PHYSIOLOGY OF WOODY PLANTS Third Edition This page intentionally left blank PHYSIOLOGY OF WOODY PLANTS Third Edition DR. STEPHEN G. PALLARDY School of Natural Resources University of Missouri Columbia, Missouri AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald’s Road, London WC1X 8RR, UK This book is printed on acid-free paper. Copyright © 2008, Elsevier 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. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: [email protected]. You may also complete your request online via the Elsevier homepage (http://elsevier.com), by selecting “Support & Contact” then “Copyright and Permission” and then “Obtaining Permissions.” Library of Congress Cataloging-in-Publication Data Pallardy, Stephen G. Physiology of woody plants / Stephen G. Pallardy.—3rd ed. p. cm. Rev. ed. of: Physiology of woody plants / Theodore T. Kozlowski, Stephen G. Pallardy. 2nd ed. c1997. ISBN 978-0-12-088765-1 1. Woody plants—Physiology. 2. Trees—Physiology. I. Kozlowski, T. T. (Theodore Thomas), 1917– Physiology of woody plants. II. Title. QK711.2.K72 2007 571.2—dc22 2007033499 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library. ISBN: 978-0-12-088765-1 For information on all Academic Press publications visit our Web site at www.books.elsevier.com Printed in the United States of America 07 08 09 10 9 8 7 6 5 4 3 2 1 Working together to grow libraries in developing countries www.elsevier.com | www.bookaid.org | www.sabre.org Theodore T. Kozlowski Paul J. Kramer This book is dedicated to Dr. Theodore T. Kozlowski and the late Dr. Paul J. Kramer (1904–1995), who pioneered the fi eld of woody plant physiology. This page intentionally left blank Contents Preface xiii Stems 19 Sapwood and Heartwood 19 CHAPTER Xylem Increments and Annual Rings 20 Earlywood and Latewood 21 1 Phloem Increments 22 Introduction 1 Wood Structure of Gymnosperms 23 Axial Elements 24 Horizontal Elements 25 Heredity and Environmental Regulation Wood Structure of Angiosperms 25 of Growth 1 Axial Elements 26 Physiological Regulation of Growth 2 Horizontal Elements 27 Some Important Physiological Processes Bark 27 and Conditions 3 Roots 28 Complexity of Physiological Processes 3 Adventitious Roots 30 Problems of Foresters, Horticulturists, Root Tips 30 and Arborists 3 Root Hairs 31 Physiology in Relation to Present and Suberized and Unsuberized Roots 32 Future Problems 4 Mycorrhizas 33 Summary 6 Reproductive Structures 35 General References 6 Angiosperms 35 Gymnosperms 36 CHAPTER Summary 37 2 General References 38 The Woody Plant Body 9 CHAPTER Introduction 9 Crown Form 10 3 Variations in Crown Form 10 Vegetative Growth 39 Stem Form 11 Vegetative Organs and Tissues 12 Introduction 39 Leaves 12 Cell and Tissue Growth 40 Angiosperms 13 Dormancy 42 Variations in Size and Structure of Leaves 15 Dormancy Concepts 42 Gymnosperms 17 Hormonal Infl uences on Bud Dormancy 44 vii viii Contents Shoot Growth 45 CHAPTER Bud Characteristics 46 Dormant and Adventitious Buds 46 4 Hormonal Infl uences on Shoot Growth 47 Reproductive Growth 87 Leaf Growth 48 Seasonal Leaf Growth Characteristics 49 Introduction 87 Leaf Area Index 50 Reciprocal Relations between Vegetative and Shoot Growth Types and Patterns 50 Reproductive Growth 88 Determinate and Indeterminate Shoots 50 Sexual Reproduction in Angiosperms 88 Epicormic Shoots 50 Flowering Periodicity 88 Preformed and Neoformed Shoots 51 Pollination 90 Recurrently Flushing Shoots 51 Fruit Set 90 Abnormal Late-Season Shoots 52 Fertilization 91 Apical Dominance 53 Postfertilization Development 91 Maximum Height 54 Polyembryony 92 Shoot Growth in the Tropics 54 Apomixis 92 Cambial Growth 55 Parthenocarpy 92 Cell Division in the Cambium 55 Growth of Fruits 92 Production of Xylem and Phloem 55 Fruit Ripening 93 Time of Growth Initiation and Amounts of Xylem Sexual Reproduction in Gymnosperms 96 and Phloem Produced 56 Cone Initiation and Development 96 Differentiation of Cambial Derivatives 56 Polyembryony 99 Increase in Cell Size 58 Parthenocarpy 99 Hormonal Infl uences on Cambial Growth 58 Duration and Timing of Cone Development 99 Cell Wall Thickening 62 Increase in Size and Dry Weight of Cones Loss of Protoplasts 62 and Seeds 100 Formation and Development of Rays 63 Maturation of Seeds 102 Expansion of the Cambium 63 Abscission of Reproductive Structures 103 Variations in Growth Increments 63 Abscission and Crop Yield 103 Seasonal Duration of Cambial Growth 64 Summary 105 Anomalous Cambial Growth 64 General References 106 Sapwood and Heartwood Formation 64 Wounding and Wound Healing 67 CHAPTER Root Growth 68 Root Elongation 69 Rate of Root growth 70 5 Seasonal Variations 70 Photosynthesis 107 Cambial Growth in Roots 71 Shedding of Plant Parts 72 Introduction 107 Leaves 73 Chloroplast Development and Structure 108 Branches 77 Pigments 109 Bark 78 Proteins 110 Roots 79 Membrane Systems 110 Measurement and Analysis of Growth 80 The Photosynthetic Mechanism 110 Analysis of Growth 80 Light Reactions 110 Relative Growth Rates 81 Photochemistry 111 Allometric Formula and the Allometric Electron Transport 112 Coeffi cient 81 NADP+ Reduction 112 Net Assimilation Rate and Other Growth Photophosphorylation 112 Parameters 82 Photoinhibition 113 Limitations of Traditional Growth Analysis for Dark Reactions 116 Woody Plants 84 Carbon Dioxide Uptake by Photosynthetic Tissues 119 Summary 85 Carbon and Oxygen Isotope Discrimination During General References 86 Photosynthesis 121 Contents ix Variations in Rates of Photosynthesis 122 Scaling of Respiration to the Ecosystem Level 185 Species and Genetic Variations 123 Respiration of Harvested Fruits 187 Photosynthesis and Productivity 124 Factors Affecting Respiration 188 Diurnal Variations 126 Age and Physiological Condition of Tissues 188 Seasonal Variations 128 Available Substrate 188 Environmental Factors 132 Light 188 Light Intensity 132 Hydration 188 Air Temperature 140 Temperature 189 Soil Temperature 142 Composition of the Atmosphere 190 Carbon Dioxide 144 Soil Aeration 191 Water Supply 147 Mechanical Stimuli and Injuries 191 Soil Drying and Photosynthesis 147 Chemicals 192 Humidity 151 Air Pollutants 193 Flooding 152 Assimilation 194 Mineral Nutrition 152 Summary 195 Salinity 155 General References 197 Pollution 156 Applied Chemicals 158 CHAPTER Plant Factors 158 Stomatal Characteristics and Capacity of 7 Photosynthetic Partial Processes 159 Carbohydrates 199 Source-Sink Relations 160 Age of Leaves and of the Plant 162 Introduction 199 Summary 164 Kinds of Carbohydrates 199 General References 166 Monosaccharides 199 Oligosaccharides 200 CHAPTER Polysaccharides 201 Carbohydrate Transformations 204 6 Phosphorylation 204 Sucrose 205 Enzymes, Energetics and Respiration 169 Starch 205 Uses of Carbohydrates 205 Introduction 169 Respiration 206 Enzymes and Energetics 169 Growth 206 Enzymes 169 Defense 207 Energetics 172 Leaching 207 Respiration 173 Exudation 208 Biological Oxidations 174 Accumulation of Carbohydrates 208 ATP 174 Carbohydrate Distribution 208 Other High-Energy Compounds 175 Storage Sites 208 Glycolysis and the Krebs Cycle 175 Autumn Coloration 211 Electron Transfer and Oxidative Summary 214 Phosphorylation 176 General References 215 Metabolic Decomposition of Respiration 176 Other Oxidases 177 CHAPTER The Pentose Shunt 178 Anaerobic Respiration 178 8 Respiratory Quotient 179 Photorespiration 179 Lipids, Terpenoids, and Related Respiration of Plants and Plant Parts 180 Substances 217 Amount of Food Used in Respiration 180 Respiration of Entire Trees 180 Introduction 217 Respiration of Various Plant Parts 181 Lipids 218 Seasonal Variations 185 Simple Lipids 218 x Contents Fatty Acids 218 CHAPTER Lipid Distribution 219 Waxes, Cutin, and Suberin 220 10 Cuticle 220 Mineral Nutrition 255 Waxes 220 Cutin and Suberin 222 Introduction 255 Internal Lipids 223 Functions of Mineral Nutrients and Effects Phospholipids 223 of Defi ciencies 256 Glycolipids 224 Nitrogen 256 Membrane Lipids 224 Phosphorus 256 Isoprenoids or Terpenoids 224 Potassium 256 Essential Oils 224 Sulfur 257 Resins 225 Calcium 257 Oleoresins 226 Magnesium 258 Monoterpenes 227 Iron 258 Carotenoids 229 Manganese 259 Rubber 229 Zinc 259 Related Compounds 231 Copper 259 Summary 231 Boron 259 General References 232 Molybdenum 260 Chlorine 260 Nickel 260 Other Mineral Nutrients 260 CHAPTER Accumulation and Distribution of Mineral Nutrients 261 9 Mineral Cycling 262 The Soil Mineral pool 262 Nitrogen Metabolism 233 Atmospheric Deposition 262 Leaching from Plants 264 Introduction 233 Throughfall and Stemfl ow 266 Distribution and Seasonal Fluctuations Weathering of Rocks and Minerals 268 of Nitrogen 234 Decomposition of Organic Matter 268 Concentration in Various Tissues 234 Temperature 269 Seasonal Changes in Nitrogen Exudation from Roots 270 Concentration 235 Losses of Mineral Nutrients From Ecosystems 270 Changes in Distribution with Age 238 Ecosystem