Table Of Contentinfrastructure
of
Microalgae
Edited by
Tamar Berner, Ph.D.
Department of Life Sciences
Bar-Ilan University
Ramat-Gan, Israel
CRC Press
Boca Raton Ann Arbor London Tokyo
Library of Congress Cataloging-in-Publication Data
Ultrastructure of microalgae/edited by Tamar Bemer.
p. cm.
Includes bibliographical references and index.
ISBN 0-8493-6323-3
1. Microalgae—Ultrastructure. I. Berner, Tamar.
QK568.M52U38 1993
589.3'872—dc20 92-21125
CIP
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PREFACE
Algae represent a very diverse group of organisms. Unlike the Chromo-
phyta (Bryophyta, Pteridopyta, and Spermatophyta), whose classification is
based on macromorphology and life cycles, algal systematics is determined
by different characteristics. Biochemical content of photosynthetic pigments,
storage products, and chemistry of the cell walls are basic features of algal
taxonomy. Since the 1950s, when electron microscopy was shown to be a
powerful tool in phycological research, it became obvious that the ultrastruc-
ture of algae was at least as valuable as biochemistry in determining the
taxonomic affinities of algal groups.
It has been a long time since Dodge's comprehensive monograph The
Fine Structure of Algal Cells (1973) was published. In view of the many
dramatic developments in microscopic techniques and in our understanding
of algal evolution and taxonomy, which have occurred since, an effort to
produce an updated, comprehensive treatise is both justified and timely.
The emphasis of this book is on the fine structure of microalgae. Although
most of the features discussed are typical of macroalgae as well, it seemed
that it would be easier to deal with the subject on a cellular level, thereby
excluding all features relating to tissue organization, as well as multicellular
structures.
Cyanophyta and Prochlorophyta are included in this book in spite of being
prokaryotes, unlike the "true'' eukaryotic algae. The reason for their inclusion
is that these two groups are studied primarily by phycologists. Glaucophyta
were excluded (except for brief mentions in Chapters 7 and 8) since they are
more like protozoa with photosynthetic cyanobacterial endosymbionts than
algae with fully integrated chloroplasts. Some small organelles are not dis
cussed since it was felt that not much study has been done since the publication
of Dodge's book.
While accepting all responsibility for the final product, I gratefully ac
knowledge the cooperation of those phycologists who generously contributed
their knowledge and time in writing chapters for this volume. I would es
pecially like to thank Zvy Dubinsky, from Bar-Ilan University, Israel, for
suggesting this volume, and Sheldon Aaronson, from Queens College, New
York, for his continuous advice and support offered throughout the project.
Tamar Berner
THE EDITOR
Tamar Berner, Ph.D., is a Lecturer of botany in the Department of Life
Sciences at Bar-Ilan University and at the School of Education in Oranim,
Haifa University, Israel. Dr. Berner graduated from the Hebrew University
in Jerusalem, where she received her Ph.D. The subject and title of her thesis,
The Ecophysiology of Algae Underneath Flint Pebbles in the Negev Desert
of Israel, was completed under the supervision of the late Professor M.
Evenari. For several years Dr. Berner studied the physiology of microalgae
and their role in high-rate oxidation ponds and the optimization of sewage
treatment through such systems. In recent years she has been working on the
ultrastructure of microalgae.
The emphasis of Dr. Berner's research is on the effect of environmental
factors on changes in the ultrastructure, especially of light and the light-shade
acclimation response, as well as different nitrogen regimes on photosynthate
accumulation as shown by electron microscopy. The main objects of study
are zooxanthellae of marine invertebrates in the coral reefs of the Red Sea
and Hawaii, and zoochlorellae in sponges of Lake Baikal.
CONTRIBUTORS
Sheldon Aaronson, Ph.D. Thomas E. Jensen, Ph.D.
Professor Professor and Chairman
Department of Biology Department of Biological Sciences
Queens College Lehman College
City University of New York City University of New York
Flushing, New York Bronx, New York
Barry S. C. Leadbeater, Ph.D.
Tamar Berner, Ph.D. School of Biological Sciences
Department of Life Sciences The University of Birmingham
Bar-Ilan University Birmingham, United Kingdom
Ramat-Gan, Israel
Charles J. O'Kelly, Ph.D.
Senior Lecturer
Guy Cox, Ph.D.
Department of Plant Biology
Electron Microscope Unit
Massey University
University of Sydney
Palmerston North, New Zealand
Sydney, Australia
Hans J. Sluiman, Ph.D.
J. C. Green, Ph.D.
Head
Plymouth Marine Laboratory
Scientific and Technical Services
Citadel Hill
Royal Botanic Garden
Plymouth, United Kingdom
Edinburgh, Scotland
United Kingdom
Isao Inouye, Ph.D.
Associate Professor Jean M. Whatley, Ph.D.
Institute of Biological Sciences Department of Plant Sciences
University of Tsukuba University of Oxford
Tsukuba, Japan Oxford, United Kingdom
TABLE OF CONTENTS
Introductory Table
Ultrastructural Characterization of Eukaryotic Algal Groups 2
Tamar Berner
Chapter 1
Cyanobacterial Ultrastructure 7
Thomas E. Jensen
Chapter 2
Prochlorophyceae 53
Guy Cox
Chapter 3
Cell Coverings of Microalgae 71
Barry S. C. Leadbeater and John Green
Chapter 4
Flagella and Flagellar Apparatuses of Algae 99
Isao Inouye
Chapter 5
Chloroplast Ultrastructure 135
Jean M. Whatley
Chapter 6
The Ultrastructure and Function of the Contractile Vacuole 205
Sheldon Aaronson
Chapter 7
Nucleus, Nuclear Division, and Cell Division 221
Hans J. Sluiman
Chapter 8
Relationships of Eukaryotic Algal Groups to Other Protists 269
Charles J. O'Kelly
Index 295
Ultrastructure
of
Microalgae
2 Ultrastruclure of Microalgae
Introductory Table*
Tamar Berner
Ultrastructural Characterization of Eukaryotic Algal Groups 1-6
No Membrane One Membrane
Rhodophyta Chlorophyta Euglenophyta
Flagellum Two anterior (whip One anterior pantone-
lash) acronematic matic flagellum
isokont flagella;
sometimes stephan-
okont
Hairs Fibrillar solid hairs Nontubular fibrillar
(Tripartite tubular hairs
hairs in some primi
tive Chlorophyta)
Chloroplast Two layered chloro Two layered chloro CER membrane not
plast envelope plast envelope continuous with nu
clear membrane
Thylakoids/band 1 2-7; sometimes
pseudograna
Girdle lamella
Storage products Outside chloroplast Within the chloro- Outside chloroplast
location plast stroma, often
as a sheath around
the pyrenoid
Chromosomes Condensed at mitosis Condensed at mitosis Persistently
condensed
Kinetochore - or +
Nucleolus Not persistent at Not persistent at Persistent at mitosis
mitosis mitosis (endosome)
Nuclear envelope
Intact
Polar gaps
Breaks down ( + )
Spindle type Closed nuclear and Nuclear and cyto Intranuclear and sub-
cytoplasmic plasmic or intra spindles
nuclear
Polar structure Polar rings; no true Centrioles
centrioles
Cytokinesis Annular wall plus Phycoplast and Cleavage
septal plug phragmoplast
* This table is based on the division of the algal phylla according to the number of membranes
of chloroplast endoplasmic reticulum (CER)
Introductory Table 3
Ultrastructural Characterization of Eukaryotic Algal Groups
(continued)
One Membrane Two Membranes
Dinophyta Cryptophyta Chrysophyta
Flagellum Transverse pantone- Two anterior panto- Two unisokont with
matic flagellum, lo nematic flagella long anterior panto-
cated in a groove or nematic and short
a girdle and shorter posterior acrone-
posterior acrone- matic flagellum
matic flagellum
Hairs Nontubular fibrillar Tripartite tubular Tripartite tubular
hairs hairs hairs
Chloroplast CER membrane not ? Outer CER mem
continuous with nu brane continuous
clear membrane with outer mem
brane of nuclear
envelope
Thylakoids/band 3 2 3
Girdle lamella - - +
Storage products Within the chloro Between chloroplast Outside chloroplast
location plast stroma, often envelope and CER
as a sheath around
the pyrenoid
Chromosomes Persistently con Persistently Condensed at mitosis
densed; attached to condensed
nuclear envelope
Kinetochore - + -
Nucleolus Not persistent at Persistent at mitosis Not persistent at
mitosis mitosis
Nuclear envelope
Intact + +
Polar gaps
Breaks down + +
Spindle type Cytoplasmic tunnels Nuclear and cyto Nuclear and cyto
plasmic plasmic
Polar structure 7 Flagellar bases; no Rhizoplast
true centriole
Cytokinesis Cleavage Cystokinetic ring Cleavage
