Ozone and Plant Cell
Ozone and Plant Cell by Victoria V. Roshchina and Valentina D. Roshchina Russian Academy of Sciences, Institute of Cell Biophysics, Russia SPRINGER-SCIENCE+BUSINESS MEDIA, B.V.
A C.I.P. Catalogue record for this book is available from the Library of Congress. ISBN 978-90-481-6340-3 ISBN 978-94-017-2523-1 (ebook) DOI 10.1007/ 978-94-017-2523-1 Printed on acid-free paper All Rights Reserved 2003 Springer 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.
CONTENTS PREFACE ix ACKNOWLEDGEMENTS x INTRODUCTION 1 Chapter 1. ATMOSPHERIC OZONE 7 1.1. PHYSICO-CHEMICAL PROPERTIES OF OZONE 7 1.2. FORMATION OF OZONE 10 1.3. DECOMPOSITION OF OZONE 15 1.3.1. Reaction of ozone with oxides of nitrogen and halogen atoms 15 1.3.2. Reaction of ozone with organic components in the atmosphere 16 1.4. CHEMICAL REACTIVITY OF OZONE IN WATER 18 1.5. PROPERTIES OF THE ACTIVE OXYGEN SPECIES FORMED IN WATER AS A RESULT OF OZONOLYSIS 20 1.5.1. Superoxide anion radical 21 1.5.2. Hydroxyl radical 22 1.5.3. Hydrogen peroxide 23 1.5.4. Singlet oxygen 24 1.6. DOSES OF OZONE REQUIRED TO INDUCE BIOLOGICAL EFFECTS IN LIVING ORGANISMS 24 CONCLUSION 26 Chapter 2.TRANSPORT OF OZONE IN PLANT CELLS AND CELLULAR REACTIONS 27 2.1. MOVEMENT OF OZONE FROM SURFACE INTO PLANT TISSUE 27 2.1.1. Cells of leaf and stem 28 2.1.1.1. Pathways through stomata and cuticle 31 2.1.1.2. Pathways through cell wall and across extracellular space 34 2.1.2. Secretory cells 37 2.1.3. Cells of microspores 39 2.2. TRANSPORT OF OZONE THROUGH CELLULAR MEMBRANES 41 2.2.1. Pathways through plasmalemma to cytoplasm and cellular organelles 42 2.2.2. Plasmalemma 44 2.2.3. Cytoplasmic matrix and cellular organelles 47 CONCLUSION 54 Chapter 3. MOLECULAR MECHANISMS OF THE OZONE INTERACTION WITH PLANT CELLULAR COMPONENTS 55 3.1. OZONOLYSIS OF BIOLOGICALLY IMPORTANT CELLULAR COMPOUNDS 55 3.1.1. Interaction of ozone with compounds in cell cover 57
vi 3.1.2. Interaction of ozone with membrane lipids 61 3.1.3. Reaction of ozone with amino acids and proteins 65 3.1.4. Reactions of ozone with amines 68 3.1.5. Interaction of ozone with nucleic acids 68 3.1.6. Reaction of ozone with NADH and NADPH 69 3.1.7. Reaction of ozone with plant regulators of growth and development 70 3.2. FREE RADICALS AS INTERMEDIATES IN THE PROCESS OF OZONOLYSIS 73 3.2.1. Free radicals in biological systems 73 3.2.2. Formation of free radicals during ozonation and their biological effects 78 3.2.3. Mechanisms by which free radicals act on cellular components 84 3.3. PEROXIDES AND HYDROPEROXIDES AS INTERMEDIATES IN THE PROCESS OF OZONOLYSIS 87 3.3.1. Hydrogen peroxide 88 3.3.2. Organic peroxides and hydroperoxides in atmospheric air 89 3.3.3. Formation of peroxides and hydroperoxides in reactions between ozone and the gas excretions of plants 90 3.3.4. Organic peroxides in plant leaves 92 3.3.5. Biological effects of peroxides 93 3.3.6. Mechanisms of the peroxide action CONCLUSION 98 102 Chapter 4. PROTECTIVE CELLULAR REACTIONS AGAINST OZONE AND STEADY-STATE PROTECTIVE SYSTEMS 103 4.1 ANTIOXIDANT ENZYMES 103 4.1.1. Superoxide dismutase (SOD) in plants 103 4.1.2. Catalase 107 4.1.3. Peroxidase 107 4.2. LOW-MOLECULAR ANTIOXIDANTS AND INHIBITORS OF REACTIVE OXYGEN SPECIES 110 4.2.1. Phenols 110 4.2.2. Tocoferol and ubiquinones 112 4.2.3. Ascorbic acid 113 4.2.4. Thiol (SH)-compounds 114 4.2.5. Uric acid, cytokinins and biogenic amines 115 4.2.6. Multiatomic alcohols, polyamines and polyacetylenes 115 4.2.7. Terpenoids 116 4.2.8. Carbon dioxide 117 4.3. PLANT EXCRETIONS AS NATURAL ANTIOZONANTS 117 4.4. INTERACTION OF ANTIOXIDANTS AND THEIR DUAL ROLE IN PLANT LIFE 124 CONCLUSION 126
Chapter 5. OZONE-INDUCED CHANGES IN PLANT METABOLISM 127 5.1.CARBON METABOLISM 127 5.1.1. Photosynthesis 127 5.1.2. Respiration 131 5.1.3. Changes in the pool of metabolites and in the activity of some enzymes. 132 5.2. LIPID METABOLISM 135 5.3. METABOLISM OF NITROGEN-CONTAINING COMPOUNDS 139 5.4. METABOLISM OF SECONDARY COMPOUNDS 144 5.4.1. Phenols 144 5.4.2. Hydrocarbons 149 5.4.3. Polyamines 151 CONCLUSION 152 Chapter 6. CELLULAR MONITORING OF OZONE 153 6.1. SENSITIVITY OF PLANT CELLS TO OZONE 153 6.1.1. Dependence of plant sensitivity on the ozone dose 154 6.1.1.1. Dose-dependent sensitivity of plant cellular reactions 156 6.1.1.2. Dose-dependent sensitivity of whole plant growth and development 164 6.1.1.3. Dose-dependent sensitivity of plant species and cultivars to ozone 168 6.1.2. Dependence of the plant ozone sensitivity on external and internal factors 169 6.2. DIAGNOSTICS OF PLANT CELL SENSITIVITY TO OZONE 173 6.2.1.Diagnostics involving cellular reactions 174 6.2.2. Early diagnostics using biochemical and physiological criteria 178 6.2.2.1. Stomata opening and gas exchange of leaves. 180 6.2.2.2. Permeability of membranes and changes in membrane-related processes 181 6.2.2.3. Formation of stress ethylene 181 6.2.2.4. Changes in photosynthesis and chlorophyll content 182 6.2.2.5. Autofluorescence of cell surfaces 184 6.2.2.6. Changes in the activity and biosynthesis of enzymes. Peroxidases as indicators of ozone stress 186 6.2.2.7. Changes in metabolic pools 187 6.2.2.8. Activation and expression of the genes encoding protective low-molecular and high-molecular metabolites 189 6.2.3. Diagnostics of cell damage 189 vii
viii 6.3. PLANT CELLS AS BIOSENSORS AND BIOINDICATORS OF OZONE 195 6.3.1. Plants as bioindicators of ozone pollution 196 6.3.2. Cellular models for studying ozone effects 200 6.3.2.1. Pollen 200 6.3.2.2. Vegetative microspores of spore-breeding plants 204 6.3.2.3. Secretory cells 206 CONCLUSION 208 CONCLUSION 209 Appendix 1. ADDRESSES OF INTERNET SITES RELATED TO OZONE ISSUES 213 Appendix 2. LIST OF PLANT SPECIES SENSITIVE TO OZONE 214 Appendix 3. AUTOFLUORESCENCE OF PLANT CELLS AS BIOINDICATORS FOR OZONE 215 REFERENCES 217 SUBJECT INDEX 253 LATIN INDEX 265
ix PREFACE The problem of the influence that ozone can cause in living organisms have now become blatantly apparent due to the depletion of the Earth s ozone layer and the increase of such anthropogenic sources of the gas as motor vehicle exhaust and modern devices, including computers, that emit ultraviolet radiation. Plants play significant roles as both indicators and origins of oxygen and ozone in Nature. Therefore, the interaction between plants and ozone is important for plant biophysics, physiology, biochemistry and ecology. Since the issue now needs to be considered on the cellular and molecular levels, the aim of this book is to present a real picture of the functioning of living plant cells upon ozone to a wide group of specialists and naturalists. In the first chapter, the properties of ozone and its transformations in both the atmosphere and water are described. The second chapter is devoted to the ways in which ozone penetrates into living plant cells, the subsequent reactions with cell components and the role of cellular cover and membranes in these processes. Information about such ozone effects as membrane damage, changes in cell permeability and membrane malfunctions is also provided. Molecular mechanisms characteristic for the interaction of ozone with cell components are described in chapter 3. To facilitate an understanding of these mechanisms, modern conceptions about free radicals, peroxides and protective reactions of cells against ozone are also considered. The fourth chapter is devoted to steady-state protectory systems that counteract ozone and other reactive oxygen species. Adaptive changes in cellular metabolism to O 3 exposure are considered in chapter 5. The information in chapter 6 focuses on the cellular monitoring of ozone, and deals with such issues as cellular sensitivity to this substance and the possibility of devising plant cellular models for use as ozone biosensors and indicators. Possible responses to ozone indicated by the various plant reactions are also described. In the conclusion, the importance and possible mechanisms of ozone action when large and small doses display diametrically opposite effects on cells are considered. This monograph will be useful for ecologists, plant physiologists, biochemists, biophysists and botanists, including lecturers and students, as well as for specialists in the field of environmental management.
x ACKNOWLEDGMENTS. The authors are extremely grateful to two corresponding members of the Russian Academy of Sciences, Professor Alexander Kuzin and Professor Arkadii Budantsev, for their useful comments and careful editing of the manuscript. In addition, we wish to thank Dr. Valerii N.Karnaukhov, Head of the Laboratory of Microspectral Analysis and Cellular Monitoring of Environment and Vice-Director of the Institute of Cell Biophysics at the Russian Academy of Sciences, for his constant support, his willing assistance with the work and his eagerness to provide any necessary discussion. We also thank Ludmila I. Mit kovskaya, Larisa F. Kun'eva and Andrei V.Rodionov for their help in producing the computer version of the manuscript.