Algae Anatomy, Biochemistry, and Biotechnology Algae Anatomy, Biochemistry, and Biotechnology Laura Barsanti Paolo Gualtieri Boca Raton London New York A CRC title, part of the Taylor & Francis imprint, a member of the Taylor & Francis Group, the academic division of T&F Informa plc. 1467_Discl.fm Page 1 Friday, September 30, 2005 2:40 PM 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, MA 01923, 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 Taylor & Francis Group and the CRC Press Web site at is the Academic Division of Informa plc. http://www.crcpress.com Preface This book is an outgrowth of many years of research aimed at studying algae, especially micro- algae.Workingonit,wesoonrealizedhowsmallanareawereallyknewwellandhowsuperficial 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, sacrificingcompleteness insodoing. 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,itisdestinedtoserveasameanstoencourageoutstandingworkinthefieldofphycology, 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, soas tolet new ideas springto mind bythe endof 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 isincompletely knownand prepared investigatorsshould 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 sothattheresearchworkercanentertheliteraturetofindoutmoreprecisedetailsfromtheoriginal sources. Thebibliographyisbynomeansexhaustive;thepaperswehavequotedaretheoneswehave found useful and which are reasonably accessible, both very recent references and older classic references 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 importantfacts,hopingthatthisbookwillcomplementthefewspecializedreviewsoffinestructure 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 interestingand useful Web addresses. Wearegratefultothephycologistswhohavecontributedoriginalpictures;theyarecitedinthe correspondingfigurecaptions.WearealsogratefultothestaffatCRCPress,BocaRaton,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. Oursinceregratitudeandaspecial thankstoValterEvangelistaforhisskillfulassistanceand abilityinpreparingthefinalformofallthedrawingsandillustrations,andforhiscarefulattention inpreparingallthetechnicaldrawings.Weappreciatedhiseffortstokeeppacewithusbothandto cope with ourever-changing demands without getting tooupset. WewillalwaysbegratefultoVincenzoPassarelli,whofrequentlysmoothedapathstrewnwith 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 notalways 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 suchadisablingsituation,shesucceededinpreparingmostofthedrawings,withcarefuldetermi- 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 begratefultohim. InJuly2004,MimmoGualtieri,theonlybrotherofPaolo,diedofanunexpectedheartattack. He left ahuge empty room inhis brother’sheart. 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. Thisbook isdedicated tothe threeofthem. About the Authors Dr.LauraBarsantigraduatedinnaturalsciencefromUniversityofPisa,Italy.Atpresentsheisa scientistat the Biophysics Institute ofthe National CouncilofResearch (CNR) inPisa. Dr.PaoloGualtierigraduatedinbiologyandcomputersciencefromUniversityofPisa,Italy.At presentheisaseniorscientistattheBiophysicsInstituteoftheNationalCouncilofResearch(CNR) in Pisa and adjunct professor at the University of Maryland, University College, College Park, Maryland. Table of Contents Chapter1 GeneralOverview. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . 1 Definition . . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 1 Classification.. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 2 Occurrence and Distribution. .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 2 Structure ofThallus. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 3 UnicellsandUnicellColonial Algae. . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 3 Filamentous Algae.. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 5 Siphonous Algae. .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 5 Parenchymatousand PseudoparenchymatousAlgae. . .. . . .. . . .. . .. . . .. . . .. . . .. 6 Nutrition.. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 7 Reproduction.. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 7 Vegetative andAsexual Reproduction .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 8 Binary Fission orCellularBisection. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 8 Zoospore, Aplanospore, and Autospore. .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 9 Autocolony Formation. . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 9 Fragmentation.. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 10 Resting Stages.. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 10 Sexual Reproduction . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 11 Haplonticor Zygotic Life Cycle. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 14 Diplontic orGametic Life Cycle . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 14 Diplohaplonticor Sporic Life Cycles . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 14 Summaries of the Ten Algal Divisions.. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 15 Cyanophyta andProchlorophyta. .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 16 Glaucophyta .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 19 Rhodophyta. .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 20 Heterokontophyta .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 20 Haptophyta. .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 21 Cryptophyta .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 23 Dinophyta. . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 24 Euglenophyta . . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 26 Chlorarachniophyta. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 26 Chlorophyta .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 27 Endosymbiosis andOriginof EukaryoticAlgae . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 29 Suggested Reading . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. 33 Chapter2 Anatomy . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . 35 Cytomorphology and Ultrastructure . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . 35 Outsidethe Cell . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. 35 Type1: Simple Cell Membrane. . .. . . .. . . .. . . .. . .. . . .. . . .. . . .. . .. . . .. . 35 Type2: Cell Surface with AdditionalExtracellularMaterial .. . . .. . . .. . .. . . .. . 36 Mucilages andSheaths. .. . . .. . . .. . .. . . .. . . .. . . .. . . .. . .. . . .. . . .. . . . 36