated as separate phyla. Why'l
3. In the absence of any unieellular algae in the Phaeophyta, how would you account
for the aneestry of this Phylum '1 Explain, giving reasons.
4. It is generally eonsidered that an alga with heteromorphie life cycle and hetero-
triehous habit would be nearest to the hypothetieal progenitor of land plants. How
about certain brown algae '1
5. Give two most important and least variable features that have formed the basis of
classifieation of Phaeophyceae into various orders.
6. Compare and contrast the anatomical features of Fucus or Sargassum with tbose of
an angiospermic plant.
7. In wbat ways does tbe life cycle of Fucales differ from tbat of any angiospermic plant?
8. Support with arguments the hypothesis tbat the evolution in tbe Iife cycle of algae
ended with a Fucus-Iike form.

SELECTED REFERENCES
BoaIch, G. T., Studies on Eclocarpus in eulture. 11. Growtb and nutrition of a baeteria-
free culture, J. mar. biol. Ass., U.K., 41, 287-304 (1961).
Bouek, G. B., The development and postfertilization fate of the eyespot and the apparent
photoreeeptor in Fucus sperm. 111 Frederiek, J. F. and Klein, R. M. (eds.)
Phylogenesis and morpbogenesis in the algae, Ann. N. Y. Acad. Sci. ,175, 673-85
(1970).
Evans, L. V., Distribution of pyrenoids among 'some brown algae, J. Cell Sci., I, 449-54
(1966).
Manton,l., Some phyletic implications of ftagellar structure in plants, Adv. Bol. Res., 2,
1-34 (1965).
Müller, D. G., Generationswechsel, Kernphasenweehsel und Sexualität der Braunalge
Ectocarpus siliculosus im Kulturversueh, Planta, 75, 39-54 (1967).
Müller, D. G., Jaenieke, L., Donike, M. and Akintobi, :r.,
Sex attractant in a brown alga:
chemical strueture, Science, Wash., 171, 815-17 (1971).
Parker, B. C. and Huber, J., Translocation in Macrocyslis. 11. Fine structure of the
sieve tubes, J. Pllycol.,I, 172-79 (1965).
 10
Rhodophyta
The Rhodophyta (red algae) owe their colour to the presence of excess of
r-phycoerythrin in their chromatophores and this red pigment masks the
colour of other photosynthetic pigments. The Rhodophyta are characteriz-
ed by six main features: (1) the flagellated motile stages are totally absent;
(2) the sexuality is highly specialized; the male gamete, called spermatium, is
motionless and at the time of fertilization is passively transported to and
lodged on the trichogyne of the female carpogonium; also, there are distinct
post-fertilization developments not found in any other algal phyla; (3) chloro-
phyll-d, biliproteins (r-phycoerythrin and r-phycocyanin) and the xantho-
phyll taraxanthin are the characteristic pigments; besides, the chromato-
phores generally contain chlorophyll-a, cx. and ß carotenes, lutein, zeaxan-
thin, neoxanthin and rarely other xanthophylls; (4) the reserve foods are
floridean star eh and galactoside floridosides and these do not accumulate
within the chromatophore but outside it, in the cytoplasm; (5) the cell wall
contains polysulphate esters of carbohydrates in addition to cellulose and
pectin; and (6) the transverse walls in multicellular forms are generally
provided with pits which permit cytoplasmic connection between adjacent
cells.

OCCURRENCE
The unicellular red alga Porphyridium grows on damp soil. Except for a
dozen genera, e.g., Balrachospermum and Lemanea, which are freshwater
forms, others are exclusively marine and grow mostly in the intertidal and
sublittoral regions. Members of Rhodophyta grow io. almost all marine
habitats, but their greater concentration occurs in the warmer seas.
Some of the calcareous algae, e.g., Corall ina and Lithothamnion, possess
stony thalli and are largely responsible for the formation of coral reefs.
Ceralocolax, Choreonema and some others are parasit es on macrophytic
marine algae.
Visible light, particularly its blue and red regions, which are mainly
effective in photosynthesis, seems to playa somewhat important role in the
ecological distribution of the sublittoral marine algae since with increase in
~ depth of water there is a corresponding decrease in the quantity and quality
- of incident light. At a depth of about 10 metres only 10 per cent of the
~. blue-green and 1 per cent of the red light are received. At still deeper levels,
_ further reduction in the quantity of these spectral regions takes place .
....; Under such light conditions, main]y Rhodophycean members can photo-
162 A TEXTBOOlC ON ALGAB

synthesize because r-phycoerythrin present in them can efficiently absorb the

available blue-green light. They also exhibit complementary chromatic
adaptation so that the colour of the incident light induces the development
of a particular photosynthetic pigment which has maximum absorption of
the incident light. Possibly for these reasons the red algae can grow at
depths where few others can subsist.

RANGE OF THALLUS ORGANIZATION

Perhaps the most beautiful and most showy of the marine algae, forms such
as Plocamium and Delesseria are areal delight to behold and collect. When
mounted on a herbarium sheet and dried, a specimen of Plocamium looks
like the creation of a painter. In general, not as large as the brown algae,
the red algae are more delicate in texture and more slimy and soft to
touch.
The Subclass Bangioideae, with the single Order Bangiales, comprises
unicellular (Porphyridium), filamentous (Goniotrichum) and parenchymatous
(Porphyra) forms. The thalli of Florideae are more elaborate, with two
main types of organization-uniaxial and muItiaxial-and in both a pseudo-
parenchymatous thallus results from the coalescence of filament branches.
Uniaxial thalli have a single central or axial filament, which may be corti-
cated, with a number of richly branched laterals organized to form a
pseudoparenchymatous structure, e.g., Dumontia and Batrachospermum. In
the multiaxial types, there are several central or axial filaments which, to-
gether with their branches, form the pseudoparenchymatous thallus, e.g.,
Helminthocladia.
The growth of the thallus in Bangioideae is diffuse whereas, with a few
exceptions, in Florideae apical growth is the rule. An apical cell, by conse-
cutive divisions, forms a transverse series of cells each of which further
divides tangentially to produce a group of pericentral cells that surround a
central cello The pericentral cells may not divide further (as in Rhodome-
laceae) or may divide vertically to form a cortex or the initials ofthe lateral
branches which co ales ce to form a compact structure with internal differen-
tiation into an outer photosynthetic zone, an inner food storing cortex, and
a central medulla containing vertically elongated hyphae, e.g., Gelidium and
Gloiosiphonia.

CELL STRUCTURE

The cell wall is made up of an inner microfibrillar cellulosic layer and an

outer pectie layer. In most cases it contains an outer coating of mucilage.
Cells may either harbour a single, stellate, axile chromatophore with a
central, naked pyrenoid (Bangioideae), or many discoid and parietal chro-
matophores devoid of pyrenoids (Florideae). The entire dass seems uni-
form in the ultrastructure of the chromatophore which is a double
membrane organelle with widely separated single thylakoids. Arecent
 RHODOPHYTA 163

electron microscopic study of Porphyridium sp. has revealed the presence of

particles called phycobilisomes, containing phycoerythrin and phycocyanin.
Phycobilisomes are arranged linearlyon the surface of the photosynthetic
lamellae, those containing phycoerythrin are spherical whereas those having
phycocyanin are discoid. In addition to phycobilisomes, many scattered
grains of 1l0ridean starch occur in tbe cel! cytoplasm.
The Rhodophyta show considerable variation in the shape, size and
number of nuclei. Cells of Bangioideae are always uninucleate with a
single nucleolus, those of Florideae are more complex. In general, if the
apicaJ cells are uninucleate, the segments derived from them will also be
uninucleate. In such forms as Griffithsia there may be as many as 4000
nuclei per cello The shape of nucleus varies depending on its position in
the cello Thus, it is spherical when centrally situated, and llat when parietally
located. Nuclei may migrate from one cell to another through a pit con-
nection. Some red algae, e.g., Porphyra, exhibit a diurnal periodicity in
their nuclear division. A few red algae have revealed the presence of
dictyosomes, endoplasmic reticulum and tonoplast-bounded vacuoles (Brody
and Vatter, 1959). Mito~hondria have recently been reported in Batra-
c/lOspermum moniliforme.

REPRODUCTION

Unicellular forms propagate vegetatively. In multicellular forms, asexual

or sexual reproduction is more common.

ASEXUAL

This is brought about by non-motile spores which are given different names
depending on the nature of the cells from which they are produced and
their number within each sporangium. Accordingly, there are four chief
types of asexual spores: (.1) monospores, formed singly in monosporangia;
(2) neutral spores, formed by direct transformation of vegetative cells into
spores; these two types of spores are characteristic of the Bangioideae;
(3) carpospores, formed either directly from the division products of the
zygote, as in Bangioideae, or indirectly from the cells of certain filaments
termed gonimoblasts that arise from the fertilized carpogonium or from an
auxiliary cell containing the zygote nucleus; and (4) bispores, tetraspores, and
polyspores produced respectively in twos, fours and multiples of fours within
a sporangium of the diploid tetrasporophyte. Except for some red algae that
form haploid carpospores growing into sexual plants, most red algae form
diploid carpospores which on germination give rise to sporophytes or diploid
plants, bearing diploid sporangia from which bispores, tetraspores or poly-
spores are produced after meiosis. These spores are therefore haploid and
germinate to form gametophytes or haploid plants.
A tetrasporangium may be cruciate if the two divisions of the cell are at
right angles to each other, tetrahedral if the two divisions are oriented in
164 A TEXTBOOK ON ALGAB

such a manner as to form a group of four tetrahedrally disposed spores or

zonate if there are three parallel transverse divisions.

SEXUAL
The motionless male cells or spermatia are formed singly within spermatangia
produced in clusters either on special branches, as in Polysiphonia, or in
definite sori, as in Apoglossum. The female cell, designated carpogonium,
is a fiask-shaped cell with a neck-like protuberance, the trichogyne. Tbe
egg nucleus is restricted to the basal portion. In Bangioideae, carpo-
gonia are sessile but in Florideae they are stalked and produced ter-
minally on a special branch, the carpogonial filament or procarp. During
fertilization a spermatial nucleus passes down through the trichogyne and
fuses with the nucleus of the egg. The sexual reproduction, from sex organ
formation to fertilization, is remarkably uniform throughout the Rhodo-
phyta, particularly the Subclass Florideae, which shows elaborate post-
fertilization changes resulting in tbe production of a new generation, the
carposporophyte, which is parasitic on the gametophyte. Such distinct
stages in post-fertilization activities as the place offormation of gonimoblast
filaments (either from fertilized carpogonium or from generative auxiliary
cell, and in thc latter case, the position and time of formation, if before or
after fcrtilization of auxiliary cell) are so important and significant that they
have provided the main basis for the classification of Florideae into various
orders.
In Bangioideae, no new generation, like carposporophyte, is produced
but the zygote divides directly into carpospores which germinate to form a
Conchocelis-like vegetative phase. The general opinion among phyco-
logists is that the mature plants of Bangioideae, e.g., Porphyra, Bangia and
others are diploid and it is during the formation of carpospores that meiosis
occurs. These carpospores produce haploid Conchocelis-Iike plants. Since
there is no unequivocal cytological evidence for the occurrence of meiosis,
whether during germination of zygote or formation of monospores or neutral
spores, the controversy about the diploidy or hapl'oidy of Porphyra or Bangia
thalli is yet to be resolved. Some recent evidence indicates that Conchocelis
may be diploid (Von Stotsch, personal communication). Morphologically,
Conc/lOcelis is regarded as the prostrate system and Porphyra thallus as the
erect system of an originally heterotrichous alga.
In some Florideae, the gametophytes are haploid and the zygote nucleus
may divide meiotically or more often mitotically in the carpogonium itself.
In the majority, however, the zygote nucleus migrates into a well-differentiat-
ed cell, known as auxiliary cell, either through pit connections or through
a tubular connection, the ooblast, specially established for this purpose bet-
ween the carpogonium and the auxiliary cell. The division of the zygote
nucleus in the auxiliary cell is always mitotic, hence the gonimoblast filaments
produced are all diploid and so also are the resulting carposporopbytes.
Gonimoblast filaments produced from the carpogonia after meiosis are hap-
 RHODOPHYTA 16 S

loid but in the others they are diploid. Accordingly the carposporophytic
generation of the former is haploid whereas that of the others is diploid.
Either terminal cells or all the cells of a gonimoblast filament may functioll
as carposporangia. Each carposporangium forms a single carpospore which
may be haploid or diploid depending upon the nature ofthe carposporophyte.
ßoth carposporophyte and tetrasporophyte formed by the germination of the
carpospores constitute the asexual generations.
Recent researches indicate that meiosis occurs in the developing apex of
young plants of Nemalionales and not in the fertilized carpogonium.

LlFE CYCLE

Some red algae (e.g., certain Nemalionales) have two alternating haploid,
gametophytic and carposporophytic generations. Remaining orders of the
Florideae have three generations-a haploid gametophyte, a diploid carpo-
sporophyte and a diploid tetrasporophyte, one following the other in that
seq uence. The gametophytes and tetrasporophytes are two morphologically
identical, free living generations, with an intercalation of a parasitic, diploid,
carposporophytic generation. The products of the zygote nucleus form the
carposporophyte whose carpospores give rise to tetrasporophytes. The
gametophytic generation results from tetraspore germination.

CLASSIFICATION AND GENERAL CHARACTERS

Most phycologists divide the Cl ass Rhodophyceae into two subclasses:

(I) the Bangioideae (or Bangiophycidae) comprising the only Order Bangiales,
and (2) the Florideae (or Florideophycidae) with 6 orders-Nemalionales,
Gelidiales, Cryptonemiales, Gigartinales, Rhodymeniales and Ceramiales.
The Bangioideae include those forms which have simple thalli with cells
containing single, axile chromatophores and are normally devoid of pit
connections. The zygote divides directly into haploid carpospores. The
thallus of the Bangioideae shows diffuse growth in contrast to the apical
growth of the Florideae.
The division of the Florideae into 6 orders is based on features of post-
fertilization development. Except for some Nemalionales, which do not
have a free living diploid generation, all the other orders have two diploid
generations, the carposporophytic and the tetrasporophytic. The latter is
free living. The Nemalionales and Gelidiales have been segregated due to
the lack of auxiliary cells in them. The Cryptonemiales and Gigartinales
form auxiliary cells before fertilization whereas Ceramiales develop them
after fertilization. In Rhodymeniales the auxiliary cells differentiate
before fertilization but develop weIl and become easily recognizable only
after fertilization. The Cryptonemiales differ from Gigartinales because
their carpogonial branches always arise from special accessory branches,
whereas in Gigartinales they are merely unmodified ordinary branches. In
Rhodymeniales the auxiliary cell develops from the outer cell of a two-
166 A TEXTBOOK ON ALGAB

celled branch but in Ceramiales it is formed from the supporting ceU.

PHYLOGENY
A fairly homogeneous, well-circumscribed group, the Rhodophyta apparently
show much eloser relationship with the Cyanophyta than with any other
algal phyla. Tbe features shared by the two phyla are: (1) the absence
of flagellated structures; (2) the formation of similar biliproteins (phycocya-
nin and phycoerythrin) as the accessory photosynthetic pigments; (3) the
presence ofpit connections; (4) the accumulation offairly similar food reserves,
e.g., floridean starch in Rhodophyta and cyanophycean starch in Cyano-
phyta; (5) the production of chemically similar mucilages in blue-green algae
and in some Bangioideae and Nemalionales; and (6) similar patterns offatty
acid synthesis which differ from those of other algae in that the fat content
does not increase with increase in the age of the thallus, and that the nitro-
gen starvation is not a factor in fatty acid accumulation (Fogg and Collyer,
1954). See also Table IV.
Table IV. Resemblauces betweeo Rbodopbyta Rod Cyaoopbyta

1. Both have water soluble phycobilins, located on phycobilisomes, as accessory

pigments
2. Free trehalose aod galactose occur only in them
3. The food reserves, cyanophycean and rhodophycean starch, are elaborated by
similar types of Iinkages and chemical routes
4. Linoleic acid and II-Iinoleic acid are present
S. Neither nitrogen starvation nor aging ofthallus has any effect on the pattcrn of
fatty acid accumulation

6. Xylans are the major components in the cellulose microfibrillar walls

7. Sulphated galactoses, uronic acids, glucose and xylose are the principal consti-
tuents of the mucilage

8. The organization ofthe photosynthetic thylakoids is similar, i.e., they occur singly
and are widely separated

9. The patterns of isoenzymes involved in the elaborat ion of polyglucosides are similar
10. Both lack flagellated stages.

However, the biliproteins of Rhodophyta differ from those of Cyanophyta

in their spectral properties. Cryptophyceae too are now known to possess
biliproteins. The biliprotein allophycocyanin is probably identical with the
phytochrome ofhigher plants, and future research may indicate the presence
ofthis or other biliproteins in algal phyla other than Rhodophyta and Cyano-
phyta. In view of these, too much phylogenetic importance should not be
attached to the biliproteins. See also Table V.
 RHODOPHYTA. 167
Table V. Difl'ereoces betweeo Rbodophyta aod Cyaoophyta

Redalgae Blue-green algae

Cell strueture Euearyotie Proearyotic

Pigments Chlorophyll-a and -d, Only chlorophyJl-a, rI\yxoxanthin, my-
zeaxanthin and neoxanthin xoxanthophyll, oscillaxanthin
Food reserves Rhodophyeean stareh, no Cyanophycean stareh, and the protein-
proteins aeeous eyanophyein
Cell walls Cellulose and other Mueopolymers constitute the main
earbohydrates present eomponent in eertain forms
Motile stages None Non-flage\1ate multicellular motile
stages, Le., hormogonia formed in
many forms, mature individuals of
some species are also motile though
non-flagellate
True sexuality Present in most members Absent, a primitive type of parasexua-
Iity, i.e., genetie recombination, oeeurs
in some members
Chromosomes Typiea\1y euearyotie True chromosomes not known, DNA
fibrils not assoeiated with histones but
present in the nucleoplasm
Golgi bodies Present Absent
Endoplasmie Present Absent
retieulum
Heteroeysts and None Many speeies have heterocysts and ean
Nitrogen fixation fix atmospheric nitrogen
Suseeptibility to Not known Cyanophages LPP-l and SM-l ean
virus infeetion infeet a few speeies

The stacking together of photosynthetic thylakoids into distinct bands is

regarded as an advanced feature and in this respect the Rhodophyta is the"
most primitive Phylum among eucaryotic algae since in the red algae the
photosynthetic lamellae are single, unorganized, widely separated and not
stacked into bands or multiple lamellae. Nonetheless, the Phylum Rhodo-
phyta is morphologically as complex as Chlorophyta. Regarding the an-
cestry of the class, two alternatives have been suggested (see Klein and
Cronquist 1967): (1) that they originated from some simple procaryotic
blue-green algae, and (2) that they were derived from some archaic eucar-
yotic algae which themselves originated from blue-green algae. The funda-
mental disparity between the eucaryotic and procaryotic cellular organiza-
tions seems to rule out the existence of any direct link between the red and
the blue-green aIgae. It would seem more plausible to speculate that the
former probably originated from some primitive non-flagellated eucaryotic
ancestor possessing biliproteins.
168 A TEXTBOOK ON ALGAE

Order: N EMALIONALES
Family: Batrachospermaceae
Genus: Batrachospermum

Occurrence
It is an inland freshwater form widely distributed in tropical, subtropical
and temperate regions. Slow-ftowing streams, waterfalls and oligotrophie
lakes are some of the habitats where it grows attached to stones. Most
species prefer relatively weil aeratcd 'clean' water streams. Some taxa are
marine.

Morphology
Adult plants, which may be up to 15 cm long, are generally bluish-green,
greyish-green, violet or olive-green, and soft and mucilaginous. To the
naked eye, each plant appears as a branching chain of beads (Fig. 10-1 A).
The alga is differentiated into a prostrate system that serves to anchor it to the
substratum and an erect branched system, made up of whorls (Fig. 10-1 B),
which Boats freely in water. Many species are attached by rhizoids.

A
Fig. 10-1. Batrachospermum moniliforme. A, part of thallus; B, two whorls of branches
(magnified).

Tbe thallus is uniaxial. Its primary axis consists of a uniseriate row of

large cylindrical cells, frequently terminating in a hemispherical apical cell.
The axial filament is differentiated into nodal and internodal regions and is
corticated. From the nodes arise two kinds of branches-the branches of
limited growth which are produced in whorls, some branches often termi-
nate in hairs; and those of unlimited growth which are gene rally formed
singly and monopodially and Iike the primary axis, are further differentiated
into nodes and internodes. The bead-like appearance of the alga is due to
 RHODOPHYTA 169

tbe whorls of nodal branches. Tbe cluster of branches at anode is called a

glomerule. Tbe nodal branches of limited growth are composed of monili-
form cells and arise from just below septa of the axial filament, or from
branches of unIimited growth whose cclls are much longer than those of
branch whorls. The cells are uninucleate and contain several parietal chro-
matophores, each with a single pyrenoid.
The growth of the axial filament and ofthe branches of unlimited growth
occurs by means of an apical cell which cuts off segments transversely and
produces a uniseriate row of cells. From each cell, near the septum, 4-6
lateral projections arise which soon become separated from the parent cell
by a wall and function as initials of branches of limited growth. After the
formation of the lateral branches, their basal cells give rise to corticating
filaments. A branch of unlimited growth mayaIso develop from one of
these basal cells.

Reproduction
Batrachospermum reproduces sexually by thc formation of spermatia and
eggs, and asexually by carpospores.
The plants are monoecious or dioecious. Tbe terminal or subtcrminal
cells of branches of Iimited growth constitute thc spermatangial initials.
Each initial produces one or two spermatangia wh ich are generally colour-
less and can therefore be easily distinguished from vegetative cells or bran-
ches.
Tbe basal cell of a branch of limited growth cuts off a segment which is
called the carpogonial initial. Tbis initial undergoes 3 or 4 transverse divi-
sions and forms a four to five-celled carpogonial branch whose terminal cell
develops into a carpogonium. Each carpogonium is differentiated into a
nucleate basal part and an enucleate neck or trichogyne, the latter being
sometimes demarcated from the former by a median constriction.
The spermatia are carried passively along water currents to the carpo-
gonia. During fertilization, the spermatium nucleus after migrating through
the trichogyne, fuses with the egg nucleus (Fig. 10-2 A, B) of thc carpogo-
nium. After fertilization, the trichogyne shrivels and the zygote nucleus
divides meiotically to form four haploid nuclei. At this time the fertilized
carpogonium develops many small protuberances. Each of the four nuclei
divides mitotically and one of the two resulting nuclei mi grates into a pro-
tuberance which then becomes separated from the carpogonium and starts
functioning as an initial of a gonimoblast filament. Thus a number of
branched gonimoblast filaments (Fig. 10-2 C) arise from the base of thc
carpogonium. The terminal cclls of the gonimoblasts later differentiate into
carposporangia within wh ich the carpospores are formed singly. The clus-
ter of gonimoblast filaments along with associated carposporangia constitute
the carposporophyte wh ich grows as a parasite on the female gametophyte.
Since meiosis occurs during division of the zygote nucleus, the carposporo-
phyte and the carpospores are haploid. Some of the vegetative cells sur-
rounding the carpogonium grow in the meanwbile and form a loose sheath
170 A TEXTBOOK ON ALGAB

of sterile branehes that encIoses the earposporophyte which along with the
sheath forms a eharaeteristie fruit body known as c)'stocarp.

spermaUum

A B c
Fig. 10-2. Batrachospermum moniliforme. A-C, fertilization and post-fertilization
stages.

Eaeh carpospore on germination forms a branehed, heterotrichous proto-

nema-like filament. This eonstitutes a juvenile stage in the life history of
Batrachospermum and was previously believed to be an independent algal
genus Chantransia. Almost all freshwater species of Chantransia have been
found to be the juvenile stage (Fig. 10-3 A) of Batrachospermum. The
Chantransia thalli reproduce asexually by means of monospores formed
singly in monosporangia (Fig. 10-3 B). Thus the monospores merely serve
as an accessory means of multiplieation of the Chantransia stage. The
Batrachospermum plant arises from the Chantransia stage as a lateral out-
growth. Interpreted in terms of the heterotrichous habit, both the Batra-
chospermum phase and the Chantransia phase eonstitute the two systems of
a heterotrichous plant, the former representing the ereet system and the
latter the prostrate system. A parallel situation is found in another red alga
Porphyra which itself represents the ereet system whereas the prostrate sys-
tem is constituted by its Conchocelis phase.

Life Cycle
Batrachospermum ineludes three kinds of haploid somatie phases in its life
eycIe: (I) the free living gametophytie phase, (2) the parasitie carposporo-
phytie phase, and (3) the free Iiving Chantransia phase. The gametophytes
 RHODOPHYTA 171

reproduce sexually through the formation ofspermatia and eggs which unite
to form zygotes. The zygote germinates meiotieally forming a large num-

Fig. 10-3. A, BatracllOspermum ectocarpum, Chantransia stage; B, B. lochmodes, mono-

sporangia.

ber of branched gonimoblast filaments with their terminal cells functioning

as earposporangia. Thus a haploid earposporophyte parasitic on the game-
tophyte results from germination of zygote. The carposporophyte in turn
gives rise to Chantrallsia phase and Chaniransia gives rise to Batrachosper-
mum as a special braneh. Both the earposporophytie and Chantransia
phases are asexual, and the latter perpetuates asexually by monospores.
Cytologieally the life eycle of the alga is haplontie 'in view of the zygote
being the only diploid phase, and morphologieally trimorphie because of
the succession of three distinet morphologieal phases.

Order: CERAMIALES
Family: Rhodomelaceae
Genus: Polysiphonia

Occurrence
Polysiphonia grows extensively in the intertidal belt and sublittoral region.
Most species are epiphytie on Fucaceae and other larger marine algae. P.
jastigiata grows attached to the fronds of Ascophyllum nodosum; this speeies
may be a semiparasite in view of the destruetion of some of the host cells
172 A TEXTBOOK ON ALGAB

near the point of its attachment. Some common species on thc West Coast
of India are P. variegata, P. urceolata and P. platycarpa. The plants grow
in dense tufts (Fig. 10-4 A).

Morphology
The genus derives its name from the polysiphonous nature of the thallus
which consists of an axial row of central siphons (Fig. 10-4 B) surrounded
by a layer of 4-24 pericentral siphons. Tbe plant body is heterotrichous
with an erect system of branches and a filamentous prostrate system anchor-
ing the plant to the substratum with the help of unicellular elongated rhi-
zoids whose tips are flattened into lobed discs or haptera. The rhizoids arise
from the pericentral cells facing the substratum. Tbe thallus is dichoto-
mously or laterally branched with two kinds of branches, the branches of
unlimited growth made up of central and pericentral siphons and those of
limited growth, known as trichoblasts, which appear like hairs, are dichoto-

Fig. 10-4. A, Polysiphonia nigrescens, tufted habit; B, P. sertularioides, apex of thallus;

C, spermatangia as seen in a section through fertile trichoblast (after Smith, 1955).

mously branched, tapering, uniseriate filaments made up of uninucleate

colourless cells. Pit connections occur in between and among the central
and the pericentral siphons (Fig. 10-4 C). The older parts ofthe thallus are
generally corticated with cells cut off periclinally by pericentral cells ofthose
regions. In surface view, under low magnification, the thallus of Polysipho-
nia seems to consist of a number of polysiphonous tiers. The orderly man-
ner in which the tiers are arranged suggests a differentiation of filaments
into nodes and internodes.
The thallus grows by means of an apical cell which by repeated divisions
rorms a row ofaxial cells. All axial cells excluding the first few cells
undergo tangential vertical divisions and form a peripherallayer of a specific
number of pericentral cells around the axial row of central cells. Both
central and pericentral cells elongate into their respective siphons having
 RHODOPHYTA 173

the usual pit connections. The trichoblast initial arises from an axial cell.
4 or 5 cells away from the apical meristem even before the pericentral cells
are formed. Branches of unlimited growth also arise similarly but unlike
the trichoblast initial, the initials of these branches behave exactIy like the
apical cell of the main axis and produce at first a transverse file ofaxial
cells from which later the peripheral layer of pericentral cells arises. A
branch of unlimited growth may sometimes arise in the axil of a trichoblast
in which case its basal cell serves as the branch initial.

Reproduction
Polysiphonia is generally heterothallic or dioecious and includes three kinds
of morphologically similar plants, the male gametophyte. the female game-
tophyte and the teuasporophyte. The female plant also bears the diploid
parasitic carposporophyte. Both carposporophyte and tetrasporophyte
reproduce asexually by the formation of carpospores and tetraspores
respectively.
Sexual. The male and female organs are borne on different plants.
Spermatangia and carpogonia occur on small fertile trichoblasts. Tbe male
trichoblast, after becoming 2-3 cells long, forks; generally one, rarely both,
of the branches so produced participates in the formation of spermatia.
If only one branch forms spermatia, the remaining sterile branch repeatedly
divides dichotomously. All cells except the two basal cells of the fertile
branch divide and form central and pericentral cells. The latter function
as spermatangial mother cells and produce one or a few spermatangia each
along their free surfaces.
The two or three basal cells of the female trichoblast give rise to central
and pericentral cells, and generally the rniddle cell of the three adaxial peri-
centrals produces on its free side a supporting cell (Fig. 10-5 A) which by
successive transverse divisions organizes a 4- or 5-celled carpogonial branch
(Fig. 10-5 B), the procarp. The terminal cell of the procarp transforms
into a carpogonium with a swollen base containing a uninucleate egg and a
long drawn-out enucleate trichogyne (Fig. 10-5 C-D). Meanwhile, the
supporting cell also cuts off two sterile ceIls, one towards its base and the
other towards its side. Both these cells form a few-celled sterile filaments
(Fig. 10-5 E). Following these developments, fertilization occurs and a
spermatium nucleus passes down the trichogyne and fuses with the egg
nucleus in the swollen bases of the carpogonium (Fig. 10-5 F). This is
accompanied by the cutting off of an auxiliary cell by the supporting cell of
the carpogonium towards its upper side (Fig. 10-5 F). Soon a tubular con-
nection is established between the auxiliary cell and the carpogonium base.
The zygote nucleus then migrates to the auxiliary cell through this tube.
By this time the trichogyne degenerates and the two pericentral cells of the
trichoblast adjacent to the supporting cell start producing a sterile sheath,
known as pericarp, around the developing carposporophyte; the pericarp
has a terminal opening, the ostiole.
The divisions of the zygote nucleus in the auxiliary cell are always mitotic
 174 A TEXTBOOK ON ALGAE

and a mass of uninucleate gonimoblasts grows out from the upper side of the
auxiliary cello Terminal cells of gonimoblasts then form carposporangia.
Meanwhile, the auxiliary cell, the supporting cell and the cells of tbe sterile

supporting cell \j

@~B
~VIh
\)()O
@'W
cHf
000
nnn
A

Fig. 10-5. Po[ysiphonia jiexicaulis. A-F, longitudinal sections through successive stages
of developing procarp (protoplasmic connections not dtawn). (After Smith, 1955.)

filaments fuse to form a large placental cell which nourishes the developing
carposporophyte. The carposporophytc with the pericarp and placentaI cell,
is known as the cystocarp (Fig. 10-6 A).
Carpospores are diploid and are produced singly within carposporangia.
Tbc carpospore germinates into a new diploid free living plant, the tetra-
sporophyte. During germination, it divides transversely to form a four-celled
uniseriate filament whose lowermost cell develops into a rhizoid and the
uppermost as an apical initi al of the tetrasporophyte. The tetrasporophyte
initial functions exactly like th e apical ceIl of the sexual plants, giving rise
 RHODOPHYTA 175

to a branched thallus composed of central and pericentral cells (Fig. 10-6 B).
The tetrasporophyte forms stalked tetrasporangia from certain pericentral
cells (Fig. 10-6 C); tbe pericentral cell divides vertically into an outer cover

tetrasporangium

c
A

covercell
axial filament

t.etrasporangiurnLL........+-'-':-fl

Fig. 10-6. A. PO[Ysiphonia nigrescens, cystocarp; B. section through apex of tetrasporo-

phyte; C. P. /astigiata. transverse section of developing tetrasporangium; D, old tetra-
sporophyte in surface view showing tetrasporangia. (Figs. B. D after Smith. 1955.)

cell and an inner sporangial mother cell. The former may not divide further,
but the latter segments transversely into a lower stalk cell and an upper
sporangial cell. The sporangial nucJeus undergoes reduction division
followed by cJeavage of the sporangial protoplast into four tetrahedral
uninucJeate spores called tetraspores (Fig. 10-6 D).
In Polysiphonia there is a pair of sex-determining allelic genes. The
alleles segregate at the first meiotic division and give rise to spores, one half
of which form male plants and the other half female plants.
176 A TEXTBOOK ON ALGAE

Life Cycle
The sequence of somatie phases in the li fe cycle of P. denudata has been
studied in cultures (Edwards, ] 968). The plants are dioecious, and the
diploid parasitie carposporophyte results from mitotie divisions of the zygote
nucleus in the auxiliary cello Carpospores germinate into diploid tetra-
sporophytes which are free living and are morphologically identieal witb
sexual plants. Each tetrasporangium of the tetrasporophyte forms four
haploid spores by reduetion division, two ofthese spores produee male plants
and the other two give rise to female plants. On the basis of alternation
between morphologieally identieal free living generations, tbe life cycle of
Polysiphonia is isomorphie. Cytologieally, it is diplobiontie, and aeeording
to the number of generations, including the parasitie earposporophyte, it is
isomorphie.

TEST QUEST/ONS

1. Can you grow Po!ysiphollia in freshwater and BatracllOspermum in marine habitats 1

Give reasons for your answer.
2. What advantage does a red al ga derive from the produetion of a carposporophyte 1
3. Why is the carposporophyte regarded as a parasitic generation 1
4. How do Bangioideae and Florideae differ in thcir post-fertilization changes 1
5. When the photosynthetie pigments (phycoeyanin and phycoerythrin) of red algae
are water soluble, why do they not leaeh out of the cells when the algae are grow-
ing in aquatic habitats?
6. What probable function do the pit connections of red algae serve 1
7. While most of the marine Rhodophyta are red, those growing in freshwater are
usually bluish-green, violet or purpie. What explanation can you offer to account
for this disparity ?
8. Grijfitlzsia and Vauclzeria both possess coenocytic cells, yet they are incIuded in
different phyla. Why 1
9. Compare thc post-fertilization changes in Rhodophyceae and Ascomycetes.
10. What arguments can you advanee to show that Conchocelis and Challtrallsia are
the prostrate systems and Porphyra and ßatrachospermum the ereet systems oftwo
heterotrichous organisms 1
11. Wh ich features of post-fertilization changes have been employed in the cIassifica-
tion of Florideae and why?
12. Algae growing in exposed and sunny habitats are generally greenish or bluish-
green whereas those found submerged in deep waters are usually red. How ean
you aecount for Ihis differenee 1
13. Compare the Iife eycIes of Batrachospermum and Po!ysiphollia.
14. In what rcspecls do red algae resemble, and differ from, blue-green algae 1
 RHODOPHYTA 177

]5. Differentiate between: (a) oogonium and carpogonium, and (b) spermocarp and
cystocarp.

SELECTED REFERENCES

Brody, M. and VaUer, A. E., Observations on cellufar structures of Porphyridium

cruentum, J. Biochem. Biophys. Cyto/., S, 289-92 (1959).
Edwards, P., The Iife-history of Po/ysiphonia denudata (Dillwyn) Kützing in culture,
J. Phyco/., 4, 35-37 (1968).
Fogg, G. E. and Collyer, D. M., The accumulation of fats as a characteristic of certain
classes of algae, Proc. VIII Internat. Bot. Congr., 17, 28 (1954).
 11
Field and Laboratory Techniques

FI ELO STU 01 ES

SOlL ALGAE
For field studies of distribution and abundance of soil algae, the general
ecological methods as used for higher vegetation are applied with appropriate
modifications.
In tropical countries many blue-green algae, e.g., Cylindrospermum,
Nostoe, Tolypothrix, Scytonema, Camptylonema and Aphanotheee form
characteristic patches or strata on the surface of soils of grass lawns, fallow
agricuItural lands, footpaths and crop fields. Their relative growth or
abundance is estimated by means of microquadrats or microtransects.
Microquadrats are laid down by placing a 20-cm square frame in the
field and drawing a corresponding outline on a graph paper. The contours
and shapes of algal strata caused by different species are then drawn to scale
on the graph paper.
If the topography of the habitat is uneven, a microtransect may yield
more reliable information than a microquadrat. A transect is usually chosen
to record some transition in vegetation. It is plotted by Iaying astring and
then locating the algal patches in quadrats laid at fixed points along the
transect. The actual contour of the habitat along the string and the location
of different patches are then plotted to an appropriate scale.
The point frame centre method for the study of higher vegetation may
likewise be appIied to the ecological study of algae.
More advanced students might Iike to study the autecology of some
selected perennial species. For this purpose, two or three suitable areas
exhibiting the growth of the alga should be selected. Collections of alga
as weH as the soil should be made at regular, weekly or fortnightly,
intervals.
Depending on the size of the stratum formed, blocks of soil of a certain
fixed size (2,5 or 10 mm 3) can be cut by means of a sharp blade and the
algal growth estimated by extracting the pigments from the block in a fixed
volume of 80% acetone. Pigments should be drawn out in at least three
changes of acetone to ensure complete extraction. Measurement of optical
density of the pigment extract in a cuvette of fixed path length in a spectro-
photometer set at a wavelength of 490 nm (corresponding to myxoxanthin),
against 80% acetone as the reference solvent, will give a rough estimate of
blue-green algae present in the original block. For other algae, wavelengths
 FJELD AND LABORA TORY TECHNJQUES 179

giving maximum absorption of some characteristic pigment, or of chloro·

phyll-a, may be used.
The algal growth should also be correlated with variations in soil charac·
teristics by regular soil analysis. Simple and elementary soil tests are des-
cribed here.

Moisture and Organic Matter Content

In a clean crucible weigh accurately about 5 gm of the soiI. Place it (lid
removed) in an oven maintained at 105°C for 24 hr. Replace the lid and
transfer the crucible to a desiccator until cooled to room temperature.
Reweigh. Express logs in weight on a dry weight percentage basis. This
represents the moisture content.
Expose the crucible containing dry soil at first to gende and then to
strong heat until it glows red. Cool to room temperature in a desiccator
and reweigh. The loss in weight this time represents the organic matter
content of the soiI. It is expressed as a percentage of the air dry soil.

pH
Prepare a 1 : 3 or 1 : 5 suspension of soil in glass-distiIIed water and shake
vigorously for 10 min after adding 2-3 gm of barium carbonate (BaC03).
Allow to settle for 5 min or filter, then pipette 10 ml of the cIear supernatant
and transfer it into a clean test tube. Add 0.2 ml of B.D.H. Universal
indicator, shake, and read pH by matching the colour developed with the
colour chart on the indicator bottle.

Nitrate Content
Shake vigorously for 15 min a 1 : 3 soil water suspension, aIIow to setde for
5 min, withdraw one drop of clear supernatant and to it add 7 drops of a
0.2% solution of diphenylamine in concentrated nitrogen-free sulphuric acid
(H2S0,,). Development of a blue colour indicates the presence of nitrates.
Express the intensity of colour, corresponding to nitrate content, on an
arbitrary scale of 1-4.

Carbonate Content
In a test tube containing 1 gm of soil, pour 5 ml of dilute hydrochloric acid
(HCI). An effervescence indicates the presence of carbonates in soH. Tbe
degree of effervescence, correspoding to carbonate content, may be expressed
arbitrarilyas low, medium or high.

Base·Deficiency
Add 5 ml of a freshly prepared Comber's reagent (saturated alcoholic solu·
tion of ammonium thiocyanate) to a test tube containing 5 gm of soi!.. If
a red colour develops, it indicates that the soi! is base-deficient. The intensity
of colour, proportional to base-deficiency, may be recorded on an arbitrary
+ + + + +.
scale of to
Certain anaerobic reducing soils, however, do not give any reaction with
180 A TEXTBOOK ON ALGAE

the Comber's reagent because the iron is present in the ferrous state. Hence,
if no red colour develops following addition of reagent to soil, then a drop
of hydrogen peroxide (H202) should be added to the mixture so as to oxi-
dize ferrous to ferric iron. This oxidation results in the development of a
colour. The change in colour intensity is primarily a function of the con-
centration of ferric iron present in the sampie. Thus this test provides a
simple method of finding out whether the iron present in the soil sampie is
in the reduced or oxidized state.

PHYTOPLANKTON
SampIes of free-floating aquatic algae (the phytoplankton) may be collected
from lakes, ponds, rivers and tanks by means of a plankton net. The rela-
tive proportions of different species represented in a plankton sam pie may
be estimated by counting them in a haemacytometer. Freshwater phyto-
plankton may contain unicellular, colonial and simple filamentous algae,
mostly beionging to the Chlorophyceae, Cyanophyceae and BaciIIariophyceae.
Marine phytoplankton is generally rieh in diatoms and members ofthe Dino-
phyceae, though sometimes blue-green algae are also found.
In addition to phytoplankton, most sam pies will contain zooplankton.
Two general characteristics ofplanktonic organisms should be noted: (1) they
are smaII and free-swimming by means of flageIIa or free-floating; and
(2) they have a large surface-volume ratio.
Freshwater phytoplankton, collected from reJatively 'clean', nutrient-defi-
cient or oligotrophie waters, exhibit a great diversity of algal species, though
the concentration of each species or of the algae as a whole is very low .. In
contrast, poIluted ponds and tanks, lakes or rivers that are rich in dissolved
nutrients or are otherwise eutrophie, may at certain times, especially during
summer, have luxuriant growth of one, or rarely a few species of aJgae, con-
stituting what are known as "water blooms". Such blooms impart their
characteristic colour to the water whieh appears like "pea-soup". Indeed
the appearance of a bloom of the blue-green alga Microcystis aeruginosa in
a lake or pond is regarded as indicative of pollution. One does not need a
plankton net to collect such bloom algae; merely examining a drop of the
bloom water under microscope will reveal thousands of cells or colonies of
the alga.
The appearance of particular blooms, and the specific composition and
proportion of different algae in a plankton sam pie, often show a marked
correlation with physical and chemical factors of the environment such as
pH of water, light intensity and availability (visibility), temperature, and
nitrate, phosphate, oxygen or silica content. Methods of water analysis are
beyond the scope of this book but the interested student may consult the
works of Mackereth (1963), American Public Health Association (1965) and
Strick land and Parsons (1965).
 FJELD AND LABORATORY TECHNIQUES 181

COLLECTION

As far as possible, algae should be collected fresh from nature and examined
in the Iiving state.

SOlL ALGAE
Commonly confined to the surface layers, soil algae should be collected by
means of a diminutive rectangular shovel having a sharp blade. If the soil
is damp, a sm all block can be removed directly by cutting with a scalpeJ.
Algae growing attached to tree barks, damp walls, or other such substrata
may Iikewise be collected by scraping with a scalpeJ.
It is often difficult to separate algae from soil particles. To obtain rela-
tively clean sam pIes, keep the soil block in a petri dish and add enough water
to saturate it. Place a few coverglasses on the soil block and leave the dish
open until the excess water has evaporated. Cover the dish with its lid and
keep it in a north window. Within one or two days, the motile algae in the
soil will creep up on the undersides ofthe coverglasses and begin to multiply.
During the next few days, some other algae will also start growing on the
coverglasses which can be removed periodically, placed on a drop of water
on a slide, and examined under a microscope.

PHYTOPLANKTON
These can be collected by towing a plankton net of fine bolting silk. Cer-
tain smaller forms (nannoplankton or microalgae), however, are not retained
in the net. These are collected by subjecting a sam pIe oflake water to gentle
filtration through a Millipore membrane filter of 0.45 or 0.80 I" pore size.
Reed sterns, twigs of submerged weeds, soH sampies and bark or stones
may be collected and placed in polythene bags, tins or cartons. Add just
enough water in the bottle to ensure a saturated atmosphere when it is closed.
On return to the laboratory, the bottles should be opened and sam pIes exa-
mined as soon as possible. Water sampIes should preferably be analyzed
first and soiI or bark algae observed last.
If it is intended to keep algae alive for any length of time, there should
be no overcrowding. The larger the mass of algae gathered, the sooner they
would die. When a concentrated sam pIe of algae is obtained by means of a
plankton net, some water should also be collected simultaneously. The plank-
ton coneentrate should be appropriately diluted with this water in the labo-
ratory, otherwise the algae will tend to die quickly. Another precaution is
to refrain from plaeing them indiscriminately in any kind of water, espe-
cially tap water, whieh is usually highly chlorinated or sometimes contains
toxie amounts of zinc or copper.

Concentration and Enumeration

1. Allow a 250-ml sampIe of formalin-preserved phytoplankton to stand
182 A TEXTBOOIC ON ALGAB

undisturbed in a measuring eylinder for a day. Then earefully siphon

off most of the supematant. Swirl the sediment and the remaining
fluid to make a homogeneous suspension.
2. Pour the suspension into a 25-ml volumetrie fiasko Rinse the eylinder
with distilled water twiee and add the rinses into the flask. Make up
to indieated mark by addition of distilled water. Mix thoroughly.
3. Insert a Whippie eyepieee mierometer into the oeular lens of a miero-
seope. This micrometerha~ its field of view divided into 100 squares,
with eaeh square further subdivided into 25 smaller squares. The area
of each square can be determined under the low power ofa mieroscope
by calibrating against a stage micrometer.
4. Take a Sedgwick-Rafter, eounting ehamber and carefullyadd 1 mlof
plankton concentrate from the volumetrie ftask in such a way that the
chamber is just eompletely filled without any air bubbles. Apply the
coversIip suppIied with the chamber. Gently plaee the ebamber on
the mieroseope stage and let it rest for 10 min so as to allow the algae
to settle at the bottom of the chamber.
S. Now count tbe various algae individually in a WhippIe field (square),
repeating the counts at least 1S times and each time making the counts
in a different, randomly chosen, square. Determine the number of in-
dividuals of eaeb species observed in 1 ml volume. The field count
volume can be ca1culated by multiplying tbe field area into I mm
(depth of chamber) into the number of squares counted. To tbis the
neeessary eorreetion regarding eoneentration faetor (X 10 in this case)
sbould be applied to determine phytoplankton eoneentration in the
original sam pie.
The foregoing method. has proved useful for the larger algae found in
phytoplankton. For smaller forms (nannoplankton), a Palmer Nannoplank-
ton counting ehamber can be used in place ofthe Sedgwick-Rafter chamber.
The Palmer chamber is 0.4 mm deep and is designed for eounting 0.1 ml
volumel>. It is calibrated against a stage micrometer under the high power
of a mieroseope (X 40 or 45) instead of at X 10 as in the Sedgwick-Rafter
chamber.

Estimation of Primary Productivity

Phytoplankton eonverts sunlight energy into ehernieal energy whieh ean be
utilized by other members of an aquatie eeosystem. Depending primarily
on the kind of algal producers present and the nutrient eoneentrations,
various ecosystems differ in respect of their primary produetivity (defined
as the rate at whieh the algal prpducers eonvert solar energy into ehemieal
form).
A simple method of measuring primary productivity involves the estab-
lishment of small aquaria containing some algal eulture medium (vide infra)
and inoeulated with a few speeies, preferably unieellular, of freshwater algae.
These are then grown for 2-3 weeks in light of known intensity. If any
(filamentous) algae grow attaebed to the bottom or side walls of tbe aqua-
 FJELD AND LABORATORY TECHNIQUES 183

rium, theycan be gently scraped by means of a rubber-tipped glass rod.

Water lost through evaporation should be replaced with distilled water daily.
The algal suspension should be aerated 2 or 3 times every day (or continu-
ously) by bubbling air into it. The following schedule is recommended:
1. Withdraw 20-mI aliquots of algal suspension from aquarium and cen-
trifuge in clean tubes at about 3000g for 15 min. Discard supernatants.
2. Suspend pellets in 2~3 ml of distilled water and transfer quantitatively
into accurately weighed clean and dry crucibles.
3. Dry crucibles in an oven at I05°C for 6 hr, withdraw, cool to room
temperature in a desiccator and weigh. Record dry weights of algae
by subtracting crucible weights.
4. Place the same crucibles in a muffle furnace at 60ü°C for 4 hr. Remove,
cool in desiccator and reweigh. The difference between dry weight and
ash weight in each ca se is an estimate of the biomass of the organisms
present.
5. Repeat the above procedure at 24-hr intervals in order to estimate
changes in biom ass, representing primary production.
6. Calculate as folIows:
Daily production
.
in =
Aquanum
. vo Iurne (I)
m X
Daily production in sampIe (gm)
aquarium (gm) SampIe volume (mi)
Suitably modified, this method can also be adapted for estimating daily
fluctuation in primary production in natural habitats such as ponds and
lakes. The observed changes in biomass should be carefully correlated with
changes in algal populations.

MARINE ALGAE

Seaweeds and other marine algae are best collected during a low tide. A
rocky shore is most suitable for such collections since it supports a wide
variety of different species occurring in rock-pools or as lithophytes.

PRESERVATION

When it is intended to store algae in the laboratory for subsequent morpho-

logical studies, they may be kiIled and preserved in a 4% solution offorma-
lin (prepared by adding 4 ml of 40% formalin to 96 ml of distilled water;
40% commercial formalin is regarded as 100% for purposes of calculation).
For preserving aquatic algae, appropriate quantity of 40% formalin may be
added directIy to the sampIe so as to obtain a final concentration of 4%.
Many terrestrial or subaerial algae can be preserved dry and stored in
paper envelopes for long periods without any apparent damage or loss of
viability.
For maintaining the algae in their natural (green) colour, any one of the
following solutions may be employed. The algae are immersed in the pre-
servative for a few days and then transferred to formalin acetic alcohol
(FAA)solution.
184 A TEXTBOOK ON ALGAE

I. 50% ethyl alcohol 90 ml

40% formalin 4 ml
Glycerol 3 ml
Glacial acetic acid 3 ml
Cupric chloride (CuCh) 9.5 gm
Uranium nitrate (UN03) 1.5 gm
Although suitable for most green algae, this preservative can also be used
for blue-green algae if 10 gm of copper acetate is substituted for the cupric
chloride and uranium nitrate.
2. Cupric sulphate (CUS04' 5H20) 0.25 gm
Water 38 ml
When completely dissolved, add
Glacial acetic acid 4 ml
40% formalin 8 ml
95% ethyl alcohol 50 ml
3. Po tass ium chrome alum 10 gm
40% formalin 6 ml
Distilled water 500 ml
These solutions can also be used when certain rare algae have to be ex-
hibited in their natural colours in a museum.
It is sometimes possible to preserve living algae on a slide for a few days.
This is achieved by cutting a lens paper somewhat sm aller than a square
coverglass, and making a square hole in the centre and placing it on a slide
having a drop of water. The living al ga is then placed in the water drop in
the cent re of the square and covered with a coverglass. Since the lens paper
is very absorbent the whole preparation can be easily kept moist. Alter-
natively, the alga can be placed in a few drops of water in a hollow cavity
slide. A coverglass is then applied and its edges sealed with paraffin oil or
Gold Size.

PREPARATION OF HERBARIUM SHEETS

It is fascinating to prepare herbarium sheets ofseaweeds and the larger fresh-

water algae. The advantage of this method is that the specimen is preserved
in its natural habit and colour.
Herbarium sheets of most marine algae, especially those having mucila-
ginous or gelatinous thalli (e.g., De/esseria, Plocamium, Gracilaria, Porphyra
and V/va) can be prepared with great ease. The specimen is floated in an
enamel tray containing freshwater or seawater, as the case may be, and a
piece of moderately thick herbarium sheet inserted in the water below the
specimen. The al ga is then spread and the tray tilted gently while the speci-
men is held on the sheet (Fig. 11-1). If the specimen is mucilaginous, it
will stick to the sheet and any excess water on the latter may be drained off
by keeping the sheet on an incIined plane for some time. The sheet is
then placed between two dry newspapers or blotting papers and after a
number of herbarium sheets of different algae have been prepared in this
 FJELD AND LA BORA TORY TECHNIQUES 185

way, they are all pressed in a herbarium press. After 24 hours the wet
newspapers are replaced by fresh, dry sheets and pressed again. Five or
6 such changes are usually sufficient to absorb all traces of water from the
sheets and specimens may then be mounted and labelIed.

Fig.11-1. Diagram iIIustrating the preparation of a herbarium sheet of an alga.

This method may be modified for non-sticky algae by affixing the speci-
men on to herbarium sheet by means of a few small pie ces of cellotape.

PERMANENT PREPARATIONS
Permanent slides of algae have a limited use and as pointed out earlier,
observations on Iiving specimens should be preferred to making permanent
preparations. Some of the more common methods are described here.

Schedule I
This method may be used for both freshwater and marine algae. It is
suitable for morphological study and identification, but not for cytological
study.
I. Place alga on a slide in a drop of freshwater or seawater.
2. Add a small drop of 40% formalin to fix. Drain out excess water.
3. Place a small lump of glycerine jelly on the alga. (Glycerine jelly is
prepared by dissolving 5 gm of gelatine in 30 ml ofwater by gentle heat
and then adding 0.125 gm phenol and 35 ml of glycerol.)
4. Transfer the slide to an incubator or oven (at 60°C) for a few minutes
for the jelly to melt. Spread the algal material appropriately.
5. Apply circular coverglass. Keep the slide again in the oven for a short
while. Wipe off excess jelly from around the coverglass and seal it with
Gold Size. Store ßat.
186 A TEXTBOOK ON ALGAB

Schedule 11
1. PIace al ga on a slide in a tiny drop of water, fix by adding a HttIe of
40% formalin and then add a sm all drop of 10% glyceroI.
2. Lcave the slide in a warm, dust-free atmosphere for a few days so that
the glycerine may concentrate.
3. Add a drop of glycerine jelly and apply a warm coverglass.

Schedule 111
1. The fresh material may be fixed in any one of the following solutions:
(i) Chrome-acetic Fixative
10% aqueous chromic acid 2.5 ml
10% aqueous acetic acid 5.0 ml
DistiIIed water 92.5 ml
(ii) Dioxan Fixative
Dioxan 50 ml
40% formalin 5 ml
Glacial acetic acid 5 ml
Distilled water 50 ml
(iii) Chrome-osmo-acetic Fixative
Chromic anhydride 1 gm
Glacial acetic acid 3 ml
1% aqueous osmic acid I ml
DistiIIed water 100 ml
2. Wash the fixed material several times in tapwater. Dehydrate gradually
by passing the material through a large number .of grades of ethyl
a\cohol (3%,5%,8%, 12% ... absolute alcohol,2 changes in each grade).
In the Iower grades the material may be kept for 3 min; in the higher
grades for about 5 mine After absolute alcohol the alga is mounted
directly in a drop of Euparal on the slide and acoverglass appIied.
3. If stained specimens are required, any one of the following stains may
be used:
(i) Aniline Blue (1% solution in 90% ethyl alcohol) used for fila-
mentous green algae; stain for 5 min after 85% ethyl alcohol
stage.
(ii) Erythrosine Bluish (1 % solution in absolute ethyl alcohol) used
for staining gelatinous envelopes or sheaths; stain for 30 sec after
95% ethyl a\cohol.
(iii) Light Green (0.2 % solution in 90% ethyl alcohol) used for
cellulose cell walls; stain for 30-60 sec after 85% ethyl alcohol.
(iv) Congo Red (0.2% solution in absolute ethyl alcohol) used for
staining mucilage sheaths of Cyanophyta; stain for 60 sec after
95% ethyl alcohol.

Schedule IV
1. Smear the slide with a thin film of Mayer's ·albumin and allow it to dry
for a few minutes. Place material on this in a drop ofwater and titt the
 PIELD AND LABORATORY TECHNIQUES 187
slide to drain out water. Pass slide momentarily over a spirit flame.
2. Holding the slide in the left hand, pour 30% ethyl alcohol over it from a
dropping bottle and immediately drain it out. Rinse the material simi-
larly with 50%, 70%, 90% ethyl alcohol and give 2 changes of absolute
ethyl alcohol. After the last change, immediately place a drop of
Euparal on the material and apply a coverglass.
If necessary, the material may be stained at the appropriate stage.

Schedule V
This method is employed for making permanent slides suitable for cyto-
logical studies.
1. Fix the material in any of the fixatives described in Schedule IH, or in a
mixture containing equal volumes of glacial acetic acid and absolute
ethyl alcohol. Material should be immersed in the fixative for 10-
15min.
2. Wash in 3 changes of water.
3. Mordant in 1.5% aqueous iron alum solution for 10-60 sec.
4. Wash thoroughly in running tap water for 2-3 mine
5. Stain the material in a drop or two of acetocarmine on a slide, or, in
the case of unicellular forms, in a centrifuge tube; warm the slide gently
over a spirit flame and apply a coverglass.
6. Dip the slide and coverglass horizontally in a petri dish containing 95%
ethyl alcohol and allow them to remain there till the coverglass separates
from the slide. Carry the coverglass and slide through two other petri
dishes containing absolute ethyl alcohol, mount in Euparal and apply
the coverglass again.
For further details of this method, and its modifications for application
to different kinds of algae, consult Godward (1966).

Schedule VI
This method has been found useful for the Fucaceae (see Evans 1962;
Godward 1966).
1. Fix material in 1 : 3 glacial acetic acid, absolute ethyl alcohol mixture
for 10-18 hr; wash thoroughly in water.
2. Treat small fragments of thallus with alM solution of lithium chloride
for 10-15 min; wash in running water for 15 mine
3. Squash in water under a coverglass and irrigate the material with aceto-
carmine cc,utaining 3 drops of ferric acetate per 25 mI.
4. Warm the slide gently over a spirit flame and squash again; absorb away
the excess stain by means of a dry blotting paper and ring the prepara-
tion with glycerine jelly.
Methods for the preparation of permanent slides of red algae have been
described in detail by Dixon (quoted in Godward, 1966).

Schedule VII
This method is suitable for many filamentous algae.
188 A TEXTBOOK ON ALGAE

1. Fix in any of the fixatives described in Schedule IH, wash thoroughly

in water.
2. Stain in Harris' haematoxylin which is prepared by dissolving 5 gm of
haematoxylin and 3 gm of ammonium alum (aluminium ammonium
sulphate) in 1000 ml of 50% ethyl alcohol by heating. To this, 6 gm of
mercuric oxide is then added and the solution boiled for half an hour,
filtered, cooled and then made up to 1000 ml in a volumetrie flask by
adding 50% ethyl alcohoI. A few (8-10) drops of HCl are then added
per 1000 ml of the solution.
3. Wash in water.
4. Examine under a microscope and if overstained, destain in acid water
and wash thoroughly in tap water.
5. Dehydrate through a graded series of 50%, 70%, 90%, and 95% dioxan
and water mixtures, allowing 30 min in each grade. Give two changes
in absolute dioxan, allowing 10-12 hr in each.
6. Mount a few filaments in a thin drop of Canada Balsam dissolved in
dioxan. Leave the slide aside for dioxan to evaporate, and apply a
coverglass.

Schedule VIII (Personal communication from G. Russell)

This method is suitable for filamentous mllrine algae.
1. Fix the material in the following fixative for 12-24 hr:
Absolute ethyl alcohol 75 ml
Glacial acetic acid 25 ml
Concentrated solution of ferric chloride 3-4 drops
2. Wash in three changes of water and store in 70% ethyl alcohol.
3. Wash in distiIIed water and arrange on a slide in a few drops of 5%
aqueous solution of sodium carbonate.
4. Warm the slide gently and then cover the material with an albumenized
coverglass.
5. Squash between two thicknesses of filter paper.
6. Invert the slide in a ridged dish containing 20% acetic acid until the
coverglass (along with the material) floats off.
7. Transfer the coverglass to a watchglass containing acetocarmine and
heat to steaming for 7-10 min.
8. Transfer coverglass to 45% acetic acid and keep in this for 1-2 min until
material becomes pale yellow.
9. Place in 95% ethyl alcohol and then give two changes of absolute ethyl
aIcohol.
10. Clear in Euparal essen ce and mount in Euparal.

HISTOCHEMICAL AND GENERAL METHODS

Temporary Fixation
Add a drop of dilute lodine Potassium Iodide (IKI) solution to a 1-2 ml
sampie of algae; this also acts as a temporary preservative. (The IKI solution
 FJELD AND LABORATORY TECHNIQUES 189

is prepared by dissolving 2 gm of KI in 20":'30 ml of water and then dis-

solving 1 gm of iodine. The solution is then made up to 100 ml with distiIIed
water.)
Algae can also be fixed and temporarily preserved by exposure to osmic
acid vapour.

General Staining
Different algae vary in their affinity for stains. More frequently employed
stains are Methylene Blue, Gentian Violet or Acid Fuchsin (up to 1% aqueous
solutions). The foJlowing simple procedure is recommended. Mount algae
in a drop of water ort a slide and apply a coverglass. Add a drop of the
stain to one edge of the coverglass and let it diffuse to the opposite edge by
removing water from the other side with a piece of dry blotting paper. In
this way a spectrum of staining is obtained, the algae near one side of the
coverglass are intensely stained and on the opposite side weakly stained.

Arresting the Movements

In the living state certain algae are actively motile or exhibit gliding move-
ments. Others may exhibit a passive Brownian movement. To slow down
such movements:
(i) add a tiny drop of IKI solution to a drop of algal suspension,
(ii) add a pinch of powdered gum arabic or dextran to a drop of a)gal
suspension and apply a coverglass,
(iii) mix a drop of chloroform water (i.e., a drop of chloroform in 5 ml
of distilled water) with a drop of algal suspension and apply cover-
glass, and
(iv) place a drop of algal suspension directly in the centre of a slide or
petri dish containing a thin sheet of 2.5% aqueous agar, apply
coverglass and observe under a microscope. This method will also
stop Brownian movement.

Flagella
Most algal flagella cannot be seen under an ordinary light microscope with-
out special staining. Following are the two commonly used methods to
ren der flagella visible:
(i) add a few particles of lead of copying ink pencil to a drop of algal
suspension, apply a· coverglass and examine, and
(ii) fix algae in dilute IKI solution or by exposure to osmic acid vapour,
apply coverglass and examine. If overstained, decolourize to desired
degree by adding dilute sodium thiosulphate.

Cell Walls
The presence or absence of a true cell wall may be ascertained by micro-
scopic observation of algal cells that had been placed in a p)asmo)ysing
solution, e.g., 10% sucrose, for a few minutes.
Tbe presence of cellulose or related compounds in a cell wall may be
190 A TEXTBOOK ON ALGAE

determined by treating the algae with Schultz's reagent (chlor-zinc-iodide)

~or 15-20 min, which gives a violet or blue colour with such compounds.
Schultz's re agent used to be regarded as a specific stain for cellulose but it
gives a positive colour response with other related compounds as weIl.
Alternativeiy, mix a drop of dilute IKI solution with a drop ofalgal sus-
pension on a slide, followed either by treatment of the mixture with a drop of
cOllcentrated sulphuric acid, or by irrigation with 70% sulphuric acid. Cellu-
lose turns blue.
Cell walls composed largely of pectic substances acquire a red colour
when stained with a fresh, dilute aqueous solution of Ruthenium Red (one
or two crystals in a small watchglass containing 2-3 rol water).

Gelatinous Envelopes and Sheaths

These are rendered conspicuous by staining with a very dilute aqueous
solution of either Ruthenium Red or Methylene Blue. Alternatively, a trace
of India ink or of Gurr's Negative Stain may be employed for the same
purpose.

Chromatophores
Algal chromatophores are rendered conspicuous by treatment of the cells
with boiling 8% aqueous silver nitrate solution for 4-5 min; this treatment
turns the chromatophores brownish-black.
According to Friedmann (I 966), the chromatophores assurne a conspi-
cuous and prominent appearance when viewed under a microscope which
has a blue filter (c.g., the Kodak Wratten Filter No. 48) inserted between
its light path. In visual work this filter has been reported to produce a
surprisingly clear image and in photomicrography its use yields sharp and
brilliant negatives. A further advantage of the blue filter is that it can be
used equally weil with different kinds of algae since most of them have a
strong yellow component (i.c., the complemelltary colour to bIue) in their
chromatophore pigments.

Nuclei and Chromosomes

These can be demonstrated by the application ofthe iron alum-acetocarmine
technique, as al ready described (Schedule V).
In temporary preparations, nuclei may sometime acquire a bluish colour
when the cells are immersed in a very dilute aqueous solution of Methylene
BIue, and then washed in water to remove the stain from the cytoplasm.

Starch and Other Reserve Materials

If a blue or violet colour develops after treatment of the material with IKI
solution, starch is indicated. Iodine in chloral hydrate gives better results
in this test.
Glycogen or cyanophycean starch generally gives a brownish colour on
treatment with iodine, whereas floridean starch gives a reddish colour with
this reagent.
 FJELD AND LABORATORY TECHNJQUES 191

Sulphated and non-sulphated polysaccharides in algal tissues may be

detected by treatment with alcian dyes. Parker and Diboll (1966) have
recommended the following schedule for localization of acid and sulphated
polysaccharides in various marine and freshwater algae:
(i) fix algae in FAA or in 10% formalin in seawater or freshwater,
dehydrate in tertiary butyl alcohol series, embed in paraffin under
vacuum, cut sections (151-' thick), and mount on slides,
(ii) stain 30-60 min in 0.5% aqueous Alcian BIue adjusted to pH 0.5
with NHCI,
(iii) wash in water, and
(iv) dehydrate, clear and mount.
The following procedure for histochemicallocalization ofpolysaccharides
in marine algae has been suggested by McCully (1970):
(i) fix 2 mm 3 pieces of seaweed (e.g., Fucus) in 10% aqueous acrolein
for 24 hr,
(i i) wash in water and dehydrate by passing through alcohol grades
(10%,20%, 30%, ... , 100%) at 0-4°C in a refrigerator,
(iii) infiltrate and embed in the resin glycol methacrylate and cut thin
sections with a good microtome,
(iv) stain sections with a 0.05% solution of toluidine blue in benzoate
buffer at pH 4.5 (alternatively, in dilute acid). Wash in tap water.
Development of red colour indicates polysaccharides.
The red colour may be presenred by air drying the sections after staining
and washing.

PREPARATION AND CLEANING OF DIATOM FRUSTULES

To study the frustules, make a diatom suspension and wash it by centrifu-
gation; resuspend material in 10-20 ml of distilled water in a 50-011 quartz
tube. Add 5-10 drops of hydrogen peroxide and expose the tube to ultra-
violet radiation (short wavelength, germicidal) for up to 2 hours. Examine
the irradiated suspension under a microscope.
If a quartz tube is not available, the peroxide containing sam pIe may be
exposed to ultraviolet light in an ordinary glass petri dish, with the lid remov-
ed during irradiation. This simple method for preparing clean frustules has
proved convenient for taxonomical studies (Swift, 1967).

CHROMATOGRAPHY OF PIGMENTS
Chromatography is a convenient technique for separating individual com-
ponents from a mixture of similar substances, e.g., amino acids, sugars and
photosynthetic pigments. The mixture is extracted in a suitable solvent and
the extract allowed to fiow over the surface of a finely divided or coarse solid
material. The different components of the mixture move in the solvent at
different speeds thus becoming separated from each other in due course.
Simple chromatographie examination of algal pigments can prove helpful
192 A TEXTBOOK ON ALGAE

in distinguishing various algal groups. The fat-soluble pigments, i.e., chloro-

phyll-a, chlorophyll-b, carotenes and xanthophylls can be easily extracted
from any alga though Chlorella is particularly suitable for this purpose.
Concentrate a large population of Chlorella by centrifugation and suspend
it in 10 or 20 ml of 80% aqueous acetone. Keep in refrigerator for 4-6
hours, filter or centrifuge to remove cell debris.
The dark-green filtrate thus obtained is the
solution of the photosynthetic pigments.
Cut a filter paper into small strips (2-3 cm
wide and 10-12 cm 10ng). Draw a pencilline
across the paper strip nearly 2 cm above the
bottom end. Develop a concentrated spot of
the pigment extract in the centre of the pencil
line (Fig. 11-2) and then dry the spot with a hot
air blower. Place a few more drops ofpigment
extract along the pencil line and dry them in
hot air. Repeat this step till the spot is con-
centrated.
Take a corked test tube having a hook
underneath the cork (see Fig. 11-2) and place in
it a few ml of a mixture of 90 parts of methanol
+ 10 parts of petroleum ether, or 5 parts of
acetone + 95 parts of petroleum ether. Hook
the paper strip to the cork in such a manner
that its bottom end just dips into the solvent
mixture (Fig. 11-2). Run the experiment for
about an hour until the solvent front just
reaches the top of the strip. Meanwhile watch Fig. 11-2. Diagrammatic
demonstration of ascend-
the separation of individual pigments in the ing paper chromatography
form of differently coloured bands along the for separation of algal
paper strip. pigments.

CULTURE

Certain morphological and reproductive stages can best be demonstrated by

suitable manipulation oflive cultures of algae. Algal cultures may either
be obtained from a Culture Collection, or raised in the laboratory. The
three major Culture Collection Centres are:
(I) The Culture Collection of Algae, Department of Botany, Indiana
University, Bloomington, Indiana, U.S.A.
(2) The Culture Centre of Algae and Protozoa, Storey's Way, Cam-
bridge, England.
(3) The Culture Collection of Algae and Microorganisms, Institute of
Applied Microbiology, University of Tokyo, Bunkyo-ku, Tokyo,
Japan.
Unialgal cultures may, however, be isolated without much difficu!ty from
 FIELD AND LABORATORY TECHNIQUES 193

fresh material collected from nature. A number of culture media have been
found suitable. The following three are likely to prove useful for the isola-
tion and muItiplication of most freshwater and soiI algae:
1. Chu No. 10 (modified), for common freshwater and soil algae:
Calcium nitrate [Ca(N03hJ 0.04 gm
Dipotassium hydrogen phosphate (K2HP04) 0.01 gm
Magnesium sulphate (MgS04'7H20) 0.025 gm
Sodium carbonate (Na2C03) 0.020 gm
Sodium silicate (Na2Si03) 0.025 gm
Ferric citrate 0.003 gm
Citric acid 0.003 gm
*As Trace elements stock solution (optional) 1.0 ml
Glass-distilled water to 1000 ml
*The A s Trace elements stock solution has the following composition in
grams per litre of glass-distilled water:
Boric acid (H3B03) 2.86
Manganese chloride (MnCh'4H20) 1.81
Zinc sulphate (ZnS04'7H20) 0.222
Molybdenum trioxide [M003(85%)] 0.0177
Cupric sulphate (CuS04'5H20) 0.079
2. Allen and Arnon 's Medium (modified) for nitrogen-fixing blue-green
algae:
Magnesium sulphate (MgS04'7H20) 0.025 gm
Calcium chloride 0.05 gm
Sodium chloride 0.20 gm
Dipotassium hydrogen phosphate 0.35 gm
As Trace elements stock solution 1.0 ml
Glass-distilled water to 1000 ml
If 0.20 gm of potassium nitrate is added, this medium will also support
the growth of many non-nitrogen-fixing blue-green algae.
3. ASM-l Medium for freshwater planktonic algae:
Sodium nitrate 170 mg
Dipotassium hydrogen phosphate 17.4 mg
Disodium hydrogen phosphate 14.2 mg
Magnesium chloride (MgCh'6H20) 40.7 mg
Magnesium sulphate 49.3 mg
Calcium chloride 22.2 mg
Ferric chloride (FeCh'6H20) 1.1 mg
Sodium ethylene diamine tetra acetate
(Na~DTA) 6.7 mg
Boric acid 2.5 mg
N'
r-
Manganese chloride (MnCh'4H20) 1.4 mg
~ Zinc chloride (ZnCh) 0.4 mg
~ Cobalt chloride (CoCh'6H20) 0.02 mg
; Cupric chloride (CuCh'2H20) 0.00014 mg
~ Glass-distilled water to 1000 mf
194 A TEXTBOOK ON ALGAE

4. Marine algae Medium:

,Smaller marine algae may be grown in either synthetic or natural sea-
water culture media. The latter have, however, been found to be more
suitable than the former. In inland places far away from sea, the synthetic
media may be used if seawater cannot be procured. The composition of a
suitable culture medium, both natural and synthetic, for growing marine
algae (e.g., Ectocarpus) is given here (see also BoaIch 1961):

Natural Sea- Synthetic

water Medium Medium
Natural seawater 1 litre
Glass-distiIIed water 1 litte
Potassium nitrate (KN03) 200 mg 200 mg
Potassium monohydrogen phosphate
(K 2HP0 4) 35 mg 35 mg
Ferric chloride (FeCb'6H20) 3 mg
Manganese chloride (MnCh'4H20) 0.2 mg
Sodium chloride (NaCI) 30,000 mg
Magnesium chloride (MgCh'6H20) 5000 mg
Calcium sulphate (CaSO.2H20) 1000 mg
Sodium ethylene diamine tetra acetate
(Na2EDTA) 20 mg
Potassium chloride (KCI) 750 mg
Potassium bromide (KBr) 15 mg
Ferrous sulphate (FeSO.7H20) 0.7 mg
Aluminium sulphate [Ah(S04hJ 0.25 mg
Cobalt sulphate (CoS04'9H20) 0.03 mg
Cupric sulphate (CuS04'5H20) 0.005 mg
Lithium chloride (LiCI'H20) 0.05 mg
Sodium molybdate (Na2Mo04'2H2Q) 2.0 mg
Rubidium chloride (RbCl) 0.5 mg
Strontium chloride (SrCh'6H20) 5.0 mg
Zinc sulphate (ZnS04'7H20) 10.0 mg
(The pH of all the above media should be adjusted to 7.0-7.5 for green
algae and 8.5-9.0 for blue-green algae.)
In all these media, phosphate should be autocIaved separately from the
other components and mixed aseptically upon cooling.
The media may be solidified with 1 or 1.5% agar. It is advisable to
sterilize agar in a small volume of water separately from the mineral salts,
to mix the two components after autocIaving, and then to pour in sterile
petri dishes.
For liquid culture, a small volume of the mineral salts medium is taken
in an Erlenmeyer flask or a test tube which is plugged with non-absorbent
cotton wool and then sterilized at 15 Ibsjin 2 for 15 min. Corning test tubes
(150 X 25 mm) with screw caps may be used if cotton plugging is desired to
be avoided.
 FJELD AND LABORATORY TECHNJQUES 195

Fresh material collected from nature is observed microscopically to see

if it is fairly unialgal or not. Some of it may be taken in a watchglass
containing sterile water and teased and separated by means of sterile needles
under a binocular disseeting microseope. The teased mass is transferred to
a rubber-stoppered test tube containing a small volume of sterile water
and shaken vigorously for 10-15 min so as to disperse the cells or filaments.
A drop or two ofthe dispersed suspension is then added to a fresh watchglass
containing a few ml of sterile water. From this, single cells or filaments are
picked up by means of a fine capillary pipette under the binoeular miero-
seope and transferred to a seeond watehglass containing water. After three
such washings, single cells or filaments are inoculated into cuIture tubes.
At least a dozen tubes, ftasks or petri dishes should be inoculated in this
way since some of the euItures may fail to grow. Out of those that would
grow, one or two may prove unialgal from the beginning; the others will
have to be discarded.
Another method is to spread a drop of algal suspension on the surface of
an agar plate and to incubate the latter for a few days until growth occurs.
The plate is then examined under a mieroseope and areas showing the
desired alga free (or remote) from unwanted organisms are selected. Small
loops of agar from these selected areas are then pieked up aseptically by
means of a platinum or nichrome wire-Ioop, or a pipette and grown in liquid
medium for subsequent examination.
Certain algae may best be isolated in water in which they grow in nature.
This water may be sterilized by passing it through a sterile MiUipore mem-
brane filter (pore size 0.45 ,,) and inoeulated with single or a few cells or
filaments of the alga eollected from the same pond. Incubation of such
crude eultures may be earried out by plaeing them in a north window, or by
iIIuminating them with a 60-watt tungsten lamp. After the euIture has
estabIished itself, it may be subeultured in a defined culture medium.
Many soil algae will grow in a biphasie soil water medium prepared by
adding 6-10 ml of tap or pond water to 2-3 gm of soH in a eotton-plugged
test tube and steriIizing in a steam sterilizer (without pressure) for a few
hours. Sometimes addition of a few ml of Chu No. 10 medium to the soH
water tubes may give better results, or the water may be entirely replaeed
by eulture medium.
Although vigorous aseptic preeautions are not needed for the erude and
elementary eulture work for demonstration purposes in a class room, yet
it is advisable that the eultures are maintained in a unialgal condition and
eontamination prevented by adequate sterilization of glassware, media,
inoeulating needles, and pipettes. For work involving unialgal. baeteria-
eontaminated euItures, use of sterile rooms or inoeulating hoods is unneees-
sary, and the inoeulations or subculturing may be performed on a working
bench in the laboratory.
Formation of zoospores, gametes, or other reproductive or vegetative
stages in certain algae may be induced by modification of environmental
and physiological conditions such as changes in intensity, quaIity or dura-
196 A TEXTBOOlC ON ALGAE

tion of light, pH, mineral balance (e.g., addition or depletion of a nitrogen

source) and exposure to dry or wet conditions. Species of Ch/amydomonas
which form Palmella stages when grown on agar plates, can be brought into
an active, motile phase by adding water. If this is done an hour before the
commencement of a practical class, drops of liquid suspension from the agar
plate can be taken for examination of living Ch/amydomonas cells or zoo-
spores. Likewise, if suspensions of cells from + and - strains of Ch/amy-
domonas are obtained in this way and mixed, stages in conjugation and sexual
fusion may readily be observed.

DEMONSTRATION OF SEXUAL CYCLE

Chlamydomonas is a good material for the study and demonstration of sexual
stages in green algae (Hoshaw, 1961). The heterothallic species C. reinhardii
and C. moewusii are especially suitable for demonstration ofthe sexual cycle.
Cultures of their + and - mating types may be obtained from the Cam-
bridge Collection or the Indiana University Culture Collection.
About a week before demonstrating sexuality, young subcultures of both
mating types should be grown separatelyon agar medium (Chu No. 10 or
soil water medium) in petri dishes (Fig. 11-3 A). When good growth has
occurred, add 15-20 ml of sterile water on the surface of the agar culture
and after a few hours, transfer 5-10 ml of the suspension into a sterile
culture ftask (Fig. 11-3 B) or tube and illuminate for 3-6 hr at 300-500 ft.c.
The cultures are then covered with a black cloth or left in the dark over-
night.
An hour before the practical class js scheduled to meet, the suspensions
are transferred to light.
One drop each of the plus and minus mating type suspensions is then
placed half an inch apart on a clean slide and the two drops are mixed by
means of a needle. As soon as the drops are mixed, clumping of gametes
starts and may be readily observed under 10w power of a microscope (Fig.
11-3 C). Numerous copulating pairs of gametes can be observed within a
few minutes of mixing the two sexes.
After observing the clumping process, add a drop of IKI solution to the
mixed suspension so as to kill the cells. Apply a coverglass and examine
under high power of a microscope to observe quadriftagellate zygotes.
Mature zygospores can be successfully obtained when inocula from the
complementary mating types are mixed and grown on the same place for
7-10 days. The plates are then stored in the dark for almost a month.
Scrape off the mature zygospores from such plates by means of a sterile
wire-loop and spread the scraping on the surface of a fresh agar plate in a few
drops of sterile water.
Invert the inoculated petri dish over a petri dish bottom containing a
few ml of chloroform to kill vegetative cells, jf any, in the scrapings. Half
aminute later, replace lid and incubate the culture plate in light for two
days. Then apply a coverglass directly on the surface of the agar and ob-
 FJELD AND LABORATORY TECHNIQUES 197

serve the different stages of germinating zygospores under high power of a

microscope (Fig. 11-3 D). MotiIe zoospores released from zygotes may be
observed after the zygospores are covered with a drop of water.

agar medium
. agar medium
•
+

cultures

clumping

~ring
~
o

~~/ .. 0
.0 :;'::::'

zoospores
:mt
0 ;' "
fusion
\ zoospore'
o 0\' zygote "'(

o +- ,~, : ' ~;;'; ,

zygote germination D
Fig. 11-3. Chlamyd(}m(}nas, sexual cycle of heterothallic species. (After Hoshaw, 1961.)

Cultures of any suitable heterothallic macrandrous species of Oedogonium

producing antheridia and oogonia on separate filaments can be used to
demonstrate stages in the sexual cycle (Hoshaw, 1961). The formation of
reproductive orsans can be easiIy induced if freshly inoculated subcultures
198 A TEXTBOOK ON ALGAB

are ilIuminated at 20-25°C, 300-500 ft.c. for a month or two, followed by

incubation in very dim light. Tbe following steps are suggested to demons-
trate sexual phases: (I) add a few ml of soil water medium in a watchglass
and place it in a petri dish; inoculate the medium with a sm all loopfu] each
of male and female filaments and mix them gently; (2) add 10-] 5 m] of a
5% aqueous NaHC03 solution to the petri dish bottom, replace Iid and keep
the who]e set-up at 25°C, 300-500 ft.c. (Fig. 11-4); the bicarbonate solution
will enrich the C02 content of the atmosphere within the dish; (3) observe

matlng dish

Fig. 11-4. Oedogonium, sexual cyc1e. (After Hoshaw, 1961.)

the filaments microscopically once every day and note the multiflagellate
spermatozoids and the process of fertilization which may take place within
three to five days after placing the mating dish in light (Fig. 11-4).
 PIELD AND LABORATORY TECHNIQUES 199

To demonstrate oospore germination, store the oospores dry for about

a year. About a week before it is intended to demonstrate germination,
the old and ripe oospores should be covered with soil water medium and
incubated in light. They should be checked daily for the liberation of
zoospores.
Sexual stages can also be readily demonstrated in the yellow-green alga,
Vaucheria sessilis, grown in soH water medium containing a pinch of calcium
carbonate. Freshly inoculated alga (in petri dishes containing liquid medium)
should be incubated at low light intensity of about 300 ft.c. at room tem-
perature. In such cultures, formation of antheridia and oogonia may be
induced by giving light-dark periods of 16 hr-8 hr, respectively. When
cultures become 12-15 days old, they may be continuously illuminated with-
out any lass of sex organs.

GROWTH ESTIMATION

The growth of algae in relation to time can be estimated by recording the

increase in cell numbers, optical density or dry weight at appropriate (fixed)
intervals. In unicellular algae such as Chlorella or Anacystis, which grow
in a well-suspended form, the cell count method is employed.

Cell Counting
This is done by means of a haemacytometer, originally devised for counting
blood cells. Instructions for the use of different kinds of haemacytometers
are supplied by their manufacturers and these should be consulted before
using a particular make. A commonly used haemacytometer is a special
type of glass slide which has a centraIJy located H-shaped groove which
gives rise to two prominent ridges. Each ridge contains a big cubical cham-
ber (LO mm long, 1.0 mm wide and 0.1 mm deep) which is partitioned
equally into 25 sm aller cubes by distinct narrow ridges, each consisting of
3 parallel lines. Some of the 25 chambers are again subdivided into 16
smaller cubes.
The length, breadth and depth of the area divided into 25 chambers are
1.0, 1.0 and 0.1 mm, or 0.1, 0.1 and 0.01 cm respectively. Hence its
volume = 0.1 X 0.1 X 0.01 cm, i.e., 10-4 cc.
Tbe volume of any one of the 25 chambers mayaIso be calculated in the
same way; it comes to 4 X 10-6 cc.
SimiJarly, the volume of one ofthe 16 smallest chambers will be25 X 10-8 cc.
Having ascertained the volumes of the chambers, the cell numbers in a
given suspension can be estimated by placing a drop on the H-shaped groove,
applying the special kind of coverglass supplied with the haemacytometer,
and counting the number of cells in the different chambers under a miero-
scope. The cell numbers per cc of the suspension can then be calculated by
multiplying by an appropriate factor.
Cell counting by means of a haemacytometer can be used to determine
the growth curve of a unicellular alga such as Anacystis nidulans. Inoculate
200 A TEXTBOOK ON ALGAE

culture flasks with aknown number of exponentially-growing cells of A.

nidulans and incubate the cultures at a constant temperature and light inten-
sity (35°C, 1000-1200 lux). Withdraw small aliquots regularly at 24-hr
interval and estimate cell numbersjml. Continue the experiment till the cell
numbers become constant or ultimately decline. Plot the 10gIO of cell num-
bers thus obtained against days after inoculation. A typical growth curve
thus obtained may consist of: (1) a lag phase, (2) an exponential phase,
and (3) a death phase (Fig. 11-5).

Stationary ~~
>.
L.
a ~
L.
o
o
~

....
>. ~
'iij
c:
t)
~
a ~
ii
\)
:;;
Co
o
L
o
6
z
t)
()

Age of Culture
Fig. 11-5. Typical growth curve oe an alga.
Optical Density Measurement
Growth estimation by optical density measurement is somewhat less reliable
than cell counting. This is because the contents or proportions of different
pigments are likely to vary with small changes in environmental factors and
these changes are reflected in corresponding fluctuations in the optical
density.
Tbe opticaI density is generally determined in a photoelectric colorimeter
(e.g., Bausch and Lomb Spectronic 20, Klett and Summerson colorimeter,
and Zeiss Spekol spectrocolorimeter) set at an appropriate wavelength.
Unicellular algae may be grown in optically matched test tubes of a size
suitable for use in a given colorimeter. The optical density of the culture
tubes can then be measured directly, using sterile medium (uninoculated) as
tbe reference solvent. Since the increase in growth of the alga, at least up
to tbe end of tbe exponential pbase, is reflected in a corresponding increase
in tbe opticaI density of its pigments, tbis method can be employed to esti-
mate growtb of the alga.
If tbe optical density of tbe culture tul;>~~ ~annot be measured directly,
 FIELD AND LABORATORY TECHNIQUES 201

then 2-3 ml aliquots may be withdrawn aseptically from the culture tube or
fiask, and their optical density measured in cuvettes of 1 cm path length.
An appropriate cell number versus optical density calibration curve, at
the particular wavelength used, must be plotted in order to correlate optical
density with cell number.
Though somewhat unreliable, this method has the great advantage that
growth in a large number of cultures can be estimated in a short time.
This method can also be applied to estimate the optical density of pig-
ment extracts of algae instead of that of whole cells.

Dry Weight Determination

The growth of such algae as do not grow in homogeneous suspensions (e.g.,
filamentous forms) may be determined by harvesting the entire contents ofa
culture tube or flask by centrifugation. The algal pellet is suspended in dis-
tilled water, shaken for a few minutes and recentrifuged. The washed mate-
rial is then quantitatively transferred to a weighed crucible. The crucible
is later placed in an oven at 100°C for 10-12 hours and dried to constant
weight. After removal from the oven, it is allowed to cool to room tem-
perature in a desiccator and reweighed. The dry weight of the aIga can
thus be calculated.

TEST QUEST/ONS

1. When algae can be cultured and studied in the laboratory, what is the need to study
them in the field ?
2. Why is it not possible to make accurate counts of the filaments of Ulothrix or Spiro-
gyra in a haemacytometer? Explain.
3. What are oligotrophie, mesotrophie and eutrophie lakes? Name some important
faetors that govern the distribution of aquatie algae.
4. What is the praetical use of algal herbarium sheets and of formalin-preserved
algae?
5. In what ways do formalin-preserved Ulothrix, Spirogyra and Oedogonium differ
from the respective frcsh (living) forms ?
6. How docs a bluc filter hclp in thc visual observation or microscopic study of the
algal chromatophores?
7. Compare the advantages and disadvantages of sterilization of algal culture media
as carried out in an autoclave, a hot air oven, and a Millipore membrane filter.
8. Differentiate between unialgal and pure cultures. What is the importance ofaxenic
and clonal cultures in algal research and for purposes of classroom demonstrations?

SELECTED REFERENCES

Anonymous, Standard Methods for the Examination of Water and Wastewater, 11th
edn., American Public Hcalth Association, New York (1965).
202 A TEXTBOOIC ON ALGAB

BoaIch, G. T., Studies on Ectocarpus in culture. 11. Growth and nutrition of a bacteria-
free culture, J. mar. biol. Ass., U.K., 41, 287-304 (1961).
Evans, L. V., Cytological studies in the genus Fucus, Ann. Bot., N.S., 26,345-60 (1962).
Friedmann, 1., Microscopy of algal chromatophores, Phyco[ogia, 6, 29-36 (1966).
Hoshaw, R. W., Sexual stagesof three green algae for laboratory study, Am. Biol. Teach.,
23,489-99 U961).
Mackereth, F. J. H., Some Methods of Water Analysis for Limnologists, Freshwater
Biological Association Scientific Publication No. 21, Ambleside, England (1963).
McCully, M. E., The histologicallocalization of the structural polysaccharides of sea-
weeds. In Frederick, J. F. and Klein, R. M. (eds.) Phylogenesis and morpho-
genesis in the algae, Ann. N. Y. Acad. Sei., 175, 702-11 (1970).
Parker, B. C. and Diboll, A. G., Alcian stains for histochemicallocalization of acid and
sulfated polysaccharides in algae, Phycologia, 6, 37-46 (1966).
Strickland, J. D. H. and Parsons, T. R., A Manual of Sea Water Analysis, 2nd edn.,
Fisheries Research Board Canada Bulletin No. 125, Ottawa (1965).
Swift, E., Cleaning of diatom frustules with ultraviolet radiation and peroxide, Phyco-
logia, 6, 161-63 (1967).
 Suggestions for Further Reading

Anonymous, Environmental requirements of blue-green algae, Proc. Sympos. Federal

Water Pollution Control Administration, Corvallis, Oregon, U.S.A. (1967).
Anonymous, 1st Internat. Sympos. Taxonomy and BioJogy of Blue-green AJgae, Univ.
Madras (1970).
Bold, H. C., Morphology of Plants, Harper and Row, New York (1967).
Boney, A. D., A Biology of Marine Algae, Hutchinson & Co. (publishers), London
(1966).
Carr, N. G. and Whitton, B. A. (eds.), The Biology of Blue-green Algae, Blackwell,
Oxford (1973).
Chapman, V. J., The Algae, Macmillan, London (1962).
Chapman, V. J., Seaweeds and Their Uses, 2nd edn., Methuen, London (1970).
Chauhan, V. D. and Thivy, F., On the occurrence of some Sargassum species in Gujarat,
Phykos, 3, 19-25 (1964).
Dodge, J. D., Fine structure and phylogeny in the algae, Sei. Prog., Lond., 61, 257-74
(1974).
Fogg, G. E., AJgal Cultures and Phytoplankton Ecology, Univ. Wisconsin Press,
Madison (1965).
Fott, B., Studies in Phycology, E. Schweizerbart, Stuttgart (1969).
Frederick, J. F. and Klein, R. M. (eds.), Phylogenesis and morphogenesis in the algae,
Ann. N.Y. Acad. Sei., 175,413-781 (1970).
Frey-Wyssling, A. and Mühlethaler, K., Ultrastructural Plant Cytology, Elsevier,
New York (1965).
Fritsch, F. E., The Structure and Reproduction of the Algae, Vols. I & 11, Cambridge
Univ. Press, London (1935, 1945).
Fuhs, G. W., The NucJear Structures of Protocaryotie Organisms (Baeteria and Cyano-
phyeeae), Springer Verlag, Berlin (1969).
Godward, M. B. E. (ed.), The Chromosomes ofthe Algae, Edward Arnold (Publishers),
London (1966).
Hutner, S. H. and Provasoli, L., Nutrition of algae, A. Rev. PI. Physiol., 15, 37-56 (1964).
Jackson, D. F. (ed.), Algae and Man, Plenum Press, New York (1966).
Jackson, D. F. (ed.), Algae, Man and the Environment, Syraeuse Univ. Press, Syraeuse
(1969).
Kachroo, P. (ed.), Proe. Sympos. Algology, Indian Couneil of AgricuItural Research,
New Delhi (1960).
Kavaler, L., The Wonders of Algae, John Day & Co., New York (1961).
Klein, R. M. and Cronquist, A., A consideration of the evolutionary and taxonomie
204 SUGGESTIONS FOR FURTHER READING

significance of some biochemical, micromorphological and physiological charac-

ters in the thallophytes, Q. Rev. Bioi., 42, 105-296 (1967).
Leedale, G. F., EuglenidajEuglenophyta, A. Rev. Microbio/., 21, 31-48 (1967).
Levine, R. P., Genetic dissection of photosynthesis, Science, Wash., 162,768-71 (1968).
Levine, R. P. and Goodenough, U. W., The genetics of photosynthesis and of the
chloroplast in Chlamydomonas reinhardi, A. Rev. Genet., 4, 397-408 (1970).
Levring, T., Hoppe, H. A. and Schmid, O. J., Marine Algae, A Survey of Research
and Utilization, Cram, Degruyter & Co., Hamburg (1969).
Lewin, R. A. (ed.), Physiology and Biochemistry of Algae, Academic Press, New York
(1962).
Lund, J. W. G., Planktonic algae and the ecology of lakes, Sei. Prog., Lond., 55,
401-19 (1967).
Manton, I., Rayns, D. G., Ettl, H. and Parke, M., Further observations on green
flagellates with scaly flagella; the genus Heteromastix Korshikov, J. mar. biol. Au.,
U.K., 45, 241-55 (1965).
Margulis, L., Evolutionary criteria in thallophytes: a radical alternative, Seience, Wash.,
161, 102(}"22 (1968).
Morris, I., An Introduction to the Algae, 2nd edn., Hutchinson & Co. (publishers),
London (1968).
Palmer, C. M., Algae in Water Supplies, U.S. Public Health Service Publication No.
657, Washington (1962).
Papenfuss, G. F., A review of the present system of cIassification of the Florideophy-
cidae, Phycologia, 5, 247-55 (1966).
Parker, B. C. and Brown, R. M. (eds.), Contributions to Phycology, Allen Press,
Lawrence, Kansas (1971).
Prescott, G. W., How to Know the Freshwater Algae, Wm. C. Brown Co., Dubuque
(1954).
Prescott, G. W., The Algae: A Review, Thomas Nelson & Son, London (1969).
Pringsheim, E. G., Pure Cultures of Algae, Cambridge Univ. Press, London (1949).
Purvis, M. J., Collier, D. C. and Walls, 0., Laboratory Techniques in Botany, 2nd edn.,
Butterworths, London (1966).
Randhawa, M. S., Zygnemaceae, Indian Council of Agricultural Research, New Delhi
(1959).
Rosowsky, J. R. and Parker, B. C. (eds.), Selected Papers in Phycology, T,Jniv. of
Nebraska, Lincoln (1971).
Round, F. E., The Biology of the Aigae, Edward Arnold (Publishers), London (1965).
Schwimmer, M. and Schwimmer, 0., The Role of Algae in Plankton and Medicine,
Grune and Stratton, New York (1955).
Singh, R. N., Role of Blue-green Algae in Nitrogen Economy of Indian Agriculture,
Indian Council of Agricultural Research, New Delhi (1961).
Smith, G. M. (ed.), Manual of Phycology, Chronica Botanica Publishing Co., Waltham.
Mass. (1951).
 SUGGESTIONS FOR FURTHER READING 205
Smith, G. M., Cryptogamic Botany, Vol. I, Algae and Fungi, 2nd edn., McGraw-HiIl
Book Co., Inc., New York (1955).
Stanier, R. Y. and van Niel, C. B., The concept of a bacterium, Arch. Mikrobioi., 42,
17-35 (1962).
Stein, J. R. (ed.), Handbook of Phycological Methods, Cambridge Univ. Press, London
(1973).
Stewart, W. D. P., Nitrogen fixing plants, Science, Wash., 158, 1426-32 (1967).
Stewart, W. D. P., Algal fixation of atmospheric nitrogen, Plant and Soil, 32, 555-86
(1970).
Taft, C. E., Water and Algae-World Problems, Educational Publishers Inc., New York
(1965).
Tamiya, H., Cell differentiation in Chlorella. In Fogg, G. E. (ed.) Sympos. Cell Differ-
entiation, Soc. E"ptl. Bioi., 17, 188-214 (1963).
Tiffany, L. H., Algae, the Grass of Many Waters, 2nd edn., Charles C. Thomas, Spring-
field, IIlinois (1958).
Tilden, J. E., The Algae and Their Life Relations, Fundamentals of Phycology, Univ.
Minnesota Press, Minneapolis (1935).
Watanabe, A. and Hattori, A. (eds.), Cultures and Collections of Algae, Japanese Soc.
of PI. Physiologists, Tokyo (1966).
Zajic, J. E. (ed)., Properties and Products of Algae, Plenum Press, New York (1970).
 Glossary

acronematic (Gk. akros, tip; nema, thread). archegonium (Gk. arche, beginning; gonos,
Flagella with slender and smooth surface offspring). Multicellular female game-
and ending in a thin hair. tangium producing egg and consisting of
acyllipid. Lipids joined to acids through neck and venter; inc1udes sterile cells in
anhydride bonds. addition to fertile egg.
akinete (Gk. a, not; kinein, to move). A areolae. Cavity-Iike depressions each
vegetative cell that becomes converted covered with aperforated membrane,
into a thick-walled non-motile resting the sieve membrane.
spore; the wall of the cell becomes the asexual (Gk. a, without; sexus, sex). Lack
wall of the spore. of apparent sexualorgans.
algicide (L. alga, seaweed; caedere, to kill). autospores (Gk. aulos, self; sporos, seed).
A substance highly toxie to algae. Non-motHe spores resembling the parent
allelie genes (Gk. allelon, one another; cell in shape and structure.
genos, descent). A number of genes autotrophie (Gk. autos, self; trephein, to
occupying the same locus on a Iinkage nourish). Capable of producing the
group or chromosome. required food substances from inorganic
cx-granules (Gk. alpha, first letter in alpha- raw materials.
bet; L. granulum, small grain). Sub- auxospore (Gk. auxein, to increase; sporos,
microscopie granules rich in glycogen seed). Spores of diatoms.
and found in the cells of blue-green algae. benthos (Gk. benthos, depth of sea). Algae
androsporangium (Gk. aner, male; sporos, growing at the bottom of sea or lakes.
seed; anggeon, vessel). A sporangium ~-granules (Gk. bela, second letter in alpha-
producing androspore. bet; L. granulum, smaH grain). Granules
androspore (Gk. aner, male; sporos, seed). of proteinaceous nature confined to
Antherozoid-Iike zoospore formed singly Cyanophyceae.
in an androsporangium; produces a blepharoplast (Gk. blepharis, eyelash; plas-
dwarf male filament, as in Oedogonia- tos, formed). Granule Iying at the base
ceae. of a flagellum; gives rise to one flagellum.
aneuploidy (Gk. a, without; eu, weil; ap/oos, canaliculi (L. canaliculus, small channel).
one rold). Chromosome number Icss or Tubular canals present on the valve
more than the exact multiple of haploid surface of diatoms.
number. carpogonium (Gk. karpos, fruit; genos,
anisogamy (Gk. anisos, unequal; gameIes, birth). Femalegametangium in red algae;
spouse). Union between two morpho- consists of a swollen base and an
logically dissimilar gametes. elongated neck or trichogyne.
antheridium (Gk. anlhos, flower; idion, carpospore (Gk. karpos, fruit; sporos, seed).
dirn). Uni- or multicellular male game- Spore produced within a carposporan-
tangium. gi um of red algae.
antherozoids (Gk. ant/lOs, flower; zoon, carposporophyte (Gk. karpos, fruit; sporos,
animal; udos, form). Sperms; male seed; phyton, plant). Carpospore-pro-
gametes. ducing second generation plant of red
aplanogametes (Gk. a, not; pIanos, wander- algae; parasite on female gametophyte,
ing; gameIes, spouse). Non-flagellate arising directly or indirectly from zygote.
gametes showing amoeboid movement. cellular (L. cellula. small room). Organ-
aplanospore (Gk. a, not;p/anos, wandering; isms made of conventional units of
sporos, seed). Non-motile spore in which structure and function known as cells,
the spore wall is not derived from the each containing several proteins and both
wall of its parent cell. DNA and RNA; differ from acellular
 GLOSSARY 207

entities which have very few proteins and of the valve which separate the rows of
either DNA or RNA but not both. areolae from each other.
central nodule (L. cenlrum, centre; nodus, cryptoblasts, cryptostomata (Gk. kryplos,
knob). A wall thickening in the centre hidden; stoma, mouth). Sterile concep-
of a valve in diatoms. taeles found in Fucales.
centrlc (L. cenlrum, centre). Refers to cyanophage (Gk. kyano5, bIue; p.hagein,
diatoms which are circular in valve view to eat). Virus that infects certain blue-
and have radial symmetry. green algae.
centrosome (Gk. kentron, centre; soma, cyanophycin granules (Gk. kyanos, blue;
body). Apolar body at each pole of a phykos, seaweed; L. granu/um, small
dividing cell; the spindIe is seen to di- grain). Proteinaceous food reserve
verge from these bodies. occurring in granular form in cells of
cbromatic adaptation (Gk. chroma, colour; blue-green algae.
L. ad, to; aplare, to fit). Capacity of cyst (Gk. kystis, bladder). Resting ceIls
some algae to synthesize pigments which with a thick envelope.
are complementary to the quality of cystocarp (Gk. kystis, bladder; karpos,
available light. Also known as Gaidukov fruit). Fruiting body of red algae; aggre-
phenomenon. gate structure consisting of carpospor-
chromatophore (Gk. chroma, colour; phere- angia and sterile covering cells.
in, to bear). Plastid containing chloro- cytopharyngeal (Gk. kYlos, hollow; pharynx,
phyll-a and other pigments but not gullet). Pertaining to cytopharynx, Le.,
chlorophylI-b. gull~t, reservoir and vacuoles in Eug/ena.
cbromocentres (Gk. chroma, colour; ken- dendroid (Gk. dendron, tree; eidos, form).
Iron, centre). Feulgen positive bodies Tree-like habit with much branching.
of unknown function found on the In certain colonial algae, tree-Iike habit
chromosomes of Phaeophyta. is achieved by repeated branching of
cbrysolamlnarin. Leucosin; the food reserve mucilaginous stalks.
of Chrysophyta. derepression. Release of the inhibition of
cistron. Functional unit of gene which enzyme synthesis.
codes for a single polypeptide. developmental (F. developper, to unfold).
coccoid (Gk. kokkos, berry; eidos, form). Changes that organism undergoes during
Pertaining to habit; non-motiIe uniceIls. its vegetative or body organization.
coenobial. See coenoblum. diaminopimelie acid. Amino acid with two
coenobium (Gk. koinos, common; bios, amino groups; has adefinite chemiea}
Iife). Colony consisting of adefinite structure.
number of ceIls arranged in a specific dichotomy (Gk. die/,a, two; lemnein, to cut).
manner. Repeatedly bifurcating pattern of bran-
coenocytic (Gk. koinos, common; kYlos, ching.
hollow). Multinucleate cell._ dictyosomes (Gk. dictyon, net; soma, body).
colligate (L. colligare, to bind together). Vesicular structures of unknown func-
Septum showing H-shaped structure. tions found in the cytopIasm of eucaryo-
colonial (L. c%nia, form). Habit showing tic ceIls. Also known as Golgi bodies.
number of ceIls held together within an diffused eentromeres. Chromosome con-
envelope. taining many scattered centromeres
conceptac1es (L. concipere, to conceive). along its length.
Cavity-Iike depressions on a receptaele diffuse (L. diffundere, to pour). Pertaining
of the Fucales that contain gametangia. to growth in which every vegetative cell
conJugatlon (L. cum, together; jungare, to is capable of growth and division.
yoke). Sexual union in Conjugales dimorphie (Gk. dis, twice; morphe, shape).
involving fusion between amoeboid Where gametophytic and sporophytic
gametes. generations are morphologically distinct.
contractile vacuoles. Organelles of osmo- dioecious (Gk. dis, twice; oikos, house).
regulation; also thought to play a role Location of male and fernale sex organs
in the excretion of waste material. on separate planls.
costae (L. costa, rib). Thickened regions diplohaplont (Gk. dip/oos, double; haploos,
 208 GLOSSARY

simple; on, being). Where diploid and gas vaeuoles. Gas-filled cavities in cells of
haploid generations alternate succes- certain planktonic blue-green algae
sively. which disappear when subjected to
diplont (Gk. dip/oos, double; on, being). pressure. Also known as pseudovaeuoles.
Diploid plant in wh ich the only haploid girdle view. Side view of a diatom that
stage consists of gametes. reveals the junction of epitheca and
eneapsulated (Gk. en, in; L. capsu/a, little hypotheca.
box). Pertaining to habit; a sac-like globule (L. g/obu/us, small globe). Male
covering enclosing an organism. reproductive organs of Charales having
endophyte (Gk. endon, within; phyton, a jacket of sterile cells around the fertile
plant). A plant living within another celIs; analogous to antheridium.
plant. glomerule (L. glomus, ball). Cluster of
endoplasmie retieulum. Fine tubular or branches at the node, as in Batrachosper-
vesicular structures traversing the cyto- mum.
plasm of eucaryotic cells. glueosamine. A glucose derivative contain-
endospore (Gk. endon, within; sporos, seed). ing amino group of the second carbon
Internally formed thin-walled spores of of the 6-carbon molecule.
Cyanophyta, analogous to aplanospores. glyeolipids (Gk. g/ykys, sweet; Iipos, fat).
epitheea (Gk. epi, upon; theke, box). The Compounds made of lipids and galactose.
larger half of the diatom frustule wh ich gonimoblasts (Gk. gonimos, productive;
covers the smaller half (hypotheea). b/astos, bud). Filaments formed from
euearyota (Gk. eu, weil; karyon, nuc1eus). the zygote in red algae; bear carpospo-
Cellular organisms, in which genetic, rangia.
respiratory and photosynthetic appara- gonospores (Gk. gonos, offspring; sporos,
tuses are organized into nucleus, seed). Spores produced following meio-
mitochondrion and chromatophore, res- sis; also called meiospores.
pectively. gynandrosporous (Gk. gyne, woman; aner,
eutrophie (Gk. eu, weil; trophe, nourish- man; sporos, seed). Species bearing both
ment). Pertaining to habitats rich in oogonia and androsporangia on the same
nutrients and organic matter for the filament.
growth of algae and other plants and haematoehrome (Gk. haima, blood; chroma,
microorganisms. colour). Orange or red oily pigment
exospores (Gk. exos, outward; sporos, seed). related to carotene and present in resting
Spores produced externally or outwardly cells and eye spots of certain algae.
as in Cyanophyta; analogous to aplano- haplont (Gk. hap/oos, simple; on, being).
spores. Haploid plant in wh ich the only diploid
eye spot. Red-coloured spot (stigma) be- stage is confined to the zygote.
Iieved generally to have a visual function. heteroeyst (Gk. heteros, other; kystis,
false branehing. Branching resulting from bladder). Specialized enigmatic cell
the degeneration of a cell in a loop or found in certain blue-green algae.
from growth of free ends of trichome heterokaryosis (Gk. heteros, other; karyon,
through filament sheath, as in some nucleus). Association of nuclei of differ-
blue-green algae. ent genetical constitutions in a vegetative
filamentous (L. /i/um, thread). Thread-like cell.
photosynthetic plants. heteromorphie (Gk. heteros, other; morphe,
flagella (L. flagellum, whip). Fine, thread- shape). Life cycle involving alternation
Iike structures by the activity of which between morphologically dissimilar gene-
the cells move. rations.
fueosan vesicles (L.fucus, seaweed; vesicula. heterothallie (Gk. heteros, other; thallos,
sm all bladder). Sac-like structures in the young). Self-incompatibility; sexual
cells of brown algae; contain phenolics fusion occurs only between gametes of
and tannins. different parentage or plants.
gamete (Gk. gameles, spouse). A sex cell; heterotriehous (Gk. heteros, other; thrix,
two gametcs of opposite sex unite to hair). Thallus differentiated into a
form a zygote. prostrate and an erect system ofbranch-
 GLOSSARY 209
ing filaments. cally similar to laminar in; sometimes
heterotrophie (Gk. heteros, other; trophe, also called chrysolaminarin.
nourishment). Organisms dependent on loop-formation. Arch or dome formation;
exogenous organie sourees for their achieved by rejuvenation of growth and
metabolism and growth. cell division in certain cells of the tri-
hologamy (Gk. holos, whole; gametes, chome of Scytonemataceae.
spouse). Fusion of mature individuals, maerandrous (Gk. makros, large; aner,
i.e., mature individuals directly act as male). Filaments producing antheridia
gametes. similar in size and morphology to those
holophytie (Gk. holos, whole; phyton, producing oogonia; sexually monomor-
plant). Plant-like mode of nutrition phic plants, as in Oedogonium.
involving photosynthesis. meiospores (Gk. meion; less; sporos, seed).
holozoie (Gk. holos, whole; zoon, animai). Spores formed after meiosis in the
Feeding like animals by ingesting solid zygote.
food. mitochondria (Gk. mitos, thread; chondros,
homothalJie (Gk. homos, same; thaI/os, grain). Cytoplasmic double-membraned
young shoot). Self-compatible; fusion organelles coneerned with energy release
ean oecur between gametes derived from in respiration.
the same plant. mitospores (Gk. mUos, thread; sporos,
hormocyst (Gk. hormos, chain; kystis, seed). Spores formed after mitosis; may
bladder). Thick-walled hormogonium be haploid or diploid.
or multicellular akinete found in a few monoeeious (Gk. monos, single; oikos,
blue-green algae. house). Plant bearing both male and
hormogone (Gk. hormos, ehain; gone, gen- female sex organs on the same
eration). Short piece of trichome con- individual.
sisting of undifferentiated vegetative cells monomorphic (Gk. monos, single; morphe,
wh ich are moniliform; hormogonium is form). Formation of only one kind of
generally motile and is meant for plant in the Iife cyde.
propagation. mucopeptides. Compounds made of carbo-
hypnospore (Gk. hypnos, sleep; sporos, hydrates and amino acids, carbohydrates
seed). Thick-walled spore; meant for are N-acetylglucosamine and N-acetyl
perennation. muramic acid; amino acids are glutamie
hypotheca (Gk. hypo, under; theke, box). acid, alanine, glycine, aspartic acid,
Inner and sm aller valve of a diatom lysine, or diaminopimelic acid.
frustule. multiaxial (L. multi, many; axis, axis). For-
idioandrosporous (Gk. idios, distinct; aner, mation of main axis of the thallus by a
male; sporos, seed). Species bearing group of branched filaments.
androsporangia and oogonia on separate multipIlcative (L. multip/icare, to make
filaments. manifold). Process leading to inerease
intercalary (L. intercalaris, inserted). in number of an organism or its cells.
Growth pattern in which newly formed muramie acid. It is a glucosamine derivative
cells are produced between two existing containing carboxyethyl group at 3-0
cells, e.g., of a filament. position, e.g., 3-0-carboxyethyl-D-glu-
isogamy (Gk. isos, equal; gamos, marriage). cosamine.
Fusion between morphologically and nannandrous (Gk. 1I(Jnos, dwarf; aner, male).
physiologically similar gametes. Sexually dimorphie plants as in Oedogo-
isomorphie (Gk. isos, equal; morphe, form). niaceae with a dwarf male.
Life eyde involving alternation between nudeolar organizing cbromosome. Chromo-
two morphologically similar generations. some concerned with formation of
fi'
laminarin. Polysaccharide food reserve in nucleolus.
~ brown algae. nitrogenase. Enzyme coneerned with eon-
fJleueosin (Gk. leukos, white). Highly re- version of molecular nitrogen to
~ fractile polysaccharide with ß-l,3 ammonia.
~ Jinkages which forms the food reserve in nueule (L. nucula, small nut). Fernale repro-
~ Chrysophyta and Xanthophyta; chemi- ductive organ of Charales.
210 GLOSSARY

oligotrophie (Gk. o/igos, few; trophe. nou- polyphosphate granules. Polymers of in-
rishment). Habitats relatively poor in organic phosphate stored in algal cells;
nu trients. occur in granular form.
oogamy (Gk. oon, egg; gamos, marriage). polyploidy (Gk. polys, many; aploos, one
Fusion of a motile sperm with a large fold; eidos, form). Cells containing
passive non-motile egg. three, four or more times the haploid
ostiole (L. ostioillm, Iittie door). Opening number of chromosomes.
or pore of a conceptaele of the Fucales. procaryota (Gk. pro, before; karyon, nuc-
ovum (L. ovum, egg). Female, non-motile leus). Cellular organisms lacking mem-
game te or egg cell. brane-bound genetic, photosynthetic and
oxidation pond. An enelosure for sewage respiratory organelles.
designed to promote digestion of sewage propagule (L. propagare, to propagate). A
with the help of oxygen released by algae few-celled branch serving as a means of
during photosynthesis. vegetative propagation, as in Sphacela-
palmella stage (Gk. palmos, Quivering). riales.
Temporarily non-motile sedentary stage protandrous (Gk. protos, first; aner, male).
in the Iife history of certain motile algae; Hermaphrodite organisms in which thc
cells remain passive and embedded in male reproductive structure matures
gelatinous matrix. earlier than female.
palmelloid (Gk. palmos, Quivering; eidos, pseudoparl'nchymatous (Gk. pseudo, false;
form). Palmella-Hke habit. para, beside; engchyma, infusion). Col-
pantonematic (Gk.pantos, all; nema, thread). lection of cells, filaments or hyphae form-
Flagellum in which the surface is covered ing a tissue which resembles parenchyma.
with hair-Hke appendages. pseudoraphe (Gk. pseudo, false; raphe,
parasexual (Gk. para, compared with; seam). False raphe; longitudinal spacl'
sexus, sex). Organisms showing genetic on the valve of a dia tom bounded on
recombination not involving regular alter- both sides by striae.
nation between karyogamy and meiosis. punctae (L. punctum, point). Perforations
parenchymatous (Gk. para, beside; eng- in the wall of diatoms.
chyma, infusion). Pertaining to a tissue raphe (Gk. raphe, seam). Longitudinal
composed of thin-walled and Iiving cells. eleft seen on the valve surface of a
perlplast (Gk. peri, around; plastos, moul- diatom.
ded). Delicate protective covering of receptacle (Gk. recipere, to receive). Tip
fiagellates that lack cell wall. of branch of thallus bearing conceptaeles
phototaxis (Gk. photos, light; taxis, as in the Fucales.
arrangement). Movement oriented in recomblnants. Offspring with new com-
response to light. bination of genes different from either
phycobilisomes (Gk. phykos, seaweed; L. parent.
bilis, bile; Gk. soma, body). Partieles repllcate (L. replicare, to fold back). A
containing phycobilin, phycocyanin and septum showing two circular infoldings,
phycoerythrin pigments. one on either side.
plakea stage (Gk. plakoeis, fiat cake). The repression. Inhibition of enzyme synthesis.
curved plate-Iike, 8-celled stage in the reproductlve (L. re, again; producere, to
development of a coenobium. lead forth). Processes leading to the
plane (L. planus, plain). Pertaining to continuation of species or races.
septa with smooth fiat surface. rhizopodia (Gk. rhiza, root; podos, foot).
plasmodesmata (Gk. plasmor, form; desma, Unicellular organisms capable of form-
bond). Cytoplasmic strands connecting ing pseudopodia or false feet for locomo-
adjoining cells and Hnking cytoplasm. tion or anchorage.
polar nodules (Gk. polos, pole; L. noduills, rhlzopodial (Gk. rhiza, root; podos, foot).
small knob). Wall thickenings at the Type of habit in which unicellular organ-
two poles in a diatom frustule. isms form pseudopodia as locomotory
polyhedral bodies. Polygonal particles organs.
found in cells of blue-green algae; their satellite chromosomes (L. satel/es, atten-
function and chemistry are not known. dant; Gk. chroma, colour; soma, body).
 GLOSSARY 211

Short segmentsof chromosomesconstric- transeription. Process in which DNA is

ted from the rest of the chromosomes. copied into messenger RNA.
semireplicate (L. semi, half; rep/icare, to transduction (L. transducere, to transfer).
fold back). Only one side of septum wall Process of genetic recombination in
showing circular infolding. which DNA of a donor bacterium is
siliealemma (L. silix, flint; Gk. lemma, carried to the recipient by a virus.
husk). TripIe membrane system con- transformation (L. trans/ormare, to change
cerned with deposition of silica walls in in shape). Process of genetic recombina-
dlatoms. tion in bacteria affected by the incor-
simuItaneous division. NucIear and cyto- poration of naked DNA from donor
plasmic divisions occurring concurrently. bacterium.
siphoneous (Gk. siphon, tube). Tubular triehothallie (Gk. thrix, hair; thallos, young
thallus in algae lacking septa or cross shoot). Intercalary growth in which
walls during vegetative phase of growth. meristem is located at the base of ahair.
spermatium (Gk. sperma, seed). Non-flag- trumpet hyphae. Long tubular achlorophyl-
ellated naked male gamete of red algae. lous cells of medulla found in brown
spermoearp (Gk. sperma, seed; karpos, algae; conduct water and nu trients.
fruit). Fruiting body consisting cf sterile unduliseptate(L. undulatus, risen like waves;
jacket of cells around a fertilized oogo- septum, partition). Septum with wavy
nium, as in Coleochaete. margins.
stigma (Gk. stigma, mark). Eye spot or uniaxial (L. unus, one; axis, axis). Plant
a red spot of the flagellates found within axis made of a single filament or its
the cell near or inside the chromato- branches.
phore. It is beIieved to have a regulatory vaeuolar apparatus (L. vacuus, empty; ad,
funchon in cell motility. to; paratus, prepared). Includes two
stipe (L. stipes, stalk). Stalk; portion of kinds of organelles, the contractile vacu-
kelp between holdfast and blade. oles and the cytopharyngeal apparatus
subaerial (L. sub, under; aer, air). Refer- consisting of gullet, reservoir, and a
ring to habitats that are weil raised above number of small contractile vacuoles
soil surface, e.g., tree barks, rocks, and surrounding the reservoir.
stones. vaeuole (L. vacuus, empty). SmalI, usually
sueeessive division. Nuclear and cyto- spherical space within acelI, bounded by
plasmic divisions occurring one after the a membrane and containing some fluid
other. sap.
synaptonemal complex. Characteristic nlve (L. valva, fold). One of the two parts
structures corresponding to bivalents in of a dia tom theca; the other is the con-
meiotic nuclei and seen under electron necting band.
microscope. valve view. Top surface of epitheca or
synzoospore (Gk. syn, with; zoon, animal; hypotheca of a diatom.
sporos, seed). Compound, multiflagellate vegetative (L. vegetare, to enliven). For-
and multinucIeate asexual spore found mation of plant body lacking reproduc-
in Vaucheria. tive structures or organs.
tetraspore (Gk. tetras, four; sporos, seed). water bloom. Luxuriant growth of one or
One of the four non-motHe haploid a few species of algae in water, often
spores formedin tetrasporangium in red imparting colour to water.
algae. zoogamete (Gk. zoon, animal; gametes,
thylakoid. Structural unit of lamellar spouse). Motile llageUate garnete.
system forming a double membrane disc; zoospore (Gk. zoon, animal; sporos, seed).
a photosynthetic lamella. Motile flagellated asexual cell.
 Index

Acelabu/aria, 35, 70 BAZIN, M. J., 60

Acetylene reduction, 59 Biddulphia, 137
Achnanthes, 135 Blue-green algae, 3, 51-67
Acyllipid, 55 differences from bacteria of, 3
AINSWORTH, G. C., 4 differences from red algae of, 167
AKINTOBI, S., 92 resemblances with red algae of, 166
AKINTOBI, T., 160 BOALCH, G. T., 146, 194
A/aria, 38, 141 Bodanella, 141
ALEXOPOULOS, C. J., 4 BOLD, H. C., 75, 82, 83, 121, 130
Algicide, 47, 48 BONNER, J. T., 84
Alginic acid, 41, 142 Boslrychia, 159
Anabaena,23,29,55,56 Botrydium, 25, 36, 124-125, 130
as endophytes, 52 resemblances with Prolosiphon of, 124
genetic recombination in, 60 resemblances with Vaucheria of, 130
in water supplies, 46 BOUCK, G. B., 154, 155
nitrogenase in, 56 BOWEN, C. C., 54
nitrogen fixation in, 58 BRODY, M., 163
role of heterocyst in sporulation of, 55 BROWN, R. M. (Jr.), 82, 83
toxicity of, 46 Bu/bochaele, 103
Anabaenopsis, 44 BURRIS, R. H., 58
Anacyslis, 29 BUSH, V. N., 67
genetic recombination in, 29, 60
genetic transformation in, 60
growth estimation of, 199-200 Calcareous algae, 161
in water supplies, 47 Camply/onema, 178
Androsporangia, 107 CANNON, R. E., 61
Androspores, 107 Carpogonium, 28, 102, 164, 165
Aneuploidy,72 Carposporophyte, 32, 164, 165, 169-171,
Anisogamy, 29, 73, 137, 144 174-176
Antherozoids, 28, 73 CARR, N. G., 59
c1ustering of, 149 CASTENHOLZ, R. W., 58
of Fucales, 156 Carleria, 73
Aphanizomenon, 46 Centrosome, 70
Aphanochaete, 73, 98 Cell wall, 8, 12, 21, 70, 77, 123, 131, 132,
Aphanolhece, 22, 26, 52, 178 133, 141, 146, ISS, 162
Apical growth, 24, 115, 142, 152, 162, 169, Cepha/euros, 46, 67, 69
172 Chaetonema, 73, 98
Aplanogametes, 28 Chaetophora, 47
Apoglossum, 164 Chamaesiphon, 28
Ascophyl/um, 42, 171 Clzantransia, 170, 171
ASlerionella, 46, 132 CHAPMAN, D. J., 32, 77
Au/osira, 36, 44, 58 CHAPMAN, V. J., 32
Auxiliary cell, 163, 164, 165 Chara, 14,32,70, 114-120
Auxospores, 29, 131, 137, 138, 139 Characium, 69
Chlamydomonas, 13, 15, 18, 20, 21, 29,
Bangia, 164 31-32,47,69,71-73,76-84,196
Balrachospermum, 24, 32, 34, 161, 162, Chlorella, 21, 28, 38, 69,87-89
168-171 chlorellin production by, 44
 INDEX 213

growth estimation of, 199 DAWSON, R. C., 40

in lichens, 69 De/esserio, 162, 184
in sewage disposal, 45 Derepression, 59
in water supplies, 47 Dermocorpa, 28
mass culture of, 40 DESIKACHARY, T. V., 120
pigment extraction from, 192 Desmareslio, 141
phycovirus infection in, 47 Desmids, 3, 26, 108
synchronous culture of, 88, 89 Diaminopimelic acid, 12
Chlorellin, 44 Diatomite, 45
ChlorochYlrium, 32, 69 Diatoms, 3, 26, 131
Chlorogloea, 52, 59 DIBOLL. A. G., 191
Ch/orogonium, 73 Diclyosphaerium, 26
Chondrus, 38, 42 Diclyola, 27, 141, 142
Chromatic adaptation, 57, 162 Diffused centromeres, 16
Chromatophores, 10, 13-15,20,63,69,70 Diffuse growth 24, 162
Chromocentres, 143 Dinobryon, 46
Chromosomes, 16,71 DONIKE, M., 160
acentromeric, 7 Draparna/dia, 71
centromeric, 6, 71 Droparnoldiopsis, 24, 32, 70, 71, 75, 114,
nucIeolar organizing, 16, 109, 110 119
organized, 16 DREW, K. M. (DREW-BAKER, K. M.),
polycentric, 109 32,40
Xand Y, 143 Dumonlia, 162
Chrysidiaslrum, 22
Chrysodendron, 22
Chrysolaminarin, 123, 131 Ecba//ocyslis, 22
Cladophora, 24, 27,33,44,46,69-71,94-98, Ecklonia, 41
131 EClocarpus, 24, 27, 28, 32, 33, 141, 146-50,
Coccolithophorids, 17 194
Cocconeis, 131 Edible algae, 38-41
COLE, K. M., 92 EDWARDS, P., 176
Co/eochaele, 24, 73, 74, 101-103 Eisenio, 41
Colonial habit, 21, 22 Encapsulated habit, 21
COLLYER, D. M., 166 Endoplasmic reticulum, 20, 79, 143, 163
Co/pomenia, 141 ERBEN, K., 126, 127
Conceptacles, 144, 151-155, 158-160 Eremosphaero, 71
Conchoce/is, 4O, 164, 170 ETTL,H.,78
Cora//ina. 161 Eucaryota, 3, 11
CRAIGIE, J. S., 20 Eug/eno, 18, 21
CRONQUIST, A., 4, 63, 77, 167 in sewage disposal, 45
Cryptoblasts (cryptostomata), 151, 153, 154 in water supplies, 47
Culture collections of algae, 192 Eunolia, 131
CuticIe, chemistry of, 20 Eustigmatophyceae, 124
CUlleria, 141 EVANS, L. V., 142, 146, 187
Cyanophage,47,6O,61, 62,167 EVANS, M. C. W., 56
Cyc/ole/la, 131, 134, 138, 139 Evection, 97
Cy/indrocopso, 92 Extracellular products, 44
Cylindrocystis, 108 as carbon and nitrogen sourees, 44
Cy/indrospermum, 29, 36, 58, 60, 178 as chelators, 44
Cystocarp, 170, 174 of nitrogen-fixing algae, 44, 59
Cysloseira, 41 Eye spot (stigma), 9, 18, 19, 78, 79
Cytokinesis, 72
Cytopharyngeal apparatus, 9
FAY,P.,55
DAWES, C. J., 109, 110 Flagella, 12, 13,70,79, 125, 127, 128, 141
214 INDEX

Floatation of diatoms, 136 HIBBERD, D. J., 124

Floridean stareh, 161, 163 HILL, J. C., 105
FOGG, G. E., 48, 53, 166 Himanthalia, 141
Fossil algae, 114, 139 HOARE, D. S., 59
Fragilaria, 135 HOFFMAN, L. R., 104
FRIEDMANN, 1., 190 Hologamy,29
FRITSCH, F. E., 8, 62, 75, 76, 98, 144, 145 Homothallic, 29, 73, 80
Frifschiella, 72, 98, 99 HOSHAW, R. W., 197
Fucinic acid, 142 Hot water springs, 52
Fucosan vesicles, 141 blue-green algae of, 52
Fucus, 33,41,42,43, 142, 151-57, 158, 160 HUBER, J., 142
Fungi, 3,4 Hya/odiscus, 131
Hydrodicfyon, 22, 71, 72,87,89,91
Hypnospores, 27
Gas vacuoles (pseudovacuoles), 53
Gelidium, 42, 162
Genetic recombination, 26 Iyenzaria, 141
distinction from true seltuality of, 60
in bacteria, 26
in blue-green algae, 26, 27,60 JAENICKE, L., 160
Genetic transformation, 26 JOHNSON, O. P. S. C., 82, 83
in blue-green algae, 60
Geosiphon pyri/orme, 65
Gigartina, 42 KHYEN, N. T., 60
Globule, 116 KLEIN, R. M., 4, 63, 77, 167
Gloeocapsa, 56, 63 KULASOORIY A, S. A., 55
G/oeofrichia, 36, 55 KUMAR, H. D., 60
G/oiosiphonia, 162
Glomerule, 169
Glucosamine, 12 Laminaria, 42, 141, 142
Glycolipid, 55 Laminarin, 141, 145
GODWARD, M. B. E., 187 LANG, N. J., 55, 56
Golgi apparatus (dictyosomes), 17,79, 143, LAZAROFF, N., 60, 66
163 Leathesia, 141
Gomphonema, 131 LEEDALE, G. F., 16, 124
Gonimoblasts, 32, 163-65 Lemanea, 161
Goniolrichum, 162 Lessonia, 41
Graci/aria, 42, 184 Leueosin, 123, 145
Granules, 20,54 LEWIN, J. C., 134
Grijfifhsia, 163 Licmophora, 132
GROSS, F., 136 Lifhodesmium, 135
Gymnodinium, 45 Lifhophyllum, 43
Lithothamnion, 43, 161
HAINE, L. A., 20 LUND, J. W. G., 136
HALFEN, L. N., 58 Lyngbya,44,52
Haptophyceae, 9, 17
HASE, E.,89
HAUPT, W., 126 MACKERETH, F. J. H., 180
Helminthoc/adia, 162 Macrocystis, 41
HENDEY, N. I., 134 MANTON, I., 142
Heribaudiella, 141 Mastigoc/adus, 51, 58
Heterocysts, 51, 55-57 McCuJly, M. E., 191
Heterothallic, 29,73, 81, J08 Melosira, 131, 138
Heterotrichous habit, 98, 103, 142, 145,164, Merismopedia, 52
170, 172 Mesotaenium, 14,71, 108
 INDEX 215

MICHALlS, L., 105 PARKER, B. C., 142, 191

Microcyslis, 44, 45, 46, 51, 180 PARSONS, T. R., 180
Mieroquadrats, 178 Parthenogenesis, 156
Microspora, 17,92 Parthenospores, 74
Miecotransects, 178 PEAT, A.,64
Mitochondrion, 19-20 Pe[~ophycus, 142
MOESTRUP, 0., 116 Pelvetia, 159
MOLLENHAUER, R., 66 Perennation, 34, 35, 74
MORIMURA, Y., 88 Periplastie envelope, 9
MORRIS, M. E., 60 Phormidium, 52
Mougeolia, 7 J, 108 Phototaxis, 79
Movements, 57, 58, 136 Phycobilisomes, 14, 163
MÜLLER, D. G., 149, 150 Phytochrome, 14,57,109
Muramie acid, 3 Pigment extraetion, 191-92
Pinnu[aria, 131, 134, 135
Pähophora, 36,46,74
NAGATA, Y., 109 Planktonie algae, 42, 47
Navicula, 131, 134, 135 Pleurocapsa, 28
Nemolion, 24 Pleurocladia, 141
Nereocyslis, 142 Pleurococcus, 74
Neuromotor apparatus, 70 Plocamium, 162, 184
Nitrogenase, 55, 56, 59 Polarity, IS7
Nitrogen-fixing bIue-green algae, 42, 44, 59 Polyhedral bodies, 20
Nilzschia, 131, 136 Polyphosphate granules, S6
NOSloc, 24, 38, 52, 58, 59, 60, 65, 171 Polysiphonia, 28, 32, 34, 164, 171-176
Nuclei, 6, 7, 16, 135, 163 Porphyra, 24, 38-40,162,164,170,184
Nucleolar organizing ehromosome, 16, 109 Porphyridium, 161, 162, 163
Nueule, 116, 117 Pras;ola, 25, 74, 92
Procarp, 164, 173
Procaryota, 3, 11, 51, 62
Oedoclodium, 103 Propagules, 143
Oedogonium, 14, 15, 17, 33, 70, 72, 73, 74, Protonema, 144, 170
103-108, 131, 198 Protopyrenoids, 109
Oligotrophie lakes, 168 Prolosiphon, 25, 27, 36, 74
Ooblast, 164 Prymnesium, 45
Oogamy, 29, 73, 144 Pseudoparenehymatous habit, 142,145, 162
Oscillaloria, 47, 57, 58, 64, 65 Pseudoraphe, 135
Oxidation ponds, 40 Pylaiella, 142, 146
Pyrenoids, 15-16, 71, 136, 142, 146, 162
Pyrocystis, 14
Palmella stage, 80
Pandorina, 74
PANKRATZ, H. S., 54 Raphes, 131, 136
PAPENFUSS, G. F., 120 RAVEN, P. H., 10, 15, 20
Parallelism, in evolution, 4 Receptacle, 144, 158
between Chloro-, Phaeo-, and Rhodo- REIMANN, B. E. F., 134
phyta, 145 Repression, 59
between sex organs of Characeae and Rhizopodia, 21, 22
bryophytes, 120 RhodochYlrium, 69
between Xantho- and Chlorophyeeae, Rhodymenia, 38
123 Ribosomes, 20
Parasexuality, 28 ROUND, F. E., 9, 75, 124
Parasitic algae, 69, 161
Parenehymatous habit, 24, 76, 91, 142,
145,162 SAFFERMAN, R. S., 47, 60
216 INDEX

SAG ER, R., 83 SWIFT, E., 191

Sargassum, 141, 157-60 SynaptonemaI complex, 143
Scenedesmus, 45 Synechococcus,41
SCHEIBE, J., 57 Synechocystis, 21
SCHULTZ, M. E., 139 Synedra, 46, 131, 135
SCHWABE, G. H., 66 Synura, 46
Scytonema, 52, 66--67, 178
Seaweed manure, 44
Separation dises, 27 T AMIY A, H., 40, 88
Septa, 17 Tetraspora, 22
Sex hormone, 108, 149 Tetrasporophyte, 163, 165, 173
Sexuality, 28 THIVY, F., 43
advance of, from isogamy to oogamy, TIWARI, D. N., 55
145 To/ypothrix,44, 52, 57, 58, 59, 178
laboratory demonStration of, 196-99 TRAINOR, F. R., 82, 139
origin of, 29-31 Transcription, 59
SHANE, M. S., 67 Transduction, 26
SHEST AKOV, S. V., 60 Trebouxia, 69, 71, 74
Silicalemma, 134 Trenrepohlia, 27, 69, 72
SILVEY, J. K. G., 56 Tribonema, 123
SINGH, R. N.,44, 55, 60 Triceratium, 131
SINHA, R.,60 Trichodesmium, 57
Siphoneous habit, 24 Trichothallic growth, 24, 142
Sirodotia, ~7 Trumpet hyphae, 142
Sirogonium, 13
SMITH, G. M., 4
SMITH, R. V., 56 U/othrix, 13, 17, 23, 24, 27, 28, 29, 30, 72,
SoH reclamation, 44 73,74,92-94
Sperrnatium, 28, 164 Ulva, 23, 24, 34, 38,76, 184
Spermo~arp, 102 Urospora, 75
Sphacelaria, 141
Sphaerella, 36, 71, 74, 85
Sphaerop/ea, 17,70,92 Vacuoles, 9, 19,53,71,77, 163
Spirogyra, 14, 16,29,70,71,73,109-112 VAN NIEL, C. B., 9
importance in water supplies, 46, 47 VATTER, A. E., 163
Spirolaenia, 108 Vaucheria, 14,25,27, 123, 125-30
Spirulina, 21, 44 Vegetative propagation, 26, 27, 143
Sporangia, 143,'144, 147, 148, 150 VISHNIAC, W., 61, 66
Spores, 27, 28, 29, 125, 128, 131, 137, 143, Volvox, 22, 84-87
163
STANIER, R. Y., 9
STARR, R. C., 87 Water blooms, 48, 64, 180
STEWART, W. D. P., 42 Westiella, 35, 36
Stichococcus,27 WHITTON, B. A., 64, 95, 97
Stigeoc/onium, 23, 24, 32, 73 WIESE,L.,82
Stipe, 24, 142, 152, 157 WOLK, P., 55
Streb/onema, 146 WYATT, J. T., 56
STRICKLAND, J. D. H., 180
Subaerial algae, 35, 69, 98
SUNDARALINGAM, V. S., 120 ZEUTHEN, E., 136
Surirella, 14 Zygnema, 14, 16,29,71,73, 108, 112-13
SUSSMAN, A. S., 4 Zygote, 30, 73, 74