APPLICATIONS OF CHLOROPHYLL FLUORESCENCE Applications of Chlorophyll Fluorescene in Photosynthesis Research, Stress Physiology, Hydrobiology and Remote Sensing An introduction to the various fields of applications of the in vivo chlorophyll fluorescence also including the proceedings of the first International Chlorophyll Fluorescence Symposium held in the Physikzentrum, Bad Honnej, F.R.C., 6-8 June 1988 edited by HARTMUT K. LICHTENTHALER Botanisches Institut (Plant Physiology and Biochemistry), University of Karlsruhe, Karlsruhe, F.R.G. KLUWER ACADEMIC PUBLISHERS DORDRECHT I BOSTON I LONDON Library of Congress Cataloging in Publication Data International Chiorophyll Fluorescence Symposium (1st: 1988 : Bad Honnef, Germany) App! icatlons of chlorophyl I fluorescence in photosynthesis research. stress physiology, hydrobiology, and remote sensing! ed,ted by Hartmut K. Lichtenthaler. p. cm. "An Introduction to the various fIelds of applications of the in VIVO chlorophyl I fluorescence ... proceedings of the First InternGtlOnal Chloroph,,11 Fluorescence SympoSium, held In the' chysikzentrum. Bad HGnnef. F.R.G .• 6-8 June 1988." Inc 1 Jdes ~ r;dexes. i. Ch'orophyll--Ana'~sis--Congresses. 2. Fluorimetry--Congresses. I. Llchter~t~31er. Hart~ut K. II. ~itle. Or<898. C5:57 1988 53" . 19' 218--d c 19 88-25223 CIP IBSN-13: 978-94-010-7771-2 e-ISBN-13: 978-94-009-2823-7 DOl: 10.1007/978-94-009-2823-7 Published by Kluwer Academic Publishers, P.O. Box 17, 3300 AA Dordrecht, The Netherlands. Kluwer Academic Publishers incorporates the publishing programmes of D. Reidel, Martinus Nijhoff, Dr W. Junk and MTP Press. Sold and distributed in the U.S.A. and Canada by Kluwer Academic Publishers, 101 Philip Drive, Norwell, MA 02061, U.S.A. In all other countries, sold and distributed by Kluwer Academic Publishers Group, P.O. Box 322, 3300 AH Dordrecht, The Netherlands. All Rights Reserved © 1988 by Kluwer Academic Publishers Soltcover reprint of the hardcover I st edition 1988 No part of the material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical including photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner. Sponsor and organizer of the first International Chlorophyll Fluorescence Symposium, Bad Honnef, June 1988. Dr. Volker Schafer Prof. Dr. Hartmut Lichtenthaler WE-Heraeus Foundation University of Karlsruhe Hanau (F.R.G.) The chairmen of the International Chlorophyll Fluorescence Symposium held at Bad Honnef from June 6 to 8, 1988: From left to right: R.J. Strasser, Geneve; H.R. Bolhar-Nordenkampf, Vienna; D. Walker, Sheffield; A. Melis, Berkeley; Z. Sestak, Praha; G. Bertolini, Ispra; W. Schmidt, Konstanz; H.K. Lichtenthaler, Karlsruhe (Scientific Coordinator); G. Guyot, Montfavet; D. Ernst, Hannover; V. Schafer, Hanau (Heraeus Foundation). The participants of the first International Chlorophyll Fluorescence Symposium in front of the Physikzentrum, Bad Honnef: G. Bertolini, R. Blaich, H.R. Bolhar-Nordenkampf, C. Buschmann, D. Ernst, G. Guyot, J. Harbinson, A.R. Holzwarth, L. Kocsanyi, G.H. Krause, H. Laasch, E. Lehoczki, H.K. Lichtenthaler, H. Maske, A. Melis, M. Methy, E. Nagel , E. Ogren, U. Rinderle, A. Rosema, V. Schafer, W. Schmidt, G. Schmuck, H. Schneckenburger, U. Schreiber, Z. Sestak, U. Sieber, D. Siefermann-Harms, P. Siffel, J. Snel, S. Somersalo, R.J. Strasser, Z. Szigeti, M. Tevini, A.F. Theisen, P.R. van Hasselt, O. van Kooten, D. Walker, E. Weis. PREFACE Thi s book is a general introduction into in vivo chlorophyll fluorescence and contains the contributions of the first International Ch 1o rophyll Fluorescence Sympos i urn he 1d in the Phys i kzentrum Bad Honnef, F.R.G. from June 6 to 8, 1988. This Symposium was made possible by a generous support from the Wilhelm and Else Heraeus Foundation, Hanau, which is gratefully acknmvledged. The book not only comprises all aspects of the applications of chlorophyll fluorescence in photosynthesis, stress physiol ogy, hydrobiology and remote sensing, but also gives access to measuring techni ques, data acqui si ti on and earl i er 1i terature references. Thus it is far more than just a common proceedings book, it is a general introduction to all forms of application of the non-destructive in vivo chlorophyll fluorescence including the newest results. In a first chapter the inverse correlation between in vivo chlorophyll fl uorescence and photosynthet i c quantum convers i on and CO? -ass i mi 1a t ion is outlined, the origin and life-time of the chlorophyll fluOrescence at room and liquid nitrogen temperatures are given as well as the induction kinetics (Kautsky effect) and the methodo 1o gi ca 1 approaches to regi ster different forms of chlorophyll-fluorescence signatures. In chapter 2 the applications of chlorophyll fluorescence in stress physiology and stress detection of plants are pointed out, starting with a general stress concept of plants and various examples of how short-term and long-term stress effects (e.g. herbicides, water stress, mineral deficiency, air pollutants) as well as regeneration of the plants' vitality and photo synthesis when the stressor is removed, can be detected and quantified. Chapter 3 deals with the application of chlorophyll fluorescence and de 1a yed fl uorescence (1 umi nescence) in hydrobi 01 ogy, 1i mno logy and oceano graphy including remote sensing of phytoplankton. Chapter 4 contains the basic approach and theoretical considerations for a future remote sensing and stress detection of terrestrial vegetation via laser-induced chlorophyll fluorescence and also gives examples and models as to how reflectance and chlorophyll fluorescence can interfere with and com plement each other. In vivo chlorophyll fluorescence has hitherto found broad application in photosynthesis research, stress physiology, hydrobiology and limnology as well as in remote sensing of phytoplankton in oceanography. It will possibly also be applied in the remote sensing and stress detection of terrestrial vegetat ion, whi ch is the research topi c of severa 1 European 1a boratori es which cooperate within the LASFLEUR programme (LAser-induced FLuorescence in EURope) . x It is hoped that the Proceedings book of the first International Chlorophyll Fluorescence Symposium "Applications of Chlorophyll Fluorescence", which medi ates between di fferent research fi e 1d s, wi 11 be a useful starting help for colleagues and graduate students who are not yet in the field as well as for the active researcher in planning new experiments. The book may help not only to avoid major mistakes and measuring artefacts but also to interpret the measured chlorophyll fluorescence signatures in the right way and find access to the application of newer techniques. I wish to thank Dr. Volker Schafer, W. & E. Heraeus Foundation for good cooperation and Ir. A.C. Plaizier, Kluwer Academic Publishers, Dordrecht for publishing the proceedings book, my colleagues for providing their newest resu lts in time, my coworkers and graduate students of the Botani sches Institut Karlsruhe for assistance and last but not least my wife Regine for her valuable aid in editing the book. u~ k'.iJik.~ Karlsruhe, June 1988 Hartmut K. Lichtenthaler Scientific Coordinator of the Chlorophyll Fluorescence Symposium TABLE OF CONTENTS CHAPTER 1. Applications in photosynthesis research G.H. Krause and E. Weis. The photosynthetic apparatus and chloro- phyll fluorescence: an introduction .......................•.... 3 O. Walker. Some aspects of the relationship between chlorophyll a fluorescence and photosynthetic carbon assimilation ............ 13 A.R. Holzwarth. Time-resolved chlorophyll fluorescence. What kind of information on photosynthetic systems does it provide?...... 21 A. Melis, G.E. Guenther, P.J. Morrissey and M.L. Ghirardi. Photo- system II heterogeneity in chloroplasts ........................ 33 O. Siefermann-Harms. Fluorescence properties of isolated chlorophyll-protein complexes .................................. 45 v v , L.P. Siffel and Z. Sestak. Low temperature fluorescence spectra of chloroplasts: methodical aspects and possible applications...... 55 U. Schreiber, C. Neubauer and C. Klughammer. New ways of assessing photosynthetic activity with a pulse modulation fl uorometer ................................................... 63 E. Weis. Steady state photosynthesis in intact plants as analyzed by chlorophyll fluorescence and far-red spectroscopy......... 71 C. Buschmann and H.K. Lichtenthaler. Complete fluorescence emission spectra determined during the induction kinetic using a diode- array detector ................................................. 77 Z. Sestak and P. Siffel. Changes in chloroplast fluorescence during 1e af deve 1o pment ............................................... 85 J. Snel. Measurement and analysis of chlorophyll fluorescence in plants using modulated actinic illumination .................... 93 L. Kocsanyi, M. Haitz and H.K. Lichtenthaler. Measurement of the laser-induced chlorophyll fluorescence kinetics using a fast acoustooptic device............................................ 99 Z. Szigeti, E. Palos, E. Lehoczki. Fluorescence properties of paraquat resistant Conyza leaves................................ 109 E. Lehoczki and Z. Szigeti. Characterization of paraquat resistant Conyza leaves through delayed fluorescence .................... 115 P. van Hasselt. Temperature dependence of in vivo chlorophyll fluorescence as effected by DCMU .................•............ 121 CHAPTER 2. Applications in stress physiology and environmental research H.K. Lichtenthaler. In vivo chlorophyll fluorescence as a tool for stress detection in plants .................................... 129 H.K. Lichtenthaler and U. Rinderle. Chlorophyll fluorescence signatures as vitality indicator in forest decline research..... 143 U. Schreiber, W. Bilger, C. Klughammer and C. Neubauer. Application of the PAM fluorometer in stress detection ..................... 151 S. Somersalo and G.H. Krause. Changes in chlorophyll fluorescence related to photoinhibition of photosynthesis and cold acclimation of green plants .................................... 157 E. Ogren and G. Oquist. Screening for photoinhibition of photosyn- thesis in the field using a portable fluorimeter ............... 165 H.R. Bolhar-Nordenkampf and E. Lechner. Winter stress and chloro- phyll fluorescence in Norway spruce .........•................. 173 R.J. Strasser, B. Schwarz and P.Eggenberg. Fluorescence routine tests t< to describe the behaviour of a plant in its environment......... 181 U. Rinderle and H.K. Lichtenthaler. The chlorophyll fluorescence ratio F690/F735 as a possible stress indicator ................. 189 A.F. Theisen. Fluorescence changes in a drying maple leaf observed in the visible and near-infrared ............................... 197 O. van Kooten, B. van Hove and K.J. van Wijk. The effect of long term exposition of poplars to low concentrations of S02 and NH3 ............................................................ 203 W. Schmidt. Luminescence of organic molecules, Theory and analytical applications in photosynthesis ................................. 211 W. Schmidt. Long term delayed luminescence in green organisms 217 R. Blaich. Early detection of damage conditions in plants by delayed chlorophyll fluorescence ............................... 223 M. Tevini, P. Grusemann and G. Fieser. Assessment of UV-B stress by chlorophyll fluorescence analysis ........................... 229 E. Nagel and H.K. Lichtenthaler. Comparison of photoacoustic and chlorophyll fluorescence signatures of green leaves ............ 239 M. Haitz and H.K. Lichtenthaler. The measurement of Rfd-values as plant vitality indices with the portable field fluorometer and the PAM-fl uorometer ........................................ 249
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