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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

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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


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


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


14/17

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


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


Chapter 7Algae and Men . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251

Sources and Uses of Commercial Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252


15/17

Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252

Cyanophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252

Rhodophyta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 256


Chlorophyta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267

Extracts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269

Agar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270

Alginate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272

Carrageenan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273

Animal Feed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 274

Fertilizers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278

Cosmetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280

Therapeutic Supplements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281

Toxin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285


Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293


16/17

In memory of Maria Antonietta Barsanti (1957 2005), Mimmo Gualtieri (1954 2004),

and Patricia Lee Walne (19322004), who deserved something better


17/17

algae anatomy biochemistry 0

Documents