algae anatomy biochemistry 0
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AlgaeAnatomy, Biochemistry,
and Biotechnology
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AlgaeAnatomy, Biochemistry,
and Biotechnology
Laura Barsanti
Paolo Gualtieri
A CRC title, part of the Taylor & Francis imprint, a member of the
Taylor & Francis Group, the academic division of T&F Informa plc.
Boca Raton London New York
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Published in 2006 by
CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742
2006 by Taylor & Francis Group, LLC
CRC Press is an imprint of Taylor & Francis Group
No claim to original U.S. Government works
Printed in the United States of America on acid-free paper
10 9 8 7 6 5 4 3 2 1
International Standard Book Number-10: 0-8493-1467-4 (Hardcover)
International Standard Book Number-13: 978-0-8493-1467-4 (Hardcover)
Library of Congress Card Number 2005014492
This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with
permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish
reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials
or for the consequences of their use.
No part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or
other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information
storage or retrieval system, without written permission from the publishers.
For permission to photocopy or use material electronically from this work, please access www.copyright.com
(http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC) 222 Rosewood Drive, Danvers, MA01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For
organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged.
Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for
identification and explanation without intent to infringe.
Library of Congress Cataloging-in-Publication Data
Gualtieri, Paolo, 1952-
Algae : anatomy, biochemistry, and biotechnology / by Laura Barsanti and Paolo Gualtieri.
p. ; cm.
Includes bibliographical references and index.
ISBN-13: 978-0-8493-1467-4 (alk. paper)
ISBN-10: 0-8493-1467-4 (alk. paper)
1. Algae. [DNLM: 1. Algae. 2. Biotechnology. QK 566 G899a 2005] I. Barsanti, L. II. Title.
QK566.G83 2005
579.8--dc22 2005014492
Visit the Taylor & Francis Web site at
http://www.taylorandfrancis.com
and the CRC Press Web site athttp://www.crcpress.com
Taylor & Francis Groupis the Academic Division of Informa plc.
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Preface
This book is an outgrowth of many years of research aimed at studying algae, especially micro-
algae. Working on it, we soon realized how small an area we really knew well and how superficial
our treatment of many topics was going to be. Our approach has been to try to highlight
those things that we have found interesting or illuminating and to concentrate more on those
areas, sacrificing completeness in so doing.
This book was written and designed for undergraduate and postgraduate students with a
general scientific background, following courses on algology and aquatic biology, as well as for
researchers, teachers, and professionals in the fields of phycology and applied phycology. In our
intention, it is destined to serve as a means to encourage outstanding work in the field of phycology,
especially the aspect of teaching, with the major commitment to arouse the curiosity of both
students and teachers. It is all too easy when reviewing an intricate field to give a student new
to the area the feeling that everything is now known about the subject. We would like this book
to have exactly the reverse effect on the reader, stimulating by deliberately leaving many doors
ajar, so as to let new ideas spring to mind by the end of each chapter.
This book covers freshwater, marine, and terrestrial forms, and includes extensive original
drawings and photographic illustrations to provide detailed descriptions of algal apparatus. We
have presented an overview of the classification of the algae followed by reviews of life cycles,
reproductions, and phylogeny to provide conceptual framework for the chapters which follow.
Levels of organization are treated from the subcellular, cellular, and morphological standpoints,together with physiology, biochemistry, culture methods and finally, the role of algae in human
society. Many instances of recent new findings are provided to demonstrate that the world of
algae is incompletely known and prepared investigators should be aware of this.
Each of the chapters can be read on its own as a self-containing essay, used in a course,
or assigned as a supplemental reading for a course. The endeavor has been to provide a
hybrid between a review and a comprehensive descriptive work, to make it possible for the
student to visualize and compare algal structures and at the same time to give enough references
so that the research worker can enter the literature to find out more precise details from the original
sources.
The bibliography is by no means exhaustive; the papers we have quoted are the ones we have
found useful and which are reasonably accessible, both very recent references and older classicreferences that we have judged more representative, but many excellent papers can be missing.
In our opinion, too many references make the text unreadable and our intention was to put in
only enough to lead the reader into the right part of the primary literature in a fairly directed
manner, and we have not tempted to be comprehensive. Our intention was to highlight the more
important facts, hoping that this book will complement the few specialized reviews of fine structure
already published and will perhaps make some of these known to a wider audience. Our efforts
were aimed at orientating the readers in the mare magnum of scientific literature and providing
interesting and useful Web addresses.
We are grateful to the phycologists who have contributed original pictures; they are cited in the
corresponding figure captions. We are also grateful to the staff at CRC Press, Boca Raton, Florida,particularly our editor, John Sulzycki, for his patience and human comprehension in addition to
his unquestionable technical ability, and to the production coordinators, Erika Dery and Kari
A. Budyk.
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Our sincere gratitude and a special thanks to Valter Evangelista for his skillful assistance and
ability in preparing the final form of all the drawings and illustrations, and for his careful attention
in preparing all the technical drawings. We appreciated his efforts to keep pace with us both and to
cope with our ever-changing demands without getting too upset.
We will always be grateful to Vincenzo Passarelli, who frequently smoothed a path strewn with
other laboratory obligations so that we could pursue the endeavors that led up to the book, and
above all because he has always tolerated the ups and downs of our moods with a smile on his
face, and a witty, prompt reply. He lighted up many gloomy days with his cheerful whistling.
We are sure it was not always an easy task.
For the multitudinous illustrations present in the book we are indebted to Maria Antonietta
Barsanti and to Luca Barsanti, the sister and brother of Laura. When the idea of the book first
arose, about four years ago, Maria Antonietta took up the challenge to realize all the drawings
we had in mind for the book. But this was just a minor challenge compared with the struggle
she had been engaged with against cancer since 1996. Despite all the difficulties of coping with
such a disabling situation, she succeeded in preparing most of the drawings, with careful determi-
nation, interpreting even the smallest details to make them clear without wasting scientific accu-racy, and still giving each drawing her unique artistic touch. She worked until the very last days,
when even eating or talking were exhausting tasks, but unfortunately last February she died
without seeing the outcome of her and our efforts. She will always have a very special place in
our hearts and our lives. Her brother Luca Barsanti completed the drawing work in a wonderful
way, making it very hard to distinguish between her artistic skill and his. His lighthearted and
amusing company relieved the last and most nervous days of our work, and also for this we will
always be grateful to him.
In July 2004, Mimmo Gualtieri, the only brother of Paolo, died of an unexpected heart attack.
He left a huge empty room in his brothers heart.
In October 2004, our beloved friend and colleague, Dr. Patricia Lee Walne, distinguished
professor of botany of the University of Tennessee in Knoxville, died after a long and serious
illness.
This book is dedicated to the three of them.
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About the Authors
Dr. Laura Barsanti graduated in natural science from University of Pisa, Italy. At present she is a
scientist at the Biophysics Institute of the National Council of Research (CNR) in Pisa.
Dr. Paolo Gualtieri graduated in biology and computer science from University of Pisa, Italy. At
present he is a senior scientist at the Biophysics Institute of the National Council of Research (CNR)
in Pisa and adjunct professor at the University of Maryland, University College, College Park,
Maryland.
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Table of Contents
Chapter 1
General Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Classification. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Occurrence and Distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Structure of Thallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Unicells and Unicell Colonial Algae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Filamentous Algae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Siphonous Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Parenchymatous and Pseudoparenchymatous Algae. . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Reproduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Vegetative and Asexual Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Binary Fission or Cellular Bisection. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Zoospore, Aplanospore, and Autospore. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Autocolony Formation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Fragmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Resting Stages. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Sexual Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Haplontic or Zygotic Life Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Diplontic or Gametic Life Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Diplohaplontic or Sporic Life Cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Summaries of the Ten Algal Divisions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Cyanophyta and Prochlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Glaucophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Rhodophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Heterokontophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Haptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Cryptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Dinophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24Euglenophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Chlorarachniophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Endosymbiosis and Origin of Eukaryotic Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Chapter 2
Anatomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Cytomorphology and Ultrastructure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Outside the Cell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35Type 1: Simple Cell Membrane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Type 2: Cell Surface with Additional Extracellular Material . . . . . . . . . . . . . . . . . . 36
Mucilages and Sheaths. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
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Scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Frustule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Cell Wall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Lorica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Skeleton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Type 3: Cell Surface with Additional Intracellular Material in Vesicles . . . . . . . . . 48
Type 4: Cell Surface with Additional Extracellular and
Intracellular Material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
First Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Second Level . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Third Level. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Flagella and Associated Structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Flagellar Shape and Surface Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Flagellar Scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Flagellar Hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Flagellar Spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63Internal Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Axoneme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Paraxial (Paraxonemal) Rod. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Other Intraflagellar Accessory Structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Transition Zone. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Basal Bodies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Root System . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Glaucophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Heterokontophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Haptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Cryptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
Dinophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Euglenophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Chlorarachniophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
How Algae Move . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Swimming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Movements Other Than Swimming . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Buoyancy Control . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
How a Flagellum Is Built: The Intraflagellar Transport (IFT). . . . . . . . . . . . . . . . . 93
How a Flagellar Motor Works . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93Internal Flagellar Structure. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
How a Paraflagellum Rod Works . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Photoreceptor Apparata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Type I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Type II . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Type III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
Photosensory Proteins and Methods for Their Investigation . . . . . . . . . . . . . . . . . . . . 100
Rhodopsin-Like Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Flavoproteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Action Spectroscopy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Absorption and Fluorescence Microspectroscopy . . . . . . . . . . . . . . . . . . . . . . . . . 102Biochemical and Spectroscopic Study of Extracted Visual Pigments . . . . . . . . . . . 103
Electrophysiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Molecular Biology Investigations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
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How Algae Use Light Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Photocycling Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Sensitivity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Noise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Direction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Guiding. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Trajectory Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
Signal Transmission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
An Example: Photoreceptor and Photoreception in Euglena. . . . . . . . . . . . . . . . . . 108
Chloroplasts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Cyanophyta and Prochlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
Glaucophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Rhodophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Heterokontophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Haptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Cryptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116Dinophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Euglenophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Chlorarachniophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Nucleus, Nuclear Division, and Cytokinesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Rhodophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Cryptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Dinophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Euglenophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 123
Ejectile Organelles and Feeding Apparata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Heterokontophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Haptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Cryptophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Dinophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Euglenophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
Chlorarachniophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Chapter 3
Photosynthesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Light. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Photosynthesis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Light Dependent Reactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
PSII and PSI: Structure, Function and Organization . . . . . . . . . . . . . . . . . . . . . . . 141
ATP-Synthase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
ETC Components . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Electron Transport: The Z-Scheme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Proton Transport: Mechanism of Photosynthetic Phosphorylation . . . . . . . . . . . . . 148
Pigment Distribution in PSII and PSI Super-Complexes ofAlgal Division. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Light-Independent Reactions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
RuBisCO . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
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Calvin Benson Bassham Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Carboxylation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Reduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Regeneration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Photorespiration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Energy Relationships in Photosynthesis: The Balance Sheet . . . . . . . . . . . . . . . . . . . . . 156
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
Chapter 4
Biogeochemical Role of Algae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Roles of Algae in Biogeochemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Limiting Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160
Algae and the Phosphorus Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
Algae and the Nitrogen Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
Algae and the Silicon Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Algae and the Sulfur Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Algae and the Oxygen/Carbon Cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Chapter 5
Working with Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
What is Light? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
How Light Behaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Scattering . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Absorption: Lambert Beer Law. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Interference. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Reflection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Refraction: Snells Law . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Dispersion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Diffraction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Field Instruments: Use and Application . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Radiometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Measurement Geometries, Solid Angles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Radiant Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Spectral Radiant Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Radiant Flux (Radiant Power) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191Spectral Radiant Flux (Spectral Radiant Power) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Radiant Flux Density (Irradiance and Radiant Exitance) . . . . . . . . . . . . . . . . . . . . . . 192
Spectral Radiant Flux Density . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Radiance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Spectral Radiance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Radiant Intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Spectral Radiant Intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Photometry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Luminous Flux (Luminous Power) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
Luminous Intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195
Luminous Energy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197Luminous Flux Density (Illuminance and Luminous
Exitance). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Luminance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
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Lambertian Surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Units Conversion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Radiant and Luminous Flux (Radiant and Luminous Power) . . . . . . . . . . . . . . . . . 199
Irradiance (Flux Density) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Radiance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Radiant Intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Luminous Intensity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Luminance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Geometries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
PAR Detectors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200
Photosynthesis Irradiance Response Curve (P versus E curve) . . . . . . . . . . . . . . . . . . . 202
Photoacclimation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
Chapter 6
Algal Culturing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Collection, Storage, and Preservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Culture Types . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Culture Parameters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Light. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
pH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Salinity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Mixing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Culture Vessels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Media Choice and Preparation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215Freshwater Media . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Marine Media . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Seawater Base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Nutrients, Trace Metals, and Chelators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226
Vitamins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Soil Extract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Buffers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 228
Sterilization of Culture Materials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Culture Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236
Batch Cultures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Continuous Cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Semi-Continuous Cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Commercial-Scale Cultures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Outdoor Ponds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Photobioreactors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
Culture of Sessile Microalgae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244
Quantitative Determinations of Algal Density and Growth . . . . . . . . . . . . . . . . . . . . . . 244
Growth Rate and Generation Time Determinations . . . . . . . . . . . . . . . . . . . . . . . . . . 249
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
Chapter 7Algae and Men . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Sources and Uses of Commercial Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
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Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
Cyanophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
Rhodophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256
Heterokontophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260
Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
Extracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Agar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270
Alginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272
Carrageenan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
Animal Feed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274
Fertilizers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Cosmetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
Therapeutic Supplements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
Toxin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285
Suggested Reading . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
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In memory of Maria Antonietta Barsanti (1957 2005), Mimmo Gualtieri (1954 2004),
and Patricia Lee Walne (19322004), who deserved something better
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