Ozone and Plant Cell. Victoria V. Roshchina. Valentina D. Roshchina SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. and

Transcription

1 Ozone and Plant Cell

2 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.

3 A C.I.P. Catalogue record for this book is available from the Library of Congress. ISBN ISBN (ebook) DOI / 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.

4 CONTENTS PREFACE ix ACKNOWLEDGEMENTS x INTRODUCTION 1 Chapter 1. ATMOSPHERIC OZONE PHYSICO-CHEMICAL PROPERTIES OF OZONE FORMATION OF OZONE DECOMPOSITION OF OZONE Reaction of ozone with oxides of nitrogen and halogen atoms Reaction of ozone with organic components in the atmosphere CHEMICAL REACTIVITY OF OZONE IN WATER PROPERTIES OF THE ACTIVE OXYGEN SPECIES FORMED IN WATER AS A RESULT OF OZONOLYSIS Superoxide anion radical Hydroxyl radical Hydrogen peroxide Singlet oxygen 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 MOVEMENT OF OZONE FROM SURFACE INTO PLANT TISSUE Cells of leaf and stem Pathways through stomata and cuticle Pathways through cell wall and across extracellular space Secretory cells Cells of microspores TRANSPORT OF OZONE THROUGH CELLULAR MEMBRANES Pathways through plasmalemma to cytoplasm and cellular organelles Plasmalemma Cytoplasmic matrix and cellular organelles 47 CONCLUSION 54 Chapter 3. MOLECULAR MECHANISMS OF THE OZONE INTERACTION WITH PLANT CELLULAR COMPONENTS OZONOLYSIS OF BIOLOGICALLY IMPORTANT CELLULAR COMPOUNDS Interaction of ozone with compounds in cell cover 57

5 vi Interaction of ozone with membrane lipids Reaction of ozone with amino acids and proteins Reactions of ozone with amines Interaction of ozone with nucleic acids Reaction of ozone with NADH and NADPH Reaction of ozone with plant regulators of growth and development FREE RADICALS AS INTERMEDIATES IN THE PROCESS OF OZONOLYSIS Free radicals in biological systems Formation of free radicals during ozonation and their biological effects Mechanisms by which free radicals act on cellular components PEROXIDES AND HYDROPEROXIDES AS INTERMEDIATES IN THE PROCESS OF OZONOLYSIS Hydrogen peroxide Organic peroxides and hydroperoxides in atmospheric air Formation of peroxides and hydroperoxides in reactions between ozone and the gas excretions of plants Organic peroxides in plant leaves Biological effects of peroxides Mechanisms of the peroxide action CONCLUSION Chapter 4. PROTECTIVE CELLULAR REACTIONS AGAINST OZONE AND STEADY-STATE PROTECTIVE SYSTEMS ANTIOXIDANT ENZYMES Superoxide dismutase (SOD) in plants Catalase Peroxidase LOW-MOLECULAR ANTIOXIDANTS AND INHIBITORS OF REACTIVE OXYGEN SPECIES Phenols Tocoferol and ubiquinones Ascorbic acid Thiol (SH)-compounds Uric acid, cytokinins and biogenic amines Multiatomic alcohols, polyamines and polyacetylenes Terpenoids Carbon dioxide PLANT EXCRETIONS AS NATURAL ANTIOZONANTS INTERACTION OF ANTIOXIDANTS AND THEIR DUAL ROLE IN PLANT LIFE 124 CONCLUSION 126

6 Chapter 5. OZONE-INDUCED CHANGES IN PLANT METABOLISM CARBON METABOLISM Photosynthesis Respiration Changes in the pool of metabolites and in the activity of some enzymes LIPID METABOLISM METABOLISM OF NITROGEN-CONTAINING COMPOUNDS METABOLISM OF SECONDARY COMPOUNDS Phenols Hydrocarbons Polyamines 151 CONCLUSION 152 Chapter 6. CELLULAR MONITORING OF OZONE SENSITIVITY OF PLANT CELLS TO OZONE Dependence of plant sensitivity on the ozone dose Dose-dependent sensitivity of plant cellular reactions Dose-dependent sensitivity of whole plant growth and development Dose-dependent sensitivity of plant species and cultivars to ozone Dependence of the plant ozone sensitivity on external and internal factors DIAGNOSTICS OF PLANT CELL SENSITIVITY TO OZONE Diagnostics involving cellular reactions Early diagnostics using biochemical and physiological criteria Stomata opening and gas exchange of leaves Permeability of membranes and changes in membrane-related processes Formation of stress ethylene Changes in photosynthesis and chlorophyll content Autofluorescence of cell surfaces Changes in the activity and biosynthesis of enzymes. Peroxidases as indicators of ozone stress Changes in metabolic pools Activation and expression of the genes encoding protective low-molecular and high-molecular metabolites Diagnostics of cell damage 189 vii

7 viii 6.3. PLANT CELLS AS BIOSENSORS AND BIOINDICATORS OF OZONE Plants as bioindicators of ozone pollution Cellular models for studying ozone effects Pollen Vegetative microspores of spore-breeding plants 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

8 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.

9 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.