Environmental physiology of plants 3rd ed a fitter (AP, 2002)

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 YO10 5YW Robert Hay Scottish Agricultural Science Agency 80 Craigs 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 ISBN 0-12-257766-3 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 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, and and RKMH for Chapters 4, and 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, encouragement 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 Acknowledgements v vii Introduction Plant growth and development The influence of the environment Evolution of adaptation Comparative ecology and phylogeny 1 14 17 Part I The Acquisition of Resources 21 Energy and Carbon Introduction The radiation environment Radiation Irradiance Temporal variation Leaf canopies Effects of spectral distribution of radiation on plants Perception Germination Morphogenesis Placement Flowering Effects of irradiance on plants Responses to low irradiance Photosynthesis at high irradiance Responses to elevated carbon dioxide concentrations Photosynthetic responses Whole-plant responses 23 23 26 26 26 29 29 33 33 34 36 38 42 43 43 57 66 66 70 Mineral Nutrients Introduction Nutrients in the soil system Soil diversity Concentrations Ion exchange 74 74 82 82 85 87 This Page Intentionally Left Blank Plate Four individual Pinus sylvestris from the Caledonian Pinewood, Loch Maree, Scotland, showing the diversity of form generated by the interactions of genotype and environment (a) (b) (c) Plate (a) Large sunfleck under the canopy of a Sequoia sempervirens forest in California (b) and (c) Temperate deciduous forest in lowland England before and after the expansion of the leaf canopy (a) (b) Plate (a) Aerial view of FACE (Free Air CO2 Enrichment) rings in a Liquidambar styraciflua forest at Oak Ridge, USA (b) Closer view of the FACE rings showing the vents from which CO2 is released Photographs courtesy of Dr R Norby (a) (b) (c) (d) Plate The Park Grass experiment at Rothamsted Experimental Station, UK (a) unfertilised plot, showing the diversity of the vegetation; (b) plot receiving high rates of ammonium fertiliser (twice that in (c), right side) and no lime, now very acid with complete dominance by the grass Anthoxanthum odoratum; (c) boundary between productive, but still diverse plot that receives complete nutrients with N as nitrate (left), and unproductive, acid plot that receives N as ammonium (right); the right-hand plot is dominated by Agrostis Neither plot is limed; (d) plot receiving high phosphate but no nitrogen, dominated by red clover Trifolium pratense (b) (c) (a) (d) Plate Experiments and techniques for studying roots (a) and (b) Mini-rhizotron equipment for capturing images of roots in soil allows continuous and long-term recording of root growth under all conditions (c) Typical image of roots captured by a mini-rhizotron (note the prominent root hairs); image represents ca mm diameter view (d) Design used by Campbell et al (1991) to study root foraging, showing algal growth on the nutrient-rich quadrants Photographs courtesy of Dr G Self (b), Dr Angela Hodge (c) and Professor Philip Grime (d) (a) Plate (b) Chemical changes in soils induced by roots (a) (right hand panel) pH changes around the roots of a maize plant grown with either nitrate or ammonium as the sole nitrogen source, demonstrating strong acidification with ammonium and a marked increase in rhizosphere pH when nitrate is used; (left hand panel) Similar changes observed when sorghum and chickpea take up nitrate from soil The chickpea acidifies the soil because it is N-fixing and therefore metabolizes ammonium rather than utilizing soil nitrate (b) Oxidation of the rhizosphere by the roots of Phragmites australis in solution culture, as revealed by oxidation of methylene blue (and in soil by the precipitation of iron oxide on the root surface) Photographs courtesy of (a) Dr V R×omheld, and (b) Dr J Armstrong and Mr D Holt (a) (b) Plate The rhizosphere SEMs of a barley root in soil, showing (a) soil particles adhering to the root surface, and prominent fungal hyphae; (b) root hairs bridging the gap between the root surface and a soil particle Photographs courtesy of Dr Colin Campbell (c) (a) (d) (b) Plate Roots and symbioses (a) Cluster roots of Grevillia robusta; (b) nodules formed by the nitrogen-fixing bacterium Rhizobium on the roots of Robinia pseudoacacia; (c) cross-section of an ectomycorrhizal root of beech Fagus sylvatica; (d) intercellular hyphae and arbuscules of an arbuscular mycorrhizal fungus in a root of bluebell, Hyacinthoides non-scripta Photographs courtesy of (a) Dr Keith Skene, (b) Dr Karyn Ridgway, (c) Dr Angela Hodge, (d) Dr James Merryweather (a) (c) (b) (d) Plate Habitats in which water management is important (a) The cactus Brachycereus nesioticus on lava, Isla Fernandina, Galapagos, dependent on water storage; (b) desert in New Mexico after rain, showing the ephemeral and deciduous flora flowering, with surrounding succulents; (c) and (d) giant trees which must raise water to >70 m: c, karri Eucalyptus diversicolor in the relatively arid south-west of Australia; d, Douglas fir Pseudotsuga menziesii in the temperate moist forest of Vancouver Island, Canada (a) (b) (c) (d) Plate 10 Plants of Death Valley, California (a—c) Larrea tridentata ssp divaricata: (a) aerial view showing evenly-spaced individuals; (b) single shrub; (c) flowering; (d) Tidestromia oblongifolia Photographs 10(b—d) courtesy of Dr Stanley Smith Plate 11 Desiccation resistance Hydrated and air-dry states of the resurrection plant Craterostigma wilmsii Photograph courtesy of Professor Jill Tarrant (a) (b) Plate 12 Regeneration of Australian plants after fire (a) Eucalyptus shoots developing from epicormic buds on the woody stem; (b) vigorous growth of understorey vegetation after fire (note the fire-blackened trunks) (b) (a) (c) Plate 13 Treeline (a) Natural treeline in the Canadian Rockies at ca 2500 m; (b) Norway Spruce (Picea abies) near the treeline after heavy snowfall, Southern Norway; (c) a tree at the treeline on Mount Fuji, Japan, showing the lower skirt that develops under the snowpack, and the flagging effect caused by windblown ice particles (b) (c) (a) (d) Plate 14 Alpine plant growth forms (a) Giant rosettes of Espeletia lopezii in the Sierra Nevada de Cocuy, Colombia; (b) Oxyria digyna flowering at 600 m in the West Highlands of Scotland; (c) and (d) Silene acaulis and Crepis nana at ca 3000 m in the Canadian Rockies, demonstrating cushion growth form Photograph 14(a) courtesy of Dr Jason Rauscher ... enhanced far-red component of leaf-transmitted shade and the enhanced short-wave content (mainly blue) of diffuse radiation Temporal variation Temporal variation in irradiance is a universal feature... response of mustard (Sinapis alba) plants grown in front of a fence‘ of green grass plants (C) or a similar canopy bleached by treatment with the weedkiller paraquat (A) (from BallarÑe et al., 1987)... expressed as the leaf area index (LAI or L), a dimensionless parameter representing the area of leaf surface over unit area of ground It is possible to relate the rate of attenuation of solar radiation