Light-Harvesting Antennas in Photosynthesis

533 Pages·2003·25.11 MB·English

by  Beverley R. Green

Light-Harvesting Antennas in Photosynthesis
Advances in Photosynthesis and Respiration
VOLUME 13
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, Osnabrück, 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 all life.Advances in Photosynthesis and Respirationis 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 all growth and
reproduction in these and almost all other organisms. As such, virtually all 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 medicine, 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
on the back cover.
Light-Harvesting Antennas
in Photosynthesis
Edited by
Beverley R. Green
University of British Columbia,
Vancouver, Canada
and
William W. Parson
University of Washington,
Seattle, U.S.A.
'~'
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
A C.I. P Catalogue record for this book is available from the Library of Congress.
ISBN 978-90-481-5468-5 ISBN 978-94-017-2087-8 (eBook)
DOI 10.1007/978-94-017-2087-8
The camera ready text was prepared by
Lawrence A. Orr, Center for the Study of Early Events in Photosynthesis,
Arizona State University, Tempe, Arizona 85287-1604, U.S.A.
Printed an acid-free paper
AII Rights Reserved
© 2003 S pri nger Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 2003
No part of this work may be reproduced, stored in a retrieval system, or transmitted
in any form or by any means, electronic, mechanical, photocopying, microfilming, recording
or otherwise, without written permission from the Publisher, with the exception
of any material supplied specifically for the purpose of being entered
and executed on a computer system, for exclusive use by the purchaser of the work.
This book is dedicated to our parents:
Mary Green (1911-2002), who supported all her daughter's professional
endeavors and was a constant presence during the writing of this book.
Rowland Green (1902-1971), who taught her to look at things with a
critical eye and ask pointed questions.
William Parson (1913-2002), who showed his son the excitement of
science.
Buffy Parson (1912- ), who reminds him that art is important too.
Editorial
Advances in Photosynthesis and Respiration
I am delighted to announce the publication of the (11) Regulation of Photosynthesis (Eva-Mari Aro
long-awaitedLight-HarvestingAntennas, Volume 13, and Berti! Andersson, editors, 2001 ); and
edited by Beverley R. Green and William W Parson (12) Photosynthetic Nitrogen Assimilation and
in our Series Advances in Photosynthesis and Associated Carbon and Respiratory Metab
Respiration. olism (Christine Foyer and Graham Noctor,
The present volume is a sequel to the following editors, 2002).
twelve volumes in the Advances in Photosynthesis
and Respiration (AIPH) series. See <http://www.wkap.nllseries.htm/AIPH/> for
further information and to order these books. Please
note that the members of the International Society of
Published Volumes Photosynthesis Research, ISPR (<http://www.
photosynthesisresearch.org/>) receive special
(1) Molecular Biology of Cyanobacteria (Donald discounts.
R. Bryant, editor, 1994);
(2) Anoxygenic Photosynthetic Bacteria (Robert
E. Blankenship, Michael T. Madigan and Carl Light-Harvesting Antennas
E. Bauer, editors, 1995);
In 1932, at the California Institute of Technology,
(3) Biophysical Techniques in Photosynthesis (Jan
Pasadena, Assistant Professor of Biophysics Robert
Amesz* and Arnold J. Hoff*, editors, 1996);
(Bob) Emerson (1903-1959) and his undergraduate
(4) Oxygenic Photosynthesis: The Light Reactions
student William (Bill) Arnold (1904- 2001) did the
(Donald R. Ort and Charles F. Yocum, editors,
most remarkable experiment. Using brief saturating
1996);
light flashes, spaced at optimum dark periods, they
(5) Photosynthesis and the Environment (Niel R.
measured 0 evolution in the green alga Chlorella by
2
Baker, editor, 1996);
manometry, a technique that had been perfected
(6) Lipids in Photosynthesis: Structure, Function earlier by Otto Warburg. The results were surprising:
and Genetics (Paul-Andre Siegenthaler and a maximum of only one 0 molecule was evolved per
2
Norio Murata, editors, 1998); 2,400 chlorophyll molecules present. The concept
(7) The Molecular Biology of Chloroplasts and that hundreds of chlorophyll molecules were
Mitochondria in Chlamydomonas (Jean David associated with each 'photoenzyme' was born.
Rochaix, Michel Goldschmidt-Clermont and Since the quantum yield of 0 evolution was high,
2
Sabeeha Merchant, editors, 1998); Hans Gaffron (1902-1979) and K. Wohl implied, in
(8) The Photochemistry ofCarotenoids (Harry A. 1936, that light energy absorbed by the bulk
Frank, Andrew J. Young, George Britton and chlorophyll molecules is transferred among these
Richard J. Cogdell, editors, 1999); molecules until it reaches the 'photoenzyme.' Now,
we know that the bulk chlorophyll molecules, bound
(9) Photosynthesis: Physiology and Metabolism
to proteins, serve as the 'antenna,' and the photo
(Richard C. Leegood, Thomas D. Sharkey and
enzyme is the 'reaction center.'
Susanne von Caemmerer, editors, 2000);
Volume 13 (Light-Harvesting Antennas), edited
(1 0) Photosynthesis: Photobiochemistry and Photo
by Beverley Green and William Parson, exposes
biophysics (Bacon Ke, author, 2001 );
before our very own eyes the structure and function
*deceased
of this intricate and marvelous machinery. The book present in organic molecules to fuel a wide range of
is unique in the degree to which it emphasizes the metabolic reactions critical for cell growth and
integration of molecular biological, biochemical and development. In addition, many photosynthetic
biophysical approaches. The collaboration of these organisms engage in energetically wasteful photo
two editors with very different areas of expertise respiration that begins in the chloroplast with an
shows the advantages of such an approach: Beverley oxygenation reaction catalyzed by the same enzyme
Green, whose photograph and a brief biography responsible for capturing carbon dioxide in
appears in this volume, is a biochemist who was one photosynthesis. This series of books spans topics
oft he major players in untangling the light-harvesting from physics to agronomy and medicine, from
pigment-protein complexes of higher plants and femtosecond (10.15 s) processes to season-long
chromophyte algae; Bill Parson, whose photograph production, from the photophysics ofr eaction centers,
and a brief biography also appears in this volume, is through the electrochemistry of intermediate electron
a 'very fast' biophysicist who is well-known for his transfer, to the physiology of whole organisms, and
studies of the sub-picosecond events in which light from X-ray crystallography of proteins to the
energy is absorbed by antenna systems and transferred morphology of organelles and intact organisms. The
to photosynthetic reaction centers. The two editors intent of the series is to offer beginning researchers,
have assembled an outstanding team of authors, all advanced undergraduate students, graduate students,
of international repute, representing a broad spectrum and even research specialists, a comprehensive, up
of interests, and coordinated the contributions to to-date picture ofthe remarkable advances across the
make a well-organized, understandable and compre full scope of research on bioenergetics and carbon
hensive volume. This book treats all aspects of metabolism.
photosynthetic light-harvesting antennas, from the
biophysical mechanisms of light absorption and
energy transfer to the structure, biosynthesis and Future Books
regulation of antenna systems in whole organisms. It
sets the great variety of antenna pigment-protein The readers of the current series are encouraged to
complexes in their evolutionary context and at the watch for the publication of the forthcoming books:
same time brings in the latest hi-tech developments.
Throughout, there is a consistent attempt to deal with (1) Photosynthesis in Algae (Editors: Anthony W.D.
both biophysical and molecular biological infor Larkum, Susan Douglas, and John A. Raven);
mation as it relates to different sides of the same (2) Chlorophylls and Bacteriochlorophylls:
question. It is 'a real book with chapters,' not a Biochemistry, Biophysics and Biological
compilation of review articles. Function (Editors: Bernhard Grimm, Robert J.
Porra, Wolfhart Rudiger and Hugo Scheer);
(3) Respiration in Archae and Bacteria. 2 volumes
The Scope of the Series
(Editor: Davide Zannoni );
(4) Chlorophyll a Fluorescence: A signature of
Advances in Photosynthesis and Respiration is a
Photosynthesis (Editors: George Papageorgiou
book series that provides, at regular intervals, a
and Govindjee);
comprehensive and state-of-the-art account of
(5) Plant Respiration (Editors: Miquel Ribas
research in various areas of photosynthesis and
Carbo and Hans Lambers);
respiration. Photosynthesis is the process by which
higher plants, algae, and certain species of bacteria (6) Photosystem II: The Water/Plastoquinone
transform and store solar energy in the form of Oxido-reductase in Photosynthesis (Editors:
energy-rich organic molecules. These compounds Thomas J. Wydrzynski and Kimiyuki Satoh);
are in turn used as the energy source for all growth (7) Photosystem 1: The NADP+/Ferredoxin
and reproduction in these and almost all other Oxidoreductase in Oxygenic Photosynthesis
organisms. Virtually all life on the planet thus (Editor: John Golbeck);
ultimately depends on photosynthetic energy (8) Photosynthesis: A Comprehensive Treatise;
conversion. Respiration, which occurs in mito Biochemistry, Biophysics and Molecular
chondria and in bacterial membranes, utilizes energy
Vlll
Biology, 2 volumes (Editors: Julian Eaton-Rye Plants; and C-3 and C-4 Plants.
and Baishnab Tripathy) Readers are requested to send their suggestions
(9) Photoprotection, Photoinhibition, Gene for these and future volumes (topics, names of future
Regulation and Environment (Editors: Barbara editors, and of future authors) to me by E-mail
Demmig-Adams, William W Adams III and ([email protected]) or fax [phone]).
Aut ar Mattoo ); In view of the interdisciplinary character of
research in photosynthesis and respiration, it is my
(1 0) The Structure and Function ofPlastids (Editors:
earnest hope that this series of books will be used in
Kenneth Hoober and Robert Wise); and
educating students and researchers not only in Plant
(II) History of Photosynthesis Research (Editor:
Sciences, Molecular and Cell Biology, Integrative
Govindjee)
Bi.ology, Biotechnology, Agricultural Sciences,
Microbiology, Biochemistry, and Biophysics, but
In addition to these contracted books, we are
also in Bioengineering, Chemistry, and Physics.
interested in publishing several other books. Topics
I take this opportunity to thank Beverley Green;
planned are: Global Aspects of Photosynthesis and
William Parson; all the authors ofVolume 13; Larry
Respiration; Protein Complexes of Photosynthesis
Orr; Jacco Flipsen, Noeline Gibson· Evan Delucia
and Respiration; Biochemistry and Biophysics of
and my wife Rajni Govindjee for th;ir valuable hel~
Respiration; Protonation and ATP Synthesis;
and support that made the publication of Light
Functional Genomics; Proteonomics, The Cyto
Harvesting Antennas possible.
chromes; Molecular Biology of Cyanobacteria;
Laboratory Methods for Studying Leaves and Whole
Govindjee
Series Editor
Advances in Photosynthesis and Respiration
University of Illinois at Urbana-Champaign
Department of Plant Biology
265 Morrill Hall, 505 South Goodwin Avenue
Urbana, IL 61801-3707, U.S.A.
E-mail: [email protected];
URL: http://www.life.uiuc.edu/govindjee
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Govindjee
The story oft he Series editors name and a bit ofh is initials of his doctoral students: the longest being
personal background. J.C.M. Govindjee, Jr (where J.C.M. stood for John
Govindjee, who uses one name only, grew up in Clingman Munday). Govindjee received his training
Allahabad (India). He was born in 1932 to (Mrs) in the area of 'Photosynthesis' first from Robert
Savitri Devi Asthana and (Mr) Vishveshwar Prasad Emerson, and then from Eugene Rabinowitch.
As thana. His father was a college teacher, and then a Govindjee obtained his Ph.D. in Biophysics at the
representative of the Oxford University Press in the University of Illinois, in 1960. He has been on its
then United Provinces (UP) of India. His family faculty since 1961. Since July, 1999 he has been
name, As thana, was dropped by his father in response Professor Emeritus ofBiochemistry, Biophysics and
to the 'Arya Samaj Movement' that was against the Plant Biology at the University of Illinois at Urbana
'Caste System,' and believed in the ideals of the Champaign. A brief formal biography ofGovindjee
Vedic times. After his father's death in 1943, his is available in Foyer CHand Noctor G (eds) (2003)
older brother Krishnaji (Professor of Physics, Photosynthetic NitrogenAssimilation andA ssociated
University ofA llahabad) was responsible for bringing Carbon and Respiratory Metabolism, Advances in
him up and served as his role model. Shri Ranjan, Photosynthesis and Respiration, Volume 12, Kluwer
who had been a graduate student of Felix Frost Academic Publishers, Dordrecht, The Netherlands.
Blackman, trained him in Plant Physiology. He was married to Rajni Verma in 1957 at Urbana,
Govindjee came to Urbana, Illinois, U.S.A., as a Illinois. They have two children: Sanjay Govindjee
Fulbright Scholar, and a Fellow of'Physico-Chemical (his wife is Marilyn) and Anita Govindjee (her
Biology' in 1956, with his name written as 'Govind husband is Morten Christiansen) and three grand
lee.' He was not happy to be called 'G. lee,' and, thus, children (Sunita Christiansen; Arjun Govindjee and
began to use 'Govindjee' as his one and only name. Rajiv Govindjee).
This has caused problems in citations, and in many Currently, Govindjee focuses his attention on two
formal settings. He has been called by many names: topics: (1) Chlorophyll a Fluorescence: A Probe of
N.F.N. Govindjee (where N.F.N. stands for No First Photosynthesis; and (2) History of Photosynthesis
Name); I. Govindjee (where I stands for Illini); Mister Research; he is equally concerned with photo
Govindjee; and once A.VP. Govindjee (where A. synthesis education (see his web site: http://
stands for Allahabad, and VP. are the initials of his www.life.uiuc.edu/govindjee).
father). Quite often, his name has appeared with the
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