Environmental Physiology of Plants
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Environmental Physiology of Plants Third Edition This Page Intentionally Left Blank Environmental Physiology of Plants Third Edition Alastair Fitter Department of Biology University of York P.O. Box 373 York YO105YW Robert Hay Scottish Agricultural Science Agency 80Craigs Road Edinburgh EH12 8NJ San Diego * San Francisco * New York Boston * London * Sydney * Tokyo This book is printed on acid-free paper. Copyright # 2002 A.H. Fitter and R.K.M. Hay All Rights Reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Academic Press A division of Harcourt Inc. Harcourt Place, 32 Jamestown Road, London NW1 7BY, UK http://www.academicpress.com Academic Press A division of Harcourt Inc. 525 B Street, Suite 1900, San Diego, California 92101-4495, USA http://www.academicpress.com ISBN0-12-257766-3 Library of Congress Catalog Number: 2001090353 A catalogue record for this book is available from the British Library Typeset by Mathematical Composition Setters Ltd, Salisbury, Wiltshire Printed and bound in Great Britain by MPG Books Ltd, Bodmin, Cornwall 02 03 04 05 06 07 MP 9 8 7 6 5 4 3 2 1 Preface to the Third Edition This project began nearly 25 years ago, and the first edition was published in 1981. Since then, plant science and ecology have undergone radical revolutions, but the need to understand the environmental physiology of plants has never been greater. On the one hand, with the excitement generated by molecular approaches, there is a real risk that young plant scientists will lack the necessary understanding of how whole plants function. On the other hand, there are global problems to tackle, most notably the consequences of climate change; those who are charting possible futures for plant communities need to have a good grasp of the underlying physiology. Environmental physiology occupies a vital position as a bridge between the gene and the simulation model. Although this edition retains the basic structure and philosophy of previous editions, the text has been completely rewritten and updated to give a synthesis of modern physiological and ecological thinking. In particular, we explain how new molecular approaches can be harnessed as tools to solve problems in physiology, rather than rewriting the book as a primer of molecular genetics. To balance the molecular aspects, we have made a positive decision to use relevant examples from pioneering and classic work, drawing attention to the foundations of the subject. New features include a more generic approach to toxicity, explicit treatment of issues relating to global climate change, and a section on the role of fire. The text illustrations, presented according to a common and improved format, are complemented by colour plates. Even though the rewriting of the book has been a co-operative enterprise, AHF is primarily responsible for Chapters 2, 3 and 7 and RKMH for Chapters 4, 5 and 6. We thank Terry Mansfield, Lucy Sheppard, Ian Woodward, Owen Atkin and Angela Hodge for their helpful comments on individual chapters in draft. Environmental physiology is a rapidly expanding field, and the extent of the literature is immense. Our aim has not been to be comprehensive and authoritative but to develop principles and stimulate new ideas through selected examples, and we remain committed to a policy of full citation to facilitate access to key publications. Where the subject area is in rapid flux, we have attempted to provide a balanced review, which will inevitably be overtaken by events; and we have consciously focused attention on studies in North America, Europe and Australia, because of our personal experience of these areas, and because we hope that this will give a greater coherence to the vi Preface to the Third Edition examples chosen. Although much excellent work is published in languages other than English, we have not relied heavily on this, since the book is intended primarily for students to whom such literature is relatively inaccessible. Finally, we would recommend the advanced monograph Physiological Plant Ecology, edited by M.C. Press, J.D. Scholes and M.G. Barker @1999; Blackwell Science, Oxford) as a useful complement to this book. This third edition is a celebration of a quarter of a century of working together towards a common goal from different viewpoints and experiences. We are very grateful to our editor, Andy Richford, whose vision, encourage- ment and persistence have kept us to the task. A.H. FITTER R.K.M. HAY Acknowledgements We are grateful for permission from the following authorities to use materials for the figures and tables listed: Academic Press Ltd., London À Figs. 3.23, 4.9, 5.10; Annals of Botany À Fig. 1.1a; American Society of Plant Physiologists À Figs. 4.2, 6.3; Blackwell Science Ltd., Oxford À Figs 2.8, 2.27, 3.7, 4.11, 4.12, 5.6, 5.15, 6.8, 6.9, Tables 4.4, 6.7; BIOS Scientific Publishers Ltd., Oxford À Fig. 4.17; Cambridge University Press À Table 5.2; Ecological Society of America À Fig. 5.13; Elsevier Science, Oxford À Fig. 4.4; European Society for Agronomy À Fig. 4.10; HarperCollins, London À Fig. 2.14; J. Wiley and Sons, New York À Fig. 5.16; Kluwer Academic Publishers, Dordrecht À Fig. 3.2; Munksgaard International Publishers and Dr Y. Gauslaa À Fig. 5.9; National Research Council of Canada À Fig. 5.11; New Phytologist and the appropriate authors À Figs. 2.26, 2.29, 3.26, 3.37, 4.19, 5.1, 5.14, 6.7, 6.10; Oikos À Table 6.4; Oxford University Press À Figs. 2.25, 4.8, 5.2; Pearson Education, Inc., New Jersey À Fig. 4.6; Physiologia Plantarum À Fig. 6.2; Professor I.F. Wardlaw À Fig. 5.5; Professor M.C. Drew À Fig. 6.6; Royal Society of Edinburgh and Dr G.A.F. Hendry À Table 6.5; Royal Society of London and Professor K. Raschke À Fig. 4.15; Springer Verlag @Berlin) and the appropriate authors À Figs. 1.7, 2.18, 5.4, 5.8, 5.12, 5.17, 6.11 Tables 5.5, 6.6; Urban & Fischer Verlag, Jena À Fig. 6.5; Weizmann Science Press of Israel and Professor Y. Gutterman À Table 4.2. To Rosalind and Dorothea Contents Preface to the Third Edition v Acknowledgements vii 1. Introduction 1 1. Plant growth and development 1 2. The influence of the environment 8 3. Evolution of adaptation 14 4. Comparative ecology and phylogeny 17 Part I. The Acquisition of Resources 21 2. Energy and Carbon 23 1. Introduction 23 2. The radiation environment 26 1. Radiation 26 2. Irradiance 26 3. Temporal variation 29 4. Leaf canopies 29 3. Effects of spectral distribution of radiation on plants 33 1. Perception 33 2. Germination 34 3. Morphogenesis 36 4. Placement 38 5. Flowering 42 4. Effects of irradiance on plants 43 1. Responses to low irradiance 43 2. Photosynthesis at high irradiance 57 5. Responses to elevated carbon dioxide concentrations 66 1. Photosynthetic responses 66 2. Whole-plant responses 70 3. Mineral Nutrients 74 1. Introduction 74 2. Nutrients in the soil system 82 1. Soil diversity 82 2. Concentrations 85 3. Ion exchange 87 x Contents 4. Cycles 90 5. Transport 91 6. Limiting steps 97 3. Physiology of ion uptake 98 1. Kinetics 98 2. Interactions 101 3. Regulation 102 4. Morphological responses 106 1. Root fraction 106 2. Root diameter and root hairs 107 3. Root density and distribution 108 4. Turnover 112 5. Soil micro-organisms 115 1. The nature of the rhizosphere 115 2. Nitrogen fixation 118 3. Mycorrhizas 120 6. General patterns of response to soil nutrients 128 4. Water 131 1. Properties of water 131 2. The water relations of plants and soils 134 1. Water potential 134 2. The water relations of plant cells 135 3. Plant water stress 138 4. Supply of water by the soil 143 5. Loss of water from transpiring leaves 148 6. Water movement in whole plants 153 3. Adaptations favouring germination and seedling establishment in dry environments 157 4. Adaptations favouring survival and reproduction under conditions of water shortage 162 1. Acquisition of water 163 2. Conservation and use of water 167 3. Tolerance of desiccation 180 4. Contrasting life histories in arid environments 183 5. Some special problems in tree=water relations 184 1. Vascular system 186 2. Leaves 188 Part II. Responses to Environmental Stress 191 5. Temperature 193 1. The temperature relations of plants 193 1. The thermal environment of plants 193 2. The temperature relations of plant processes 195 3. Plant development 197 4. Plant growth and metabolism 200 5. Responses to changes in the thermal environment 203 Contents xi 2. Plant adaptation and resistance to low temperature 205 1. The influence of low temperature on plants 205 2. Characteristic features of cold climates: Arctic and Alpine environments, temperate winters 207 3. Adaptations favouring plant growth and development in Arctic and Alpine regions 211 4. Adaptations favouring survival of cold winters: dormancy 224 5. Adaptations favouring survival of cold winters: plant resistance to freezing injury 225 6. Life in a warmer world: the case of the Arctic Tundra 229 3. The survival of plants exposed to high temperatures 231 4. Fire 236 1. The influence of fire on plants and communities 236 2. Life histories in the Kwongan: ephemerals, obligate seeders and resprouters 239 6. Toxicity 241 1. The nature of toxicity 241 2.