Table Of ContentPlant Mitochondria: From Genome to Function
Advances in Photosynthesis and Respiration
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VOLUME 17
Series Editor:
GOVINDJEE
University of Illinois, Urbana, Illinois, U.S.A.
Consulting Editors:
Christine FOYER, Harpenden, U.K.
Elisabeth GANTI, 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, OsnabrOck, Germany
Hartmut MICHEL, Frankfurt am Main, Germany
Kimiyuki 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 Respiration is a book series that
provides a comprehensive and state-of-the-art account of research in photosynthesis and respira
tion. 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 pho
torespiration that begins in the chloroplast with an oxgenation reaction catalyzed by the same
enzyme responsible for capturing carbon dioxide in photosynthesis. This series of books spans top
ics from physics to agronomy and medicine, from femtosecond processes to season long produc
tion, from the photophysics of reaction centers, through the electrochemistry of intermediate
electron transfer, to the physiology of whole organisms and from X-ray crystallography of proteins
to the morphology of 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.
Plant Mitochondria:
From Genome to Function
Edited by
David A. Day
A. Harvey Millar
and
James Whelan
School of Biomedical and Chemical Sciences, The University of
Western Australia, Australia
Springer-Science+Business Media, B.V.
A C.I.P. Catalogue record for this book is available from the library of Congress.
ISBN 978-90-481-6651-0 ISBN 978-1-4020-2400-9 (eBook)
DOI 10.1007/978-1-4020-2400-9
Printed on acid-free paper
Cover photograph: The cover illustration is a collage of two photographs. It shows an Arabidopsis flower
expressing the reporter gene GUS under the control of the soybean Aox2b promoter (expression is denoted
by the blue colour), superimposed on a confocal micrograph of an Arabidopsis cell expressing
green fluorescent protein (GFP) targeted to mitochondria.
All Rights Reserved
© 2004 Springer Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 2004.
Softcover reprint of the hardcover 1st edition 2004
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.
Editorial
Advances in Photosynthesis and Respiration
T am delighted to announce the publication, in the (Christine Foyer and Graham Noctor, editors,
Advances in Photosynthesis and Respiration (AIPH) 2002);
Series, the first book related to plant respiration (Plant (13) Light Harvesting Antennas (Beverley Green and
Mitochondria: From Genome to Function, edited by William Parson, editors, 2003);
David A. Day, A. Harvey Millar and James Whelan). (14) Photosynthesis in Algae (Anthony Larkum,
Volumes 15 and 16 (both edited by Davide Zannoni) Susan Douglas and John Raven, editors, 2003);
dealt with "Respiration in Bacteria and Archaea." ( 15) Respiration in Archaea and Bacteria: Diversity
Volume 17 is a sequel to the sixteen volumes in the of Prokaryotic Electron Transport Carriers
AIPH Series. Volume 18 (edited by Hans Lambers and (Davide Zannoni, editor, 2004);
Miquel Ribas-Carbo), soon to be published, will also (16) Respiration inArchaea and Bacteria 2: Diversity
deal with Plant Respiration, but will focus on physio of Prokaryotic Respiratory Systems (Davide
logical and ecological aspects. Zannoni, editor, 2004)
See <http://www.wkap.nllseries.htm/ AI PH> for
Published Volumes
further information and to order these books. Please
(1) Molecular Biology of" Cyanobacteria (Donald A. note that the members of the International Society
of Photosynthesis Research, ISPR (<http://www.
Bryant, editor, 1994);
(2) Anoxygenic Photosynthetic Bacteria (Robert E. photosynthesisresearch.org>) receive special discounts.
Blankenship, Michael T. Madigan and Carl E.
Bauer, editors, 1995); Plant Mitochondria: From Genome to
(3) Biophysical Techniques in Photosynthesis (Jan Function
Ameszt and Arnold J. Hofft, editors, 1996);
(4) Oxygenic Photosynthesis: The Light Reactions This book provides an up-to-date and in-depth review
(Donald R. Ort and Charles F. Yocum, editors, of the the molecular aspects of plant mitochondrial
1996); function, with a focus on gene regulation and protein
(5) Photosynthesis and the Environment (Neil R. composition. The chapters are written by experts in the
Baker, editor, 1996); field who are actively researching in the areas on
(6) Lipids in Photosynthesis: Structure. Function which their chapters focus. Each chapter, except 1
and Genetics (Paul-Andre Siegenthaler and and 2, has multiple authors, usually from different
Norio Murata, editors, 1998); laboratories and countries, ensuring a balanced and
(7) The Molecular Biology of Chloroplasts and comprehensive review of the different aspects of
Mitochondria in Chlamydomonas (Jean David mitochondrial molecular biology. The three editors are
Rochaix, Michel Goldschmidt-Clermont and from the very active mitochondrial research group
Sabeeha Merchant, editors, 1998); established at the University of Western Australia and
(8) The Photochemistry ol Carotenoids (Harry A. have used their complementary knowledge of bio
Frank, Andrew J. Young, George Britton and chemistry, molecular genetics and proteomics to com
Richard J. Cogdell, editors, 1999); pile an exciting view of cutting edge research on plant
(9) Photosynthesis: Physiology and Metabolism mitochondria. The book will complement the more
(Richard C. Leegood, Thomas D. Sharkey and physiologically-orientated Volume 18 edited by Hans
Susanne von Caemmerer, editors, 2000); Lambers and Miquel Ribas-Carbo.
( l 0) Photosynthesis: Photobiochemistry and The two main themes running through the book are
Photobiophysics (Bacon Ke, author, 200 I); the interconnection between gene regulation and pro
(ll) Regulation ofP hotosynthesis (Eva-Mari Aro and tein function, and the integration of mitochondria with
Berti! Andersson, editors, 200 l ); other components of plant cells. The book begins
(12) Photosynthetic Nitrogen Assimilation and with two overviews on the dynamics of mitochondrial
Associated Carbon and Respiratory Metabolism structure, function, morphology and inheritance. It
., Deceased.
VI Advances in Photosynthesis and Respiration
then discusses the biogenesis of mitochondria, the In 1932, the year I was born, Walter Stiles and William
regulation of gene expression, the mitochondrial Leach wrote their small (124-page) book Respiration
genome and its interaction with the nucleus, and the in Plants (London: Methuen). The theories ofWarburg
targeting of proteins to the organelle. This is followed and Wieland on the oxidation reduction and the enzy
by a discussion of the contributions that mutations, matic nature of respiration, under both aerobic and
involving mitochondrial proteins, have made to our anaerobic conditions, were discussed in this book. It
understanding of the way the organelle interacts with was the only book I had read during my student days.
the rest of the plant cell, and the new field of In 1953, one year after I obtained my BSc degree, a
proteomics and the discovery of new functions. Also new book was published (W.O. James ( 1952) Plant
covered are the pathways of electron transport, with Respiration. Oxford, at the Clarendon Press). It is this
special attention to the non-phosphorylating bypasses, book that I studied after I had obtained my MSc
metabolite transport, and specialized mitochondrial degree. On the personal side, I was thrilled to note that
metabolism. Finally, the impact of oxidative stress on James discussed (see pp. 99 and I 00) unpublished
mitochondria and the defense mechanisms that are work of my Professor (Ranjan); this work was done
employed to allow survival are discussed. This book is while Ranjan was in Blackman's lab. It was a 282-
for the use of advanced undergraduates, graduates, page, thorough and modern book. James also related
postgraduates, and beginning researchers in the areas respiration, although very briefly, to photosynthesis
of molecular and cellular biology, integrative biology, and commented "Possible interactions with respiration
biochemistry, bioenergetics, proteomics, and plant and have been the bugbear of photosynthetic measure
agricultural sciences. ments since their beginning." He did mention the
related work of Bessel Kok, Robert Emerson, Jack
Myers, James Franck, Hans Gaffron, and Melvin
Calvin and Andy Benson, among others. A 19-page
Plant Respiration Books: A Bit of bibliography was very helpful to me in obtaining the
Early History necessary information. The advances made during the
last 50 years are really remarkable and exciting. To
It is a noble employment to rescue from oblivion those understand the historical evolution of research in
who deserve to be remembered (Pliny the Younger, "plant respiration," the reader is encouraged to consult
Letters V). the following three books: (I) Harry Beevers (1961)
Respiratory Metabolism in Plants (Evanston, Illinois:
I mention here the books by Blackman, Stiles and Row, Peterson and Company); (2) Helgi Opik (1980)
Leach, and James. Although we may not remember the The Respiration of Higher Plants (London:
old, the new is built upon the old directly or indirectly, E. Arnold); (3) Roland Douce and David A. Day (e ds)
knowingly or unknowingly. My own training in Plant (1985) Higher Plant Cell Respiration. Encyclopedia
Respiration, during 1952-1956 at the University of of Plant Physiology, New Series, Volume 18, Berlin:
Allahabad, was under Shri Ranjan, who had been a Springer-Verlag.
student of Felix Frost Blackman. Blackman studied Fascinating reading awaits us in the AIPH books
both respiration and photosynthesis. G.E. Briggs edited by David Day, Harvey Millar and James
(of Cambridge, UK) has published a nice book on the Whelan (Volume 17) and by Hans Lambers and
work of Blackman after his death (Late F.F. Blackman Miquel Ribas-Carbo (Volume 18). Good hunting!
(1954) Analytic Studies in Plant Respiration.
Cambridge, at the University Press.). A great deal of
the experimental work in Blackman's laboratory was
done by one of his students from India, P. Parija. Most The Scope of the Series
of this work was done in the late 1920s at Cambridge.
Regarding the importance of respiration, I quote Advances in Photosynthesis and Respiration is a book
Blackman and Parija (1928) "Of all protoplasmic series that provides, at regular intervals, a comprehen
functions, the one which is, by tradition, most closely sive and state-of-the-art account of research in various
linked with our conception of vitality is the function areas of photosynthesis and respiration. Photosyn
for which the name of respiration has been accepted." thesis is the process by which higher plants, algae, and
Advances in Photosynthesis and Respiration Vll
certain species of bacteria transform and store solar (4) Photosystem II: The Water/Plastoquinone
energy in the form of energy-rich organic molecules. Oxidoreductase in Photosynthesis (Editors:
These compounds are in turn used as the energy Thomas J. Wydrzynski and Kimiyuki Satoh);
source for all growth and reproduction in these (5) Photosystem /: The Plastocyanin/Ferredoxin
and almost all other organisms. As such, virtually Oxidoreductase in Oxygenic Photosynthesis
all life on the planet ultimately depends on photosyn (Editor: John Golbeck);
thetic energy conversion. Respiration, which occurs in (6) Photoprotection, Photoinhibition, Gene Regulation
mitochondria and in bacterial membranes, utilizes and Environment (Editors: Barbara Demmig
energy present in organic molecules to fuel a wide Adams, William W. Adams III and Autar Mattoo );
range of metabolic reactions critical for cell growth (7) Photosynthesis: A Comprehensive Treatise;
and development. In addition, many photosynthetic Biochemistry, Biophysics and Molecular Biology,
organisms engage in energetically wasteful photores 2 volumes (Editors: Julian Eaton-Rye and
piration that begins in the chloroplast with an Baishnab Tripathy);
oxygenation reaction catalyzed by the same enzyme (8) The Structure and Function of Plastids (Editors:
responsible for capturing carbon dioxide in photo Kenneth Hoober and Robert Wise); and
synthesis. This series of books spans topics from (9) History of Photosynthesis Research (Editors:
physics to agronomy and medicine, from femtosecond Govindjee, John Allen, J. Thomas Beatty and
oo-
15 s) processes to season-long production, from Howard Gest).
the photophysics of reaction centers, through the
electrochemistry of intermediate electron transfer, In addition to these contracted books, we are inter
to the physiology of whole organisms, and from ested in publishing several other books. Topics under
X-ray crystallography of proteins to the morphology consideration are: Molecular Biology of Stress in
of organelles and intact organisms. The intent of Plants; Global Aspects of Photosynthesis and
the series is to offer beginning researchers, advanced Respiration; Protein Complexes of Photosynthesis and
undergraduate students, graduate students, and Respiration; Protonation and ATP Synthesis;
even research specialists, a comprehensive, up-to Functional Genomics; The Cytochromes; Laboratory
date picture of the remarkable advances across the Methods for Studying Leaves and Whole Plants; and
full scope of research on bioenergetics and carbon C-3 and C-4 Plants.
metabolism. Readers are requested to send their suggestions for
these and future volumes (topics, names of future
editors, and of future authors) to me by e-mail
<[email protected]> or fax ( + 1-217-244-7246).
Future Books In view of the interdisciplinary character of
research in photosynthesis and respiration, it is my
The readers of the current series are encouraged earnest hope that this series of books will be used
to watch for the publication of the forthcoming in educating students and researchers not only in Plant
books (not necessarily arranged in the order of future Sciences, Molecular and Cell Biology, Integrative
appearance): Biology, Biotechnology, Agricultural Sciences,
Microbiology, Biochemistry, and Biophysics, but also
in Bioengineering, Chemistry, and Physics.
(I) Plant Respiration. From Cell to Ecosystem I take this opportunity to thank David Day and his
(Edited by Hans Lambers and Miquel Ribas co-editors (Harvey Millar and James Whelan) for
Carbo); their timely and prompt editorial work. I thank all the
(2) Chlorophylls and Bacteriochlorophylls: Bio authors of volume 17: without their authoritative chap
chemistry, Biophysics and Biological Function ters, there will be no book. I owe Noeline Gibson spe
(Editors: Bernhard Grimm, Robert J. Porra, cial thanks for her friendly and wonderful working
Wolfhart Rudiger and Hugo Scheer); connection with the production of this book. Thanks
(3) Chlorophyll a Fluorescence: A Signature of are also due to Jacco Flipsen (also of Kluwer
Photosynthesis (Editors: George Papageorgiou Academic Publishers), and Jeff Haas (Director of
and Govindjee); Information Technology, Life Sciences, University of
Vlll Advances in Photosynthesis and Respiration
Illinois) :tiJr their support. My wife Rajni Govindjee and her husband Morten Christiansen provided
deserves my special praise for being a role model for facilities at the time this book was being prepared for
my life and well-being. Our daughter Anita Govindjee publication.
Govindjee
Series Editor
Advances in Photosynthesis and Respiration
University of Illinois at Urbana-Champaign,
Department of Plant Biology
265 Morrill Hall, MC-116, 505 South Goodwin Avenue,
Urbana, IL 61801-3707, USA
E-mail: <[email protected]>
URL: <http://www.life.uiuc.edu/govindjee>
Series Editor: Govindjee
Govindjee is Professor Emeritus of Biochemistry, the primary photochemistry; role of bicarbonate in the
Biophysics and Plant Biology at the University of electron and proton transport; thermoluminescence,
Illinois at Urbana-Champaign (UIUC), Illinois, USA. delayed and prompt fluorescence (particularly life
He received his PhD in Biophysics from the University times), and their use in understanding electron trans
of Illinois at Urbana-Champaign in 1960, with a thesis port and photoprotection against excess light. He has
on the "Action Spectra of the Emerson Enhancement co-authored Photosynthesis (1969); and has edited (or
Effect in Algae," under Eugene Rabinowitch. From co-edited) Bioenergetics of Photosynthesis (1975);
1960 to 1961, he served as a United States Public Photosynthesis (in 2 volumes, 1982); Light Emission
Health (USPH) Postdoctoral Fellow; from 1961 to by Plants and Bacteria ( 1986), among other books. He
1965, as Assistant Professor of Botany; from 1965 to is a member of the American Society of Plant Biology,
1969 as Associate Professor of Biophysics and American Society for Photobiology, Biophysical
Botany; and from 1969 to 1999 as Professor of Society of America, and the International Society of
Biophysics and Plant Biology, all at the UIUC. In Photosynthesis Research (ISPR). Govindjee's scien
1999, he became Professor Emeritus of Biochemistry, tific interest, now, includes Fluorescence Lifetime
Biophysics and Plant Biology at UIUC. Julian Eaton Imaging Microscopy (FLIM) (with Robert Clegg) and
Rye, Prasanna Mohanty, George Papageorgiou, Alan regulation of excitation energy transfer in oscillating
Stemler, Thomas Wydrzynski, Jin Xiong, Chunhe Xu light (with Lada Nedbal). In addition, Govindjee is
and Barbara Zilinskas are among his more than 20 interested in "History of Photosynthesis Research,"
PhD students. His honors include: Fellow of the and in "Photosynthesis Education." His personal
American Association of Advancement of Science background appears in Volume 13 (edited by B. Green
( 1976); Distinguished Lecturer of the School of Life and W Parson); and contributions to photosynthesis
Sciences, UIUC ( 1978); President of the American and fluorescence in algae in Volume 14 (A. Larkum,
Society of Photobiology ( 1980-1981 ); Fulbright J. Raven and S. Douglas, editors) of the Advances
Senior Lecturer ( 1996-1997); and honorary President in Photosynthesis and Respiration (AIPH). He serves
of the 2004 International Photosynthesis Congress as the Series Editor of AIPH, and as the "Historical
(Montreal, Canada). Govindjee's research has Corner" Editor of Photosynthesis Research. For fur
focused on the function of "Photosystem II" ther information, see his web page at <http://www.
(water-plastoquinone oxidoreductase), particularly on life.uiuc.edu/govindjee>.
