CONTRIBUTORS PAULO De T. ALVIM RAIMUNDO SANTOS BARROS
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CONTRIBUTORS PAULO de T. ALVIM VICENTE H. F. MORAES RAIMUNDO SANTOS BARROS G. Ο. MOTT D. P. BARTHOLOMEW D. B. MURRAY JAKOB BENNEMA M. C. NAMBIAR JAN-DIRK FERWERDA LOUIS G. NICKELL R. FORDHAM H. L. POPENOE S. K. HAHN WALTER REUTHER S. Β. KADZIMIN L. B. SINGH MOACYR MAESTRI EGBERT A. TAI J. L. MONTEITH L. A. WILSON SHOUICHI YOSHIDA Ecophysiology of Tropical Crops Edited by PAULO de T. ALVIM Centro de Pesquisas do Cacau Comissâo Executiva do Piano da Lavoura Cacaueira Itabuna, Bahia, Brazil T. T. KOZLOWSKI Russell Laboratories Department of Forestry University of Wisconsin Madison, Wisconsin ACADEMIC PRESS New York San Francisco London 1977 A Subsidiary of Harcourt Brace Jovanovich, Publishers COPYRIGHT © 1977, 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 Library of Congress Cataloging in Publication Data Main entry under title: Ecophysiology of tropical crops. "Based to a large extent on material presented at an international symposium . .. held in Manaus, Brazil in May, 1975." Includes bibliographies and index. 1. Tropical crops-Ecology-Congresses. 2. Tropical crops-Physiology-Congresses. I. Alvim, Paulo de T. II. Kozlowski, Theodore Thomas, Date SB111.A2E26 63l'.0913 76-55967 ISBN 0-12-055650-2 PRINTED IN THE UNITED STATES OF AMERICA List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. PAULO de T. ALVIM (279), Centro de Pesquisas do Cacau, Comissàb Executiva do Piano da Lavoura Cacaueira, Itabuna, Bahia, Brazil RAIMUNDO SANTOS BARROS (249), Divasâo de Fisiologia Vegetal, Centro de Pesquisas do Cacau, Itabuna, Bahia, Brazil D. P. BARTHOLOMEW (113), Department of Agronomy and Soil Science, University of Hawaii, Honolulu, Hawaii JAKOB BENNEMA (29), Department of Soil Science and Geology, Agricultural University, Wageningen, The Netherlands JAN-DIRK FERWERDA (351), Department of Tropical Crop Science, Agricul- tural University, Wageningen, The Netherlands R. FORDHAM (333), Commodity Development Unit, Mars Ltd., Slough, England S. K. HAHN (237), International Institute of Tropical Agriculture, Ibadan, Nigeria S. B. KADZIMIN (113), Department of Agronomy and Horticulture, University of Agriculture Malaysia, Serdang, Selangor, Malaysia MOACYR MAESTRI (249), Departmento de Biologia, Universidade Federal de Viçosa, Minas Gérais, Brazil J. L. MONTEITH (1), Department of Physiology and Environmental Studies, University of Nottingham School of Agriculture, Sutton Bonington, Loughborough, Leics, England xi xii List of Contributors VICENTE H. F. MORAES (315), Centro National de Pesquisas da Seringueira, EMBRAPA, Manaus, Brazil G. Ο. MOTT (157), Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida D. B. MURRAY (383), Faculty of Agriculture, University of the West Indies, St. Augustine, Trinidad, West Indies M. C. NAMBIAR (461), Central Plantation Crops Research Institute, Kasaragod, Kerala, India LOUIS G. NICKELL (89), Research Division, W. R. Grace & Co., Washington Research Center, Columbia, Maryland H. L. POPENOE (157), Institute of Food and Agricultural Sciences, University of Florida, Gainesville, Florida WALTER REUTHER (409), Department of Plant Science, University of Cali- fornia, Riverside, California L. B. SINGH (479), Horticultural Research Institute, Saharanpur, Uttar Pradesh, New Delhi, India EGBERT A. TAI (441), Ministry of Agriculture and Supply, St. John's, Antigua, West Indies L. A. WILSON (187), Faculty of Agriculture, Department of Crop Science, University of the West Indies, St. Augustine, Trinidad, West Indies SHOUICHI YOSHIDA (57), International Rice Research Institute, Manila, Phil- ippines Preface The threat of impending food shortages at a time when population is increasing at an alarming rate emphasizes the importance of tropical crops to the health and economy of people within the developing tropical countries and elsewhere. Improving the productive potential of tropical crops on a sustained basis will depend on increased understanding of tropical climates and soils and the way in which environmental and cultural factors affect plant growth through the intermediation of physiological processes. These considerations, as well as a rapidly expanding body of research results, prompted our bringing together in a single volume the current state of knowledge and opinion on ecophysiology of the major tropical crop plants. The opening chapter reviews fundamental ideas about numerical description of plant development and considers effects of climatic factors (e.g., temperature, light, and water) on physiological processes in plants. The second chapter presents an overview of physical and chemical characteristics of tropical soils. Separate chapters then follow on ecophysiology of the major crop plants, particularly those suitable for the wet tropics, including rice, sugarcane, pine- apple, grasslands, root crops, sweet potato, coffee, cacao, rubber, banana, tea, oil palm, coconut palm, citrus, cashew, and mango. The book is both authoritative and interdisciplinary. It should be useful to students and researchers, as well as those interested in production of tropical crops. The subject matter will be of interest to plant ecologists, plant physiolo- gists, biochemists, horticulturists, agronomists, meteorologists, soil scientists, food technologists, and plant breeders. xiii xiv Preface This volume is based to a large extent on material presented at an Interna- tional Symposium on Ecophysiology of Tropical Crops held in Manaus, Brazil in May, 1975. Invitations were extended to leading investigators concerned with growth and ecological requirements of tropical crops. Their contributions reflect many years of experience with tropical crops in Africa, Southeast Asia, Latin America, and the Caribbean Islands. We thank the following organizations in Brazil for generous financial support and facilities for the Symposium: National Council for Scientific and Techno- logical Development (CNPq.), Superintendency of the Free Zone of Manaus (SUFRAMA), the Executive Committee of the Plan for Cacao Production (CEPLAC), and the Inter-American Institute of Agricultural Sciences (IAIAS). We also thank each author for his contribution, patience, and cooperation during the production phases. The senior editor expresses his deep gratitude to the following who assisted in organizing the Manaus Symposium: Luiz Carlos Cruz (CEPLAC), W. Martin Aitken (CEPLAC), Antonio Dantes Machado (CNPq.), and Marcilio Junqueira (SUFRAMA). Mr. T. L. Noland and Mr. R. J. Norby assisted with the preparation of the subject index. Paulo de T. Alvim T. T. Kozlowski CHAPTER 1 Climate J. L. MONTEITH I. Introduction 1 II. Specification of Development 2 III. Temperature 3 A. Physical Aspects 3 B. Physiological Aspects 4 C. Day-Degrees 6 IV. Light 8 A. Physical Aspects 8 B. Physiological Aspects 10 V. Water 14 A. Physical Aspects 14 B. Physiological Aspects 18 C. Ecological Aspects 19 VI. Conclusions 21 A. Integration 21 B. Omissions 24 References 25 I. INTRODUCTION How does the climate of a region determine what crop species will thrive? To what extent are yields determined by sequences of weather from month to month and from year to year? These are formidable questions, but they are central to agricultural meteorology; they have been attacked from many differ- ent directions; and they are still largely unanswered. 1 2 J. L. Monteith As we are concerned with environmental physiology of tropical crops, the effects of climate must be discussed with reference to specific physiological processes. The first section of this chapter is therefore concerned with some fundamental but simple points about the numerical description of plant develop- ment; subsequent sections will consider the effects of temperature, light, and water supply on growth and yield. By adopting this treatment, I have tried to avoid three major pitfalls of conventional crop ecology: (1) statistical treatments of crop-weather relations, which lack generality and usually conceal an igno- rance of mechanisms; (2) conventional growth analysis in terms of net assimila- tion rates and relative growth rates, valid for single, isolated plants, but not for plant communities in which leaves are competing for light; and (3) complex dynamic models of crop growth, which are becoming so bulky and unwieldy that they may suffer the fate of the dinosaurs. II. SPECIFICATION OF DEVELOPMENT In mathematical terms, plants consist of an assembly of discrete units each of which has measurable attributes such as number, size, and weight. If the development of a plant is defined as the outcome of processes of differentiation and growth (Wareing and Phillips, 1970), the rate of differentiation can be expressed by an increase in primordial numbers, whereas the growth of an organ is expressed and measured by an increase in its size or weight. Cell division and expansion provide a basic example of the distinction between numbers and size, but it is applicable to any higher level of organization — to numbers of leaf primordia or florets, to lengths of stems, areas of leaves, or weights of tubers. The physical dimensions of differentiation are