CONTRIBUTORS TO THIS VOLUME H. E. BARRS H. G. JONES B. CALLANDER P. E. KRIEDEMANN W. R. CHANEY J. J. LANDSBERG D. DOLEY S. G. PALLARDY F. DUHME H. RICHTER T. M. HINCKLEY G. R. SQUIRE P. R. JARVIS R. Ο. TESKEY D. WHITEHEAD WATER DEFICITS AND PLANT GROWTH EDITED BY T. T. KOZLOWSKI DEPARTMENT OF FORESTRY UNIVERSITY OF WISCONSIN MADISON, WISCONSIN VOLUME VI Woody Plant Communities 1981 ACADEMIC PRESS A Subsidiary of Harcourt Brace Jovanovich, Publishers New York London Toronto Sydney San Francisco COPYRIGHT © 1981, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED. NO PART OF THIS PUBLICATION MAY BE REPRODUCED OR TRANSMITTED IN ANY FORM OR BY ANY MEANS, ELECTRONIC OR MECHANICAL, INCLUDING PHOTOCOPY, RECORDING, OR ANY INFORMATION STORAGE AND RETRIEVAL SYSTEM, WITHOUT PERMISSION IN WRITING FROM THE PUBLISHER. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data Main entry under title: Water deficits and plant growth. Includes bibliographies and indexes. Contents: v. 1. Development, control, and measurement—v. 2. Plant water consumption and response—[etc.]—v. 6. Woody plant communties. 1. Plants—Water requirements. 2. Growth (Plants). 3. Plant-water relationships. A. Plant diseases. I. Kozlowski, T. T. (Theodore Thomas), Date. QK870.W38 582'.013 68-14658 ISBN 0-12-424156-5 (v. 6) AACR2 PRINTED IN THE UNITED STATES OF AMERICA 81 82 83 84 9 8 7 6 5 4 3 2 1 LIST OF CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors' contributions begin. H. D. BARRS (325), Division of Irrigation Research, C.S.I.R.O., Griffith, New South Wales, Australia B. CALLANDER (471), Department of Physiology and Environmental Stud- ies, School of Agriculture, University of Nottingham, Sutton Bonington, Loughborough, Leicestershire, England WILLIAM R. CHANEY (1), Department of Forestry and Natural Re- sources, Purdue University, West Lafayette, Indiana 47907 D. DOLEY (209), Botany Department, University of Queensland, St. Lucia, Queensland, Australia F. DUHME (153), Lehrstuhl für Landschaftsökologie, Technische Univer- sität München, D-8050 Freising-Weihenstephan, Federal Republic of Germany T. M. HINCKLEY* (153), School of Forestry, Fisheries and Wildlife, Uni- versity of Missouri, Columbia, Missouri 65211 P. G. JARVIS (49), Department of Forestry and Natural Resources, Uni- versity of Edinburgh, Edinburgh EH9 3JU, Scotland H. G. JONES (419), East Mailing Research Station, East Mailing, Maidstone, Kent, England P. E. KRIEDEMANN (325), Division of Irrigation Research, C.S.I.R.O., Griffith, New South Wales, Australia J. J. LANDSBERG (419), Long Ashton Research Station, Long Ashton, Bristol, England * Present Address: College of Forest Resources, University of Washington, Seattle, Washington 98195 ix χ List of Contributors STEPHEN G. PALLARDY (511), School of Forestry, Fisheries and Wild- life, University of Missouri, Columbia, Missouri 65211 H. RICHTER (153). Botanisches Institut, Universität für Bodenkultur, Vienna, Austria G. R. SQUIRE (471), Department of Physiology and Environmental Stud- ies, School of Agriculture, University of Nottingham, Sutton Bonington, Loughborough, Leicestershire, England R. O. TESKEY* (153), School of Forestry, Fisheries and Wildlife, Univer- sity of Missouri, Columbia, Missouri 65211 D. WHITEHEAD (49), Forest Research Institute, New Zealand Forest Ser- vice, Private Bag, Rotorua, New Zealand * Present Address: College of Forest Resources, University of Washington, Seattle, Washington 98195 PREFACE The emphasis in this volume is on the water relations of woody plants in a community context. The ecosystems discussed include forest trees as well as trees grown for fruit crops and leaf crops. The opening chapter presents a quantitative overview of sources of water available to woody plants. Separate chapters follow on water rela- tions of coniferous forests, temperate hardwood forests, tropical and sub- tropical forests and woodlands, closely related woody plants, apple or- chards, citrus orchards, and tea plantations. For each of the plant communities discussed, emphasis is given to hydrological cycles; water use and transpiration; absorption of water; effects of environmental factors on soil and plant water balance; effects of water deficits on physiological processes; vegetative and reproductive growth; yield of harvested products; drought resistance; and cultural practices affecting plant water balance and yield. The subject matter is sufficiently varied so as to make this volume useful to both researchers and those involved in the practice of growing woody plants for wood and fruit crops and for their esthetic values. In this volume water deficits are characterized in bars or megapascals (Mpa) depending on preferences of individual authors (1 bar =0.1 Mpa). The contributors to this volume were chosen for their research pro- ductivity and extensive experience with the particular plant communities discussed. I wish to express my sincere thanks to each author for his scholarly contribution as well as cooperation during the production phases. T. T. KOZLOWSKI xi CONTENTS OF OTHER VOLUMES VOLUME I. DEVELOPMENT, CONTROL, AND MEASUREMENT 1. Introduction T. T. KOZLOWSKI 2. Water Structure and Water in the Plant Body A. S. CRAFTS 3. Terminology in Plant and Soil Water Relations S. A. TAYLOR 4. Evaporation of Water from Plants and Soil C. B. TANNER 5. Availability and Measurement of Soil Water W. R. GARDNER 6. Plant Factors Influencing the Water Status of Plant Tissues I. R. COWAN and F. L. MILTHORPE 7. Drought-Resistance Mechanisms JOHNSON PARKER 8. Determination of Water Deficits in Plant Tissues H. D. BARRS AUTHOR INDEX—SUBJECT INDEX VOLUME II. PLANT WATER CONSUMPTION AND RESPONSE 1. Water Consumption by Agricultural Plants Μ. Ε. JENSEN 2. Water Consumption by Forests A. J. RUTTER 3. Water Deficits and Physiological Processes A. S. CRAFTS 4. Water Deficits and Growth of Herbaceous Plants C. T. GATES 5. Water Deficits and Growth of Trees R. ZAHNER xiii xiv Contents of Other Volumes 6. Water Deficits in Vascular Disease P. W. TALBOYS AUTHOR INDEX—SUBJECT INDEX VOLUME III. PLANT RESPONSES AND CONTROL OF WATER BALANCE 1. Shrinking and Swelling of Plant Tissues Τ. Τ. KOZLOWSKI 2. Soil Moisture and Seed Germination D. HlLLEL 3. Water Deficits and Reproductive Growth MERRILL R. KAUFMANN 4. Protoplasmic Resistance to Water Deficits JOHNSON PARKER 5. Water Deficits and Enzymatic Activity GLENN W. TODD 6. Water Deficits and Nutrient Availability FRANK G. VIETS, JR. 7. Water Deficits and Nitrogen Metabolism AUBREY W. TAYLOR 8. Water Deficits and Hormone Relations AVINOAM LIVNE AND YOASH VAADIA 9. Physiological Basis and Practical Problems of Reducing Transpiration ALEXANDRA POLJAKOFF-MAYBER AND J. GALE 10. Soil Water Conservation D. HlLLEL AND E. RAWITZ AUTHOR INDEX—SUBJECT INDEX VOLUME IV. SOIL WATER MEASUREMENT, PLANT RESPONSES, AND BREEDING FOR DROUGHT RESISTANCE 1. Measurement of Water Content and the State of Water in Soils S. L. RAWLINS 2. Structure and Functioning of Stomata W. G. ALLA WAY AND F. L. MILTHORPE 3. Stomatal Conductance in the Control of Gas Exchange F. J. BURROWS AND F. L. MILTHORPE Contents of Other Volumes xv 4. Water Deficits and Photosynthesis J. S. BOYER 5. Water Supply and Leaf Shedding T. T. KOZLOWSKI 6. Water Deficits and Flow of Latex B. R. BUTTERY AND S. G. BOATMAN 7. Water Deficits and Nitrogen-Fixing Root Nodules JANET I. SPRENT 8. Plant Breeding for Drought Resistance E. A. HURD AUTHOR INDEX—SUBJECT INDEX VOLUME V. WATER AND PLANT DISEASE 1. Water Relations of Diseased Plants PETER G. AYRES 2. Water Stress as a Predisposing Factor in Plant Disease DONALD F. SCHOENEWEISS 3. Abiotic Diseases Induced by Unfavorable Water Relations R. D. DURBIN 4. Water and Spore Liberation C. T. INGOLD 5. Water and the Infection Process C. E. YARWOOD 6. Effect of Soil Moisture on Survival and Spread of Pathogens D. M. GRIFFIN 7. Moisture and Seed Decay C. M. CHRISTENSEN 8. Moisture and Deterioration of Wood D. W. FRENCH AND C. M. CHRISTENSEN 9. Moisture as a Factor in Epidemiology and Forecasting A. F. VAN DER WAL AUTHOR INDEX—SUBJECT INDEX CHAPTER 1 SOURCES OF WATER William R. Chaney DEPARTMENT OF FORESTRY AND NATURAL RESOURCES, PURDUE UNIVERSITY, WEST LAFAYETTE, INDIANA I. Introduction 1 World Water Resources 1 II. Sources of Moisture for Woody Plants 4 A. Atmospheric Moisture 4 B. Soil Water 19 C. Internal Water Reservoirs 32 References 40 I. INTRODUCTION Of the myriad of environmental factors influencing distribution and growth of woody plants, water is of paramount importance and is usually the most limiting throughout the world. This situation exists even though global supplies of water are immense. However, the interaction of air circulation patterns, topography, temperature, and edaphic factors result in uneven distribution and availability of this moisture. Consequently, the forms of moisture utilized by trees, and the adaptations necessary for its utilization, are quite diverse. This chapter will consider the use and importance to woody plants of moisture that occurs in the atmosphere, the soil, and internal tissues. The physical, anatomical, and physiological factors that affect the availability and absorption of the various forms of moisture will be emphasized. WORLD WATER RESOURCES Of the world's total water supply, only an extremely small part is available to woody plants during any one year. Approximately 93% of the earth's 165 trillion acre-feet of water is in the oceans and seas. The re- 1 Water Deficits and Plant Growth, Vol. VI Copyright © 1981 by Academic Press, Inc. All rights of reproduction in any form reserved. ISBN 0-12-424156-5