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Chlorophyll a Fluorescence Advances in Photosynthesis and Respiration Chlorophyll a Fluorescence Advances in Photosynthesis and Respiration VOLUME 19 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, Frankfur t am Main, Germany Kirmiyuki SATOH, Okayama, Japan James Siedow, Durham, Nor th 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. Chlorophyll a Fluorescence A Signature of Photosynthesis Edited by George C. Papageorgiou National Center for Scientific Research Demokritos, Athens, Greece and Govindjee University of Illinois, Urbana, Illinois, U.S.A. 4y Springer A C.I.P. Catalogue record for this book is available from the Library of Congress. ISBN 1-4020-3217-X (HB) ISBN 1-4020-3218-8 (e-book) Published by Springer, P.O. Box 17, 3300 AA Dordrecht, The Netherlands. Sold and distributed in North, Central and South America by Springer, 101 Philip Drive, Norwell, MA 02061, U.S.A. In all other countries, sold and distributed by Springer, P.O. Box 322, 3300 AH Dordrecht, The Netherlands. 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. Legend for solar photon flux densities and pigments spectra to 1000 nm Solar photon flux densities at the top of the atmosphere and at the Earth’s surface, with estimated in vivo absorbance spectra of selected photosynthetic pigments from plants, algae and cyanobacteria, and fluorescence spectrum of chlorophyll a. The solar spectra were truncated beyond 1000nm, and the pigment absorption spectra below 400 nm. Sources: Top-of-the-atmosphere irradiance: 150-200 nm, Andrew Lacis, NASA Goddard Institute for Space Studies (GISS); 200-400 nm, Judith Lean (Naval Research Laboratory); 400-2500 nm, Brian Cairns, NASA GISS. Surface irradiance: 200-400 nm, J. Lean (Lean and Rind, 1998); 400-2500 nm, B. Cairns. Atmospheric gases, David Crisp (personal communication, NASA Jet Propulsion Laboratory). Chlorophyll a and Chlorophyll b absorbance measurements, made by Junzhong Li (H. Du and coworkers, 1998), in vitro, were shifted in wavelengths to match in vivo peaks, and absorbances were normalized to between 0 and 1. Carotenoid absorption spectra are estimated in vivo absorption spectra in green algae (Govindjee, 1960). Phycoerythrin and phycocyanin absorption spectra are unpublished absorption spectra from Govindjee’ s laboratory, and from Ke (2001). Chlorophyll a fluorescence spectrum, from spinach chloroplasts, is from Govindjee and Yang (1966). The color bar is from Dan Burton (Color Science web page: http://www.physics.sfasu.edu/astro/color.html). See chapter 1 by Govindjee for references cited above. The figure for the cover is copyrighted and prepared by Nancy Kiang (NASA Goddard Institute for Space Studies) and Govindjee (University of Illinois at Urbana-Champaign). Printed on acid-free paper All Rights Reserved © 2004 Springer 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. Printed in the Netherlands. This book is dedicated to L. N. M. Duysens, a pioneer of Photosynthesis During his scientific life, Louis (Lou) Nico Marie ideas of Lou Duysens. Fundamental concepts that fol- Duysens quietly contributed some of the most lowed from these experiments include the carotenoid seminal ideas in photosynthesis, with still central to (bacterio-)chlorophyll energy transfer in plant influence over our thinking today. His approach was and bacterial light harvesting systems, the efficient a perfect blend of deep theoretical understanding energy transfer among a large number of quasi-iden- and experimental innovation. Typical of Duysens' tical chlorophylls to reach the small population of early work was his insightful application of Forster reaction centers, the Take model' of photosynthetic theory to light harvesting by the accessory pigment unit function, the control of the fluorescence yield beds, which firmly established the physics underly- of Photosystem II by the redox state of the quencher ing the function of the photosynthetic unit. Using his Q (now known as QA), and chlorophyll excitation own state-of-the-art absorption spectrophotometer, quenching by carotenoids. These concepts underlie he discovered the spectroscopic signature of oxida- the veritable industry of fluorescence-based, non- tion and reduction of the photochemical reaction invasive methods now used to study photosynthesis center in bacteria. Later, adapting this technique to in vivo and to analyze plant productivity, from the plant (algal) systems, he discovered the alternating lab bench to satellite monitoring. effects of green and red light on the redox state of Lou's understanding of physical principles is im- cytochrome fin red algae, to firmly establish the now mense and so facile that his colleagues and competi- familiar model of Photosystem I and Photosystem tors struggled to keep up with him. His seemingly II acting in series. These major contributions exem- simple analysis of the thermodynamic limits of pho- plified Lou Duysens' philosophy towards technical tosynthetic energy conversion sparked a literature developments. Technology never drove nor limited that lasted several decades, with each incremental his research, but was harnessed to address specific advance simply rediscovering what he had said at scientific questions and hypotheses. the outset. Indeed, this is still an area that few fully With his easy understanding of photochemical comprehend. and photophysical principles, Duysens pioneered Since Lou's retirement, many ultrafast spectro- the use of Chlorophyll fluorescence as a powerful scopic techniques have emerged, revealing intricate probe of photosynthetic function, in all classes of details of function in the photosynthetic apparatus, photosynthetic organisms, to discover many of the which can be well understood in terms of the fan- essential events of primary energy conversion in tastic atomic resolution structures of photosynthetic photosynthesis: excitation energy transfer in the pigment-protein structures that are now available. In light-harvesting antenna and charge separation in the this new era of understanding, it is amazing to see reaction center. Over many years, in his laboratory in how the concepts, developed by Duysens even 50 Leiden (The Netherlands), a multitude of state-of-the years ago, have survived as solid foundations of our art fluorescence methods was created, driven by the current models of photosynthetic activity. v Editorial Advances in Photosynthesis and Respiration Volume 19: Chlorophyll a Fluorescence: A Signature of Photosynthesis I am delighted to announce the publication, in the 10. Photosynthesis: Photobiochemistry and Photo- Advances and Photosynthesis and Respiration biophysics (Bacon Ke, author, 2001); (AIPH) Series, the first book that focuses on the red 11. Regulation ofPhotosynthesis (Eva-Mari Aro and light that plants, algae and cyanobacteria emit when Bertil Andersson, editors, 2001); exposed to UV and visible light. This new volume, 12. Photosynthetic
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