BASIC GROWTH ANALYSIS

BASIC GROWTH ANALYSIS

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BASIC GROWTH ANALYSIS Plant growth analysis for beginners RODERICK HUNT BSc, PhD, CBiol, MIBiol Independent Research Worker, Natural Environment Research COl/llcil, Unit of Comparative Plallt Ecology Honorary Lecturer in BotallY, University of Sheffield LONDON UNWIN HYMAN Boston Sydney Wellington

:9 Roderick Hunt, 1990 Softcover reprint of the hardcover 1st edition 1990 This book is copyright under the Berne Convention. No reproduction without permission. All rights reserved. Published by the Academic Division of Unwin Hyman Ltd 15/17 Broadwick Street, London W1V 1Fp, UK Unwin Hyman Inc., 8 Winchester Place, Winchester, Mass. 01890, USA Allen & Unwin (Australia) Ltd, 8 Napier Street, North Sydney, NSW 2060, Australia Allen & Unwin (New Zealand) Ltd in association with the Port Nicholson Press Ltd, Compusales Building, 75 Ghuznee Street, Wellington 1, New Zealand First published in 1990 British Library Cataloguing in Publication Data Hunt, Roderick 1945- Basic growth analysis 1. Plants. Growth. Analysis. Quantitative methods. 1. Title. 581.3'1 ISBN-13: 978-0-04-445373-4 e-isbn-13: 978-94-010-9117-6 DOl: 10.1007/978-94-010-9117-6 Library of Congress Cataloging in Publication Data Hunt, Roderick, 1945- Basic growth analysis: plant growth analysis for beginners/ Roderick Hunt. p. cm. Bibliography: p. Includes index. ISBN-13: 978-0-04-445373-4 1. Growth (plants)-experiments. 2. Growth (Plants)-Mathematical models. 3. Growth (Plants)-Measurement. 1. Title.. QK731.H85 1989 581.3T015118-dc20 89-16462 CIP Typeset in 10 on 12 point Palatino by MCS, Salisbury, England and

Preface This handbook is intended as an introductory guide to students at all levels on the principles and practice of plant growth analysis. Many have found this quantitative approach to be useful in the description and interpretation of the performance of whole plant systems grown under natural, semi-natural or controlled conditions. Most of the methods described require only simple experimental data and facilities. For the classical approach, GCSE biology and mathematics (or their equivalents) are the only theoretical backgrounds required. For the functional approach, a little calculus and statistical theory is needed. All of the topics regarding the quantitative basis of productivity recently introduced to the Biology A-level syllabus by the Joint Matriculation Board are covered. The booklet replaces my elementary Plant Growth Analysis (1978, London: Edward Arnold) which is now out of print. The presentation is very basic indeed; the opening pages give only essential outlines of the main issues. They are followed by brief, standardized accounts of each growth-analytical concept taken in turn. The illustrations deal more with the properties of well-grown material than with the effects of specific environmental changes, even though that is where much of the subject's interest lies. However, detailed references to the relevant parts of more comprehensive works appear throughout, and a later section on 'Interrelations' adds perspective. Some 'Questions and answers' may also help to show what topics will arise if the subject is pursued further. RH Sheffield, 1989

Acknowledgements The permissions of the authors and publishers of the various illustrative examples are gratefully acknowledged. Though not specilically connected with this project, Drs D. R. Causton and G. C. Evans stand foremost among the many friends who have been of personal assistance to these endeavours over the years. Mrs A. M. N. Ruttle has provided incomparable secretarial support and Professor J. N. R. Jeffers kindly commented on the manuscript. The Unit of Comparative Plant Ecology is part of the Terrestrial and Freshwater Sciences Directorate of the Natural Environment Research Council. We are grateful to the following individuals and organizations who have kindly given permission for the reproduction of copyright material (ligure numbers in parentheses): R. Sattler and Springer-Verlag (Table 1.3); British Ecological Society (Table 2.1); D. R. Causton and Edward Arnold (2.2,4.3,4.5,4.11); W. J. Leverich and the University of Chicago Press (Tables 2.2, 3.2); Annals of Botany Company (2.3, 2.4, 4.4, 4.6, 5.2, 5.4, 8.5, Tables 4.7, 8.3); The Editor, The New Phytologist (3.1, 3.2, 3.3, 4.1, 5.1, 5.9, 8.1, 8.2); T. Ingestad and The Editor, Physiologia Plantarum (Tables 3.1, 5.3); E. Ashby and the Annals of Botany Company (3.4); S. Arvidson and Ellis Horwood Ltd (3.5); The Editor, Journal of General Microbiology (3.6); The Editor, Journal of Ecology (Tables 4.1, 8.1); L. T. Evans and Cambridge University Press (Tables 4.4, 4.6, Figure 6.1); C. B. Johnson and Butterworths (Table 4.5); Cambridge University Press (4.7); J. K. A. Bleasdale and Macmillan Publishers Ltd (4.8); The Editor, Journal of the British Grassland Society (4.9); F. H. Whitehead and The Editor, The New Phytologist (4.10); J. Coombs, 1982 Pergamon Press PLC (Table 5.1); The Editor, Australian Journal of Agricultural Research (Table 5.2); R. F. Williams and the Annals of Botany Company (5.3); R. F. Williams and Cambridge University Press (Table 5.4); P. J. Welbank and the Annals of Botany Company (5.6); D. P. Stribley and The Editor, The New Phytologist (5.7); P. J. Dudney and the Annals of Botany Company (5.8); J. Kvet (Table 6.1); J. Kvet and The Editor, PhotosYllthetica (6.2, Table 6.2); D. R. Tottman (7.1); The Editor, Physiologia Plantarum (8.3); R. M. Shibles and Cambridge University Press (8.4); Institute of Terrestrial Ecology (Ch.9 Checklist).

Contents Preface Acknowledgements page vii ix 1 Introduction 1 Growth 2 Plant growth 3 Plant growth analysis 5 'Classical' and 'functional' approaches 8 Harvesting technique 10 Units 11 Notation 12 Computing, mathematics, statistics 13 Concepts and levels 14 2 Absolute growth rates 17 Absolute growth rate in size (AGR) 18 Absolute growth rate in number 20 Absolute growth rate in populations 22 Concluding remarks 24 3 Relative growth rates 25 Relative growth rate (RGR) 26 Other relative growth rates 28 Intrinsic rate of population increase 30 Relative growth rates in biotechnology 32 Concluding remarks 34 4 Simple ratios 35 Leaf area ratio (LAR) 36 Speciftc leaf area (SLA) 38 Leaf weight ratio (LWR) 40 Leaf area index (LAI) 42 xi

Efficiency of energy conversion digestible energy 44 Harvestable dry matter (harvest index) 46 Harvestable protein 48 Root-shoot allometric coefficient 50 Other allometric coefficients 52 Concluding remarks 54 5 Compounded growth rates 55 Unit leaf rate (ULR) 56 Other leaf efficiencies 58 Crop growth rate (CGR) 60 Speciftc absorption rate (SAR) 62 Speciftc utilization rate (SUR) 64 Sub-cellular efficiencies 66 Unit production rate (UPR) 68 Component production rate (CPR) 70 Concluding remarks 72 6 Integral durations 73 Leaf area duration (LAD) 74 Biomass duration (BMO) 76 Concluding remarks 78 7 Other independent variables 79 8 Interrelations 83 In general 84 Between ratios 86 Between rates Within individual plants 8& 90 Within populations and communities 92 An integrated scheme 96 Concluding remarks 98 9 Questions and answers 99 Tables of synopses 107 Synopsis of the main derived quantities Synopsis of primary quantities and associated terms 108 110 References 111 xii