K. Omasa, H. Saji, S. YOllssefian, N. Kondo (Eds.)

Air Pollution and Plant Biotechnology Prospects for Phytomonitoring and Phytoremediation

With 103 Figures

~ Springer

Kenji Omasa, Ph.D. Professor,. The University of Tokyo Yahoi 1-1-1, Bunkyo-ku, Tokyo 113-8657, Japan

Hiharu Saji, Ph.D. Head of Molecular Ecotoxicology Section Environmental Biology Division National Institute for Environmental Studies Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Shohab Youssefian, Ph.D. Associate Professor, Akita Prefectural University Minami 2-2, Ohgata-mura, Akita 010-0444, Japan

Noriaki Kondo, Ph.D. Professor, The University of Tokyo Hongo 7-3-1, Bunkyo-ku, Tokyo 113-0033, Japan

ISBN -13: 978-4-431-68390-2 e-ISBN-13 :978-4-431-68388-9 DOl: 10.1007/978-4-431-68388-9

This work is subject to copyright. All rights are reserved whether the whole or part of the materials is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks.

The use of registered names, trademarks etc., in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.

Omasa, Saji, Youssefian and Kondo: Air Pollution and Plant Biotechnology

© 2002 Springer-Verlag Tokyo

All rights reserved. No part of this publication may be reproduced, stored in any electronic or mechanical form, including photocopy, recording or otherwise, without the prior written permission of the publisher.

First Indian Reprint 2005 Second Indian Reprint 2008

ISBN -13: 978-4-431-68390-2

This edition is manufactured in India for sale only in India, Pakistan, Bangladesh, Nepal and Sri Lanka and any other country as authorized by the publisher.

This edition is published by Springer (India) Private Limited, A part of Springer Science+Business Media, Registered Office: 906-907, Akash Deep Building, Barakhamba Road, New Delhi -110 001, India.

Foreword

Organisms have encountered numerous environmental changes since their first appearance on earth some 4 billion years ago. Such changes generally occurred gradually at slow rates, but sometimes they occurred transiently, violently, and at rapid rates. In most cases, changes in environmental factors were induced by geological variations or by cosmic events, such as bombardment by giant meteorites. Pollutants produced by human activities would be classified under rapid change in the global environment. Organisms themselves also brought about changes in the environment, as typically observed with the increasing concentrations of atmospheric dioxygen produced by oxygenic photosynthetic organisms that originated from cyanobacteria. When faced with such environmental changes, organisms that possessed systems enabling them to tolerate the new conditions or factor could survive, whereas organisms lacking these tolerant systems either disappeared from. the new biosphere or escaped into the limited biosphere of the previous environment. Only those organisms that were tolerant to the new environmental factor could subsequently develop the appropriate mechanisms to utilize this factor and to occupy a dominant position in the new biosphere.

A typical environmental factor, to which organisms have shown an evolutionary acquisition of tolerance and which they have been able to effe-ctively utilize, is atmospheric dioxygen. Prior to the appearance of cyanobacteria, the amount of dioxygen in the atmosphere, produced by UV-photolysis of water, was minute, with estimated concentrations of 0.002%, around 10.4 that of present atmospheric levels. Even these extremely low dioxygen concentrations appear to have been lethal to anaerobes, the sole organisms 3 billion years ago~ Many anaerobes, including anaerobic photosynthetic bacteria, contain superoxide dismutase, an essential enzyme that scavenges superoxide radicals and protects against dioxygen damage. Hence, prior to the accumulation of dioxygen in the atmosphere resulting

V

VI Foreword

from the activity of oxygenic photosynthetic organisms, some anaerobes must already have possessed systems that enabled them to tolerate the dioxygen-induced damage that resulted mainly from oxidative breakdown of target molecules by reactive species of oxygen. Clearly, therefore, systems that scavenge such reactive molecules are absolutely indispensable for the survival of organisms in atmospheres with even extremely low dioxygen concentrations.

Cyanobacteria, the first biological donors of dioxygen, are thought to have arisen from the fusion of anaerobic purple bacteria and green sulfur photosynthetic bacteria, through which the two photochemical reaction centers became associated and the additional water oxidation system in the thylakoids was acquired. Fusion with superoxide dismutase-containing photosynthetic bacteria presumably provided the cyanobacteria with protection against the dioxygen that they themselves produced. However, this dioxygen from cyanobacteria was a dangerous pollutant gas, as are present pollutants, for neighboring anaerobes, and caused lethal oxidative damage to cells that had limited potential to scavenge reactive species of oxygen. Some such anaerobes were able to escape to anaerobic environments, which are still present today. On the other hand, those anaerobes that acquired the scavenging systems of reactive species of oxygen were able to survive even in the atmosphere containing dioxygen at higher levels, produced by prokaryotic algae and then by the eukaryotic algae and terrestrial plants. The oxygen-tolerant organisms thus evolved and subsequently acquired the aerobic respiratory systems that enabled them to produce 19-fold higher amounts of bioenergy, in the form of adenosine 5'-triphosphate (ATP), than that generated by fermentation.

The atmospheric oxygen was, at first, a toxic poliutant to anaerobes, but it finally came to be used as an effective electron acceptor for respiration and as a substrate for the biosynthesis of various essential metabolites via oxidase and oxygenase reactions. However, dioxygen is still toxic to all organisms including aerobes, and no organisms on earth survive without effective systems and mechanisms of scavenging the reactive species of oxygen, of rapidly repairing and synthesizing de novo the oxidized target molecules, and of fine-tuning the protecting systems into an orchestrated response to oxidative stress.

Among all organisms, plants are exposed to the severest environments with respect to oxidative damage. Cellular concentrations of dioxygen in leaf tissues (more than 2.5 x 10.4 M) are the highest of all organisms; for comparison, the concentration in the vicinity of mitochondria in mammalian hepatic cells is 10.7 to 10.8 M. Furthermore, plants are always exposed to strong sunlight, with a maximal 2xlO·3mol m·2 S·l in the visible range, which induces photosensitized oxidations. To maintain photosynthetic activity, plants are equipped with the most effective mechanisms of photooxidative stress protection. Even so, photosynthesis is appreciably photoinhibited in nature either by defects in the protection system or by overproduction of reactive species of oxygen generated by other environmental stresses. Atmospheric pollutants may also enhance oxidative damage, especially under sunlight, indicating that the reactive species of oxygen participate in such

Foreword VII

pollutant damage, as also presented in this monograph. The effect of environmental stresses, such as excess light, UV, drought, salt, temperature, and CO2 on photosynthesis is generally to stimulate the photoproduction of reactive species of oxygen, by which the pollutants synergistically accelerate photodamage.

No one could possibly doubt the importance of maintaining plant photosynthesis levels for protection of the global environment. Annual CO2

emissions by fossil fuel combustion and cement production (6.4 x 109 tons as carbon) comprise only 6% of the annual global fixation of CO2 by photosynthetic organisms (1.1 x 1011 tons, in net production as carbon). Thus, even a minor decrease in the rate of the global CO2 fixation, as affected by pollutants and other environmental stresses, would greatly accelerate the increase in atmospheric CO2

and result in further amplification of the numerous stress factors that affect photosynthesis. Conversely, even a minor increase in the rate of CO2 fixation could negate the increase in atmospheric CO2 resulting from fossil fuel combustion.

The molecular breeding of pollutant-tolerant plants, as described in this monograph, is indispensable for the sustainable protection of the global environment. To achieve this objective, the mechanisms of tolerance, not only to pollutants but also to environmental stresses need to be considered in the light of breeding strategies. An in-depth understanding of the cross talk between environmental and pollutant-induced stresses would most certainly allow the breeding of plants suited to specific environments. Furthermore, the incorporation of the capacity of organisms living in extreme environmental conditions into plants would also be an effective strategy of developing novel tolerant plants. In these respects I believe that the present monograph, Air Pollution and Plant Biotechnology, will serve as a milestone in mankind's future progress in identifying ways of maintaining and improving our global environments.

Kozi Asada Department of Biotechnology Faculty of Life Engineering Fukuyama University Fukuyama, Hiroshima, Japan

Preface

Air pollution is ubiquitous in industrialized and densely populated districts. It is not only toxic per se to organisms, but is also responsible for a wide range of environmental crises, including acidic deposition, stratospheric ozone-layer destruction, and global warming. It is therefore critical that we both monitor and reduce the levels of atmospheric pollutants to preserve our natural environment.

Plants can serve as useful tools for such purposes, and some species have already been exploited as detectors (for phytomonitoring) or as scavengers (for phytoremetiiation) of air pollutants. However, accelerating progress in biotechnology, especially in the fields of plant tissue culture and genetic manipulation, now makes it possible to improve various plant characteristics rapidly so as to extend their utility in these areas. An essential prerequisite to the development of such novel plants is a systematic accumulation of up-to-date infoI:IIlation on the physiological and biochemical responses of plants, on the regulation of plant gene expression, and on the effects and mechanisms of pollutant metabolism in plant cells.

To address the need for such information, we present in this volume current topics that deal with various aspects of phytpmonitoring and phytoremediation. The. book is thus divided into four main sections: section I concerns plant responses and phytomonitoring; section II deals with resistant plants and phytoremediation; section III examines systems for imaging diagnosis of plaJ;\t responses and gas exchange; and section IV focuses on the generation and use of novel transgenic plants. We review the basic physiological and biochemical properties of plants as a necessary background to several of these topics, especially where attempts have been made to apply modern methods of biotechnology to air pollution control. We also evaluate current concepts and techniques for the early detection of plant stress and for the screening of tissues and plants with characteristics relevant to phytoremediation and phytomonitoring. This book will

IX

x Preface

therefore be of considerable value to researchers and students who are interested in these new technologies and who are considering areas in which to utilize their knowledge of and expand their skills in plant biotechnology.

Kenji Omasa Hikaru Saji Shohab Youssefian Noriaki Kondo

Contents

Foreword ...................................................... V .

Preface······················································· IX

Contributors ................................................ XIX

I. Plant Responses and Phytomonitoring

1. Responses of Whole Plants to Air Pollutants' ....................... 3

Isamu Nouchi

1. Introduction··················································· 3 2. Sulfur Oxides ................................................. 4

3. Ozone······················································· 8 4. Peroxyacetyl Nitrate (PAN) ..................................... 15 5. Nitrogen Oxides ........•..................................... 18

6. Fluoride····················································· 21 7. Acid Rain" .......................... , ...................... 22

8. Combination of Air Pollutants ................................... 26

2. Plants as Bioindicators of Air Pollutants' ......................... 41

Isamu Nouchi

XI

XII Contents

1. Introduction·················································· 41 2. Bioindicators for Sulfur Dioxide' ................................. 42

3. Bioindicators for Hydrogen Fluoride .............................. 42 4. Bioindicators for Ethene (Ethylene) ............................... 43 5. Bioindicators for Ozone ........................................ 43

6. Bioindicators for Peroxyacetyl Nitrate (PAN) ....................... 53 7. Conclusion .................................................. 56

3. Phytomonitoring for Urban Environmental Management· ........... 61

Margaret Burchett, Rachid Mousine, and Jane Tarran

1. Introduction···················,······························ 61 2. Scope of Project .............................................. 64

3. Methodology: Ecoepidemiological ................................ 67 4. Findings: Ecoepidemiological .................................... 69 5. Field Experiment· ............................................. 83

6. Implications for Environmental Management· ....................... 86

4. Effects of Air Pollutants on Lipid Metabolism in Plants ............. 93

Takeshi Sakaki

1. Introduction·················································· 93 2. Leaf Glycerolipids and Their Metabolism .......................... 94 3. Lipid Oxidation by Air Pollutants' ................................ 95

4. Metabolic Alteration of Lipids by Air Pollutants ..................... 97 5. Conclusions and Prospects for Biotechnology ...................... 104

5. Effects of Ethylene on Plant Responses to Air Pollutants' . . . . . . . . . . . 111 Nobuyoshi Nakajima

1. Introduction················································· 111 2. Ozone-Induced Ethylene Synthesis' .............................. 112

3. Effects of Ethylene Under Acute Ozone Exposure' .................. 114

4. Effects of Ethylene Under Chronic Ozone Exposure' ................ 116

Conlents XIII

5. Sulfur Dioxide-Induced Ethylene Production' ...................... 116 6. Conclusion ................................................. 117

6. Effects of Air Pollutants on Gene Expression in Plants' ............ 121

Akihiro Kubo

1. Introduction················································· 121 2. Effects of Air Pollutants on Gene Expression' .................... " 122

3. Biological Significance of Gene Expression in Response to Air Pollutants ............................................. 130

4. Application of the Detection of Gene Expression to Environmental Biotechnology . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 133

5. Conclusions················································· 134

7. Biotechnology for Phytomonitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 141

Hikaru Saji

1. Introduction················································· 141 2. Use of Biotechnology to Generate Plants with Altered Sensitivity

to Air Pollutants ............................................. 142

3. Molecular Sensors of Air Pollutants' ............................. 148

4. Conclusions················································· 149

n. Resistant Plants and Phytoremediation

8. Absorption of Organic and Inorganic Air Pollutants by Plants' . . . . .. 155 Kenji Omasa, Kazuo Tobe, and Takayuki Kondo

1. Introduction················································· 155

2. A Simple Gas Diffusion Model for Analyzing Gas Absorption by Plant Leaves' . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 156

3. Analysis of Foliar Absorption of Pollutant Gases by the Gas Diffusion Model" ............................. ,. " ., ................ 159

XIV Contents

4. Stomatal Control of Gas Absorption and Susceptibility of Plants to Air Pollutants· ........................ ~ ......................... 165

5. Conclusion················································· 173

9. Uptake, Metabolism, and Detoxification of Sulfur Dioxide .. . . . . . . . . 179

Noriaki Kondo

1. Introduction················································· 179 2. Absorption of Sulfur Dioxide ................................... 180

3. Phytotoxicity of Sulfur Dioxide ................................. 183

4. Metabolism of Sulfur Dioxide' .................................. 186 5. Conclusion .•.............•...........................•.•... 191

10. Elevated Levels of Hydrogen Sulfide in the Plant Environment: Nutrient or Toxin .....................................•..... 201

Luit J. De Kok, C. Elisabeth E. Stuiver, Sue Westerman,

and Ineke Stulen

1. Introduction················································· 201 2. Elevated H2S and Plant Growth ................................. 202

3. Uptake and Metabolism of H2S .................................. 204

4. Atmospheric H2S, Sulfur Nutrition, and Sulfur Assimilation' .......... 208 5. H2S Metabolism Versus Toxicity ................................ 212 6. Concluding Remarks' ......................................... 212

11. Metabolism and Detoxification of Nitrogen Dioxide and Ammonia in Plants' ...........•...................................... 221

Tadakatsu Yoneyama, Hak Y. Kim, Hiromichi Morikawa, and

Hari S. Srivastava

1. Introduction················································· 221 2. Absorption and Metabolism of N02 •••••••••••••••••••••••••••••• 222

3. Toxicity and Detoxification of N02 •••••••••••••••••••••••••••••• 227

4. Absorption and Metabolism of NH3 .............................. 229

5. Conclusions and Perspectives' .................................. 230

Contents xv

12. Plant Resistance to Ozone: the Role of Ascorbate ................. 235

Jeremy Barnes, Youbin Zheng, and Tom Lyons

1. Introduction················································· 235 2. Genetic Basis of Ozone Resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236

3. Factors Governing Ozone Resistance ............................. 240

4. Conclusions················································· 246

13. Detoxification of Active Oxygen Species and Tolerance in Plants Exposed to Air Pollutants and CO2 •••••••••••••••••••••.••••••• 253

Shigeto Morita and Kunisuke Tanaka

1. Introduction················································· 25~ 2. Response of Antioxidative Systems to Ozone' ...................... 25:

3. Response of Antioxidative Systems and Tolerance to S02 ............. 261 4. Response of Antioxidative Systems to CO2 •••••••••••••••••••••••• 26:

14. Countermeasures with Fertilization to Reduce Oxidant-Induced Injury to Plants' ............................. 26

Haruko Kuno and Kazushi Arai

1. Introduction················································· 26'

2. Effects of Nutritional Components on Ozone-Induced Visible Injury .... 271

3. Countermeasures for Reducing Damage Caused by Oxidants to Spinach: Methods for Fertilizer Application ............................... 27:

III. Image Diagnosis of Plant Response and Gas Exchange

15. Image Instrumentation of Chlorophyll a Fluorescence for Diagnosing Photosynthetic Injury ........................... 287

Kenji Omasa and Kotaro Takayama

1. Introduction················································· 287 2. Chlorophyll a Fluorescence ................................ , ... 288

XVI Contents

3. Image Instrumentation System ............•..................... 293

4. Diagnosis of Environmental Stresses .......•..................... 296

5. Conclusions················································· 305

16. Field-Portable Imaging System for Measurement of Chlorophyll Fluorescence Quenching' ........................ 309

Barry Osmond and Yong-Mok Park

1. Introduction················································· 309 2. A Prototype (Almost) Portable System' ........................... 310 3. Some Applications and Results' ................................. 312 4. Further Developments' ........................................ 315

17. Assessment of Environmenta! Plant Stresses Using Multispectral Steady-State Fluorescence Imagery····· ............ '" ......... 321

Moon S. Kim, Charles L. Mulchi, James E. McMurtrey,

Craig S. T. Daughtry, and Emmett W. Chappelle

1. Introduction················································· 321 2. Steady-State Fluorescence Characteristics of Vegetation ...... " ...... 322 3. Multispectral Steady-State Fluorescence Techniques' ................ 326 4. Effects of Moderately Elevated 0 3 and CO2 •••••••••••••••••••••••• 332

5. Effects of Varying Content of Flavonols .......................... 336 6. Concluding Remarks' ..................... , ................... 338

18. Diagnosis of Stomatal Response and Gas Exchange of Trees by Thermal Remote Sensing .................................. 343

Kenji Omasa

1. Introduction················································· 343 2. Information Obtained from Leaf Temperature ...................... 344

3. Image Instrumentation of Leaf Temperature' ....................... 348

4. Diagnosis of Trees by Leaf Temperature Image' .................... 351 5. Conclusion ................................................. 356

Contents XVII

IV. Generation of Transgenic Plants

19. Manipulation of Genes Involved in Sulfur and Glutathione Metabolism ................................................ 363

Shohab Youssefian

1. Introduction················································· 363

2. Molecular Regulation of Sulfur Assimilation and Glutathione BiosyntheJis ................................................ 365

3. Manipulation of Genes and Production of Transgenic Plants' .......... 369 4. Concluding Remarks' ......................................... 377

20. Manipulation of Genes for Nitrogen Metabolism in Plants' ......... 383

Hiromichi Morikawa, Misa Takahashi, and Gen-Ichiro Arimura

1. Introduction················································· 383 2. Genetic Manipulation of NR Genes .............................. 385 3. Genetic Manipulation of NiR Genes' ............................. 386 4. Genetic Manipulation of GS Genes" ............................. 388

5. Nitrogen Dioxide Assimilation in Transgenic Plants Containing Chimeric NiR cDNA, and GS1 and GS2 cDNA" ................... 391

21. Manipulation of Genes for Antioxidative Enzymes ................ 403

Mitsuko Aono

1. Introduction······;·········································· 403 2. Manipulation of Genes for Superoxide Dismutases .......... > •• > • > > • 405

3. Manipulation of Genes for Glutathione Reductase' .................. 407

4. Manipulation of Genes for Other Antioxidative Enzymes ............. 408

5. Manipulation of more than One Gene Encoding Antioxidative Enzymes ................................................... 409

6. Conclusions and Perspectives' .................................. 411

XVIII Contents

22. Application of Genetic Engineering for Forest Tree Species' ........ 415

Saori Endo, Etsuko Matsunaga, Keiko Yamada-Watanabe,

and Hiroyasu Ebinuma

1. Introduction················································· 415 2. Genetic Improvement of Forest Tree Species' ...................... 416

3. Transformation Studies in Forest Tree Species' ..................... 417

4. Transformation Research of Populus Species as a Model System for Forest Tree Species ........................................ 420

5. Improving Stress Tolerance in Hybrid Aspen by Genetic Engineering ... 423

6. Prospects of Improving Traits of Forest Tree Species Through Genetic Engineering .......................................... 430

23. Environmelltal Risk Assessment of Transgenic Plants: A Case Study of Cucumber Mosaic Virus-Resistant Melon in Japan' ....... 435

Yutaka Tabei

1. Introduction to the Risk Assessment of Genetically Modified Organisms .................................................. 435

2. Guidelines for Risk Assessment of Transgenic Crops in Japan ......... 437

3. Environmental Risk Assessment of Cucumber Mosaic Virus-Resistant Transgenic Melon ............................................ 438

4. Conclusion ................................................. 444

Index························································ 449

Contributors

Aono, Mitsuko, Environmental Biology Division, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Arai, Kazushi, Environmental Ecology Division, Tokyo Metropolitan Forestry Experiment Station, Hirai 2753-1, Hinode, Nishitama-gun, Tokyo 190-0182, Japan

Arimura, Gen-Ichiro, Department of Mathematical and Life Sciences, Graduate School of Science, Hiroshima University, Kagamiyama 1-3-1, Higashi­Hiroshima, Hiroshima 739-8526, Japan

Barnes, Jeremy, Department of Agricultural and Environmental Science, Ridley Building, Newcastle University, Newcastle upon Tyne, NEI 7RU, UK

Burchett, Margaret, Centre for Ecotoxicology, Faculty of Science, University of Technology, Sydney (UTS), Westbourne St, Gore Hill, NSW 2065, Australia

Chappelle, Emmett W., Biospheric Sciences Branch, Laboratory for Terrestrial Physics, NASAlGSFC, Greenbelt, MD 20771, USA

Daughtry, Craig S. T., Hydrology and Remote Sensing Laboratory, USDA Agricultural Research Service, Beltsville, MD 20705, USA

De Kok, Luit J., Laboratory of Plant Physiology, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands

Ebinuma, Hiroyasu, Pulp and Paper Research Laboratory, Nippon Paper Industries Co., LTD, Oji 5-21-1, Kita-ku, Tokyo 114-0002, Japan

Endo, Saori, Pulp and Paper Research Laboratory, Nippon Paper Industries Co., LTD, Oji 5-21-1, Kita-ku, Tokyo 114-0002, Japan

XIX

xx Contributors

Kim, Hak Y., Institute of Agricultural Science and Technology, Kyungpook National University, 1370 Sankyuk-dong, Puk-ku, Taegu 702-701, Korea

Kim, Moon S., Instrumentation and Sensing Laboratory, USDA Agricultural Research Service, Beltsville, MD 20705, USA

Kondo, Noriaki, Department of Biological Sciences, Graduate School of Science, The University of Tokyo, Hongo 7-3-1, Bunkyo-ku, Tokyo 113-0033, Japan

Kondo, Takayuki, Air Quality Section, Toyama Prefectural Environmental Science Research Center, Nakataikouyama 17-1, Kosugi, Toyama 939-0363, Japan

Kubo, Akihiro, Environmental Biology Division, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Kuno, Haruko, Environmental Ecology Division, Tokyo Metropolitan Forestry Experiment Station, Hirai 2753-1, Hinode, Nishitama-gun, Tokyo 190-0182, Japan

Lyons, Tom, Department of Agricultural and Environmental Science, Ridley Building, Newcastle University, Newcastle upon Tyne, NE1 7RU, UK

Matsunaga, Etsuko, Pulp and Paper Research Laboratory, Nippon Paper Industries Co., LTD, Oji 5-21-1, Kita-ku, Tokyo 114-0002, Japan

McMurtrey, James E., Hydrology and Remote Sensing Laboratory, USDA Agricultural Research Service, Beltsville, MD 20705, USA

Morikawa, Hiromichi, Department of Mathematical and Life Sciences, Graduate School of Science, Hiroshima University, Kagamiyama 1-3-1, Higashi­Hiroshima, Hiroshima 739-8526, Japan

Morita, Shigeto, Faculty of Agriculture, Kyoto Prefectural University, Shimogamo-Hangicho 1-5, Sakyo-ku, Kyoto 606-8522, Japan

Mousine, Rachid, Centre for Ecotoxicology, Faculty of Science, University of Technology, Sydney (UTS), Westbourne St, Gore Hill, NSW 2065, Australia

Mulchi, Charles L., Department of Natural Resource Sciences, University of Maryland, College Park, MD 20742, USA

Nakajima, Nobuyoshi, Biodiversity Conservation Research Project, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Contributors XXI

Nouchi, Isamu, Agro-Meteorology Group, National Institute for Agro­Environmental Sciences, Kannondai 3-1-3, Tsukuba, Ibaraki 305-8604, Japan

Omasa, Keoji, Department of Biological and Environmental Engineering, Graduate School of Agricultural and Life Sciences, The University of Tokyo, Yayoi 1-1-1, Bunkyo-ku, Tokyo 113-8657, Japan

Osmond, Barry, Biosphere 2 Center, Columbia University, 32540 S Biosphere Road, PO Box 689, Oracle AZ 85623, USA

Park, Yong-Mok, Department of Life Science, College of Natural Science and Engineering, Chongju University, Chongju, 360"764, Korea

Saji, Hikaru, Environmental Biology Division, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Sakaki, Takeshi, Department of Bioscience and Technology, School of Engineering, Hokkaido Tokai University, Minami-sawa 5-1-1-1, Minami-ku, Sapporo 005-8601, Japan

Srivastava, Hari S., Department of Plant Science, Rbhilkhand University, Bareilly 243006, India

Stuiver, C. Elisabeth E., Laboratory of Plant Physiology, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands

Stulen, Ineke, Laboratory of Plant Physiology, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands

Tabei, Yutaka, Plant Biotechnology Department, National Institute of Agrobiological Sciences, Kan-nondai 2-1-2, Tsukuba, Ibaraki 305-8602, Japan

Takahashi, Misa, Department of Mathematical and Life Sciences, Graduate School of Science, Hiroshima University, Kagamiyama 1-3-1, Higashi­Hiroshima, Hiroshima 739-8526, Japan

Takayama, Kotaro, Department of Biological and Environmental Engineering, Graduate School of Agricultural and Life Sciences, The University of Tokyo, Yayoi 1-1-1, Bunkyo-ku, Tokyo 113-8657, Japan

Tanaka, Kunisuke, Faculty of Agriculture, Kyoto Prefectural University, Shimogamo-Hangicho 1-5, Sakyo-ku, Kyoto 606-8522, Japan

XXII Contributors

Tarran, Jane, Centre for Ecotoxicology, Faculty of Science, University of Technology, Sydney (UTS), Westboume St, Gore Hill, NSW 2065, Australia

Tobe, Kazuo, Laboratory of Intellectual Fundamentals for Environmental Studies, National Institute for Environmental Studies, Onogawa 16-2, Tsukuba, Ibaraki 305-8506, Japan

Westel1Dan, Sue, Laboratory of Plant Physiology, University of Groningen, P.O. Box 14, 9750 AA Haren, The Netherlands

Yamada.Watanabe, Keiko, Pulp and Paper Research Laboratory, Nippon Paper Industries Co., LTD, Oji 5-21-1, Kita-ku, Tokyo 114-0002, Japan

Yoneyama, Tadakatsu, Department of Applied Biological Chemistry, Graduate School of Agricultural and Life Sciences, The University of Tokyo, Yayoi I-I­I, Bunkyo-ku, Tokyo 113-8657, Japan

Youssefian, Shohab, Biotechnology Institute, Faculty of Bioresource Sciences, Akita Prefectural University, Minami 2-2, Ohgata-mura, Akita 010-0444, Japan

Zheng, Y oubin, Department of Plant Agriculture, Bovey Building, Gordon Street, University of Guelph, Guelph, Ontario, Canada