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Photosynthesis in Algae Advances in Photosynthesis and Respiration Photosynthesis in Algae Advances in Photosynthesis and Respiration VOLUME 14 Series Editor: GOVINDJEE University of Illinois, Urbana, Illinois, U.S.A. Consulting Editors: Christine FOYER, Harpenden, u.K. Elisabeth GANTT, College Park, Maryland, U.S.A. John H. GOLBECK, University Park, Pennsylvania, U.S.A. Susan S. GOLDEN, College Station, Texas, U.S.A. Wolfgang JUNGE, Osnabrilck, Germany Hartmut MICHEL, Frankfurt am Main, Germany Kirmiyuki SATOH, Okayama, Japan James Siedow, Durham, North Carolina, U.S.A. The scope of our series, beginning with volume 11, reflects the concept that photosynthesis and respiration are intertwined with respect to both the protein complexes involved and to the entire bioenergetic machinery of aII life. Advances in Photosynthesis and Respiration is a book series that provides a comprehensive and state-of-the-art account of research in photo­ synthesis and respiration. Photosynthesis is the process by which higher plants, algae, and certain species of bacteria transform and store solar energy in the form of energy-rich organic molecules. These compounds are in turn used as the energy source for aII growth and reproduction in these and almost aII other organisms. As such, virtually aII life on the planet ultimately depends on photosynthetic energy conversion. Respiration, which occurs in mitochondrial and bacterial membranes, utilizes energy present in organic molecules to fuel a wide range of metabolic reactions critical for cell growth and development. In addition, many photosynthetic organisms engage in energetically wasteful photorespiration that begins in the chloroplast with an oxygenation reaction catalyzed by the same enzyme responsible for capturing carbon dioxide in photosynthesis. This series of books spans topics from physics to agronomy and medici ne, from femtosecond processes to season long production, from the photophysics of reaction centers, through the electrochemistry of intermediate electron transfer, to the physiology of whole orgamisms, and from X-ray christallography of proteins to the morphology or organelles and intact organisms. The goal of the series is to offer beginning researchers, advanced undergraduate students, graduate students, and even research specialists, a comprehensive, up-to-date picture of the remarkable advances across the full scope of research on photosynthesis, respiration and related processes. The titles published in this series are listed at the end of this volume and those of forthcoming volumes an the back cover. Photosynthesis in Algae Edited by Anthony W.D. Larkum School of Biological Sciences, University of Sydney, Australia Susan E. Douglas National Research Council, Institute for Marine Biosciences, Canada and John A. Raven Department of Biological Sciences, University of Dundee, u.K. Springer-Science+Business Media, BV A C.I.P. Catalogue record lor this book is available Irom the Library 01 Congress. ISBN 978-94-010-3772-3 ISBN 978-94-007-1038-2 (eBook) DOI 10.1007/978-94-007-1038-2 The cover picture is a conlocal scanning microscope picture 01 the gyres 01 chloroplasts in an undescribed species 01 Spirogyra which has "Iri Ils" along the margins. The light areas are putative grana (see Chapter 2). The picture was taken by Brian Gunning in lalse colour using a Biorad MRC-600 conlocal microscope with a projection 0126 optical sections imaged at 0.4 micrometre locusing intervals with a 1OOx Planapo oil-immersion Zeiss lens. Printed an acid-free paper AII Rights Reserved © 2003 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 2003 Softcover reprint of the hardcover 1 st edition 2003 No part 01 this work may be reproduced, stored in a retrieval system, or transmitled in any lorm or by any means, electronic, mechanical, photocopying, microlilming, recording or otherwise, without writlen permission Irom the Publisher, with the exception 01 any material supplied specilically lor the purpose 01 being entered and executed on a computer system, lor exclusive use by the purchaser 01 the work. Editorial Advances in Photosynthesis and Respiration Volume 14: Photosynthesis in Algae I am extremely delighted to announce the (12) Photosynthetic Nitrogen Assimilation and publication of the long-awaited Photosynthesis in Associated Carbon and Respiratory Metab­ Algae edited by Anthony Larkum, Susan Douglas olism (Christine Foyer and Graham Noctor, and John Raven. It is Volume 14 in our Series editors, 2002); and Advances in Photosynthesis andRespiration (AIPH) (13) Light Harvesting Antennas (Beverley Green and is a sequel to the previous thirteen volumes in the and William Parson, editors, 2003). series. See <http://www.wkap.nl/series.htrnlAIPH> for further information and to order these books. Please Published Volumes note that the members ofthe International Society of Photosynthesis Research, ISPR (<http://www. (1) Molecular Biology ofCyanobacteria (Donald Photosynthesisresearch.org», receive special R. Bryant, editor, 1994); discounts. (2) Anoxygenic Photosynthetic Bacteria (Robert E. Blankenship, Michael T. Madigan and Carl E. Bauer, editors, 1995); Photosynthesis in Algae (3) Biophysical Techniques in Photosynthesis (Jan Amesz* and Arnold J. HoW, editors, 1996); Algae are a fascinating group of organisms, which refuse to be classified. One may simply place them (4) Oxygenic Photosynthesis: The LightReactions in the Protists, one of the six Kingdoms of Nature. (Donald R. art and Charles F. Yocum, editors, However this ignores the fact that they arc 1996); photosynthetic and are undeniably related in some (5) Photosynthesis and the Environment (Neil R. way to the land plants (Plantae). It also places the Baker, editor, 1996); algal group, which represents some of the largest (6) Lipids in Photosynthesis: Structure, Function organisms on the planet in a Kingdom that is typified and Genetics (Paul-Andre Siegenthaler and by microscopic unicellular organisms! Of course Norio Murata, editors, 1998); this has not deterred workers in photosynthesis, (7) The Molecular Biology of Chloroplasts and almost all ofwhom will have a flask or two ofalgae Mitochondria in Chlamydomonas (Jean David growing somewhere in their laboratories, even though Roehaix, Michel Goldschmidt-Clermont and they devote their main work to higher plants. As Sabeeha Merchant, editors, 1998); pointed out in the Preface to this volume, it also does (8) The Photochemistry ofCarotenoids (Harry A. not prevent frequent sallies into the domain of Frank, Andrew J. Young, George Britton and Bacteria, to make use ofthephotosynthetic properties Richard J. Cogdell, editors, 1999); of Cyanobacteria, even though these organisms are (9) Photosynthesis: Physiology and Metabolism no longer called 'algae.' (Richard C. Leegood, Thomas D. Sharkey and We are fortunate in having three outstanding and Susanne von Caemmerer, editors, 2000); charming editors whose interests have allowed them a clear overview ofa complex field and has allowed (10) Photosynthesis: Photobiochemistry and Photo­ them to choose an excellent set of authors out of a biophysics (Bacon Ke, author, 2001); army ofgreat researchers in the field. Tony Larkum (11) Regulation of Photosynthesis (Eva-Mari Aro is basically a plant physiologist but has worked at all and Bertil Andersson, editors, 2001) levels ofphotosynthesis from whole ecosystems such "deceased as coral reefs to genes for the light-harvestingproteins of dinoflagellates and prochlorophytes. Susan research in various areas of photosynthesis and Douglas is a molecular biologist who has sequenced respiration. Photosynthesis is the process by which cryptophyte genes and most recently took a major higher plants, algae, and certain species of bacteria role in sequencing the three chromosomes of the transform and store solar energy in the form of relic nucleus (nucleomorph) of the cryptophyte energy-rich organic molecules. These compounds Guillardia theta. John Raven is an eclectic algal are in turn used as the energy source for all growth physiologist, whose major contributions have been and reproduction in these and almost all other in quantifying the roles of various pathways in the organisms. As such, virtually all life on the planet carbon metabolism of algae as well as contributing ultimately depends on photosynthetic energy prolifically to discussion on the evolution ofCyano­ conversion. Respiration, which occurs in mito­ bacteria, algae and land plants. chondria and in bacterial membranes, utilizes energy Knowing the editors and the topic of the specific present in organic molecules to fuel a wide range of volume are important to me. We have included brief metabolic reactions critical for cell growth and biographies and photographs of the editors in this development. In addition, many photosynthetic volume. My interest in algae dates back to the time organisms engage in energetically wasteful photo­ when I was a graduate student of Robert Emerson respiration that begins in the chloroplast with an (who himself was a student of Otto Warburg); the oxygenation reaction catalyzed by the same enzyme green alga Chlorella was the choice ofresearch then. responsible for capturing carbon dioxide in photo­ A 1957 group photographand a shamelesslyproduced synthesis. This series of books spans topics from listing of some of our findings are given following physics to agronomy and medicine, from femtosecond this 'Editorial.' I have been fortunate to know Tony (10. 15 s) processes to season-long production, from the most. I have been his guest once at
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