PLANT PHYSIOLOGY A TREATISE Volume ΙΑ. Cellular Organization and Respiration Volume IB. Photosynthesis and Chemosynthesis Volume II. Plants in Relation to Water and Solutes Volume III. Inorganic Nutrition of Plants Volume IVA. Metabolism: Organic Nutrition and Nitrogen Metabolism Volume IVB. Metabolism: Intermediary Metabolism and Pathology Volume VA. Analysis of Growth: Behavior of Plants and Their Organs Volume VB. Analysis of Growth: The Responses of Cells and Tissues in Culture Volume VIA. Physiology of Development: Plants and Their Reproduction Volume VIB. Physiology of Development: The Hormones Volume VIC. Physiology of Development: From Seeds to Sexuality Plant Physiology A T R E A T I SE EDITED BY Ε C STEWARD Laboratory of Cell Physiology, Growth, and Development Cornell University Ithaca, New York P h y s i o l o gy of D e v e l o p m e nt : Volume VIA : P l a n ts a nd T h e ir R e p r o d u c t i on 1971 A C A D E M IC P R E S S, New York and London COPYRIGHT © 1971, BY ACADEMIC PRESS, INC. ALL RIGHTS RESERVED NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM, BY PHOTOSTAT, MICROFILM, RETRIEVAL SYSTEM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM THE PUBLISHERS. 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 7DD LIBRARY OF CONGRESS CATALOG CARD NUMBER: 59-7689 PRINTED IN THE UNITED STATES OF AMERICA CONTRIBUTORS TO VOLUME VIA J. P. NITSCH,* Centre National de la Recherche Scientifique, Laboratoire de Physiologie Pluricellulaire, Gif-sur- Yvette, France JOHN R. RÂPER, The Biological Laboratories, Harvard University, Cam- bridge, Massachusetts W. W. SGHWABE, Wye College, University of London, Wye nr. Ashford, Kent, England T. A. STEEVES, Department of Biology, University of Saskatchewan, Saska- toon, Saskatchewan, Canada R. H. WETMORE, The Biological Laboratories, Harvard University, Cam- bridge, Massachusetts * Deceased. vii PREFACE Growth (as considered in Volume V) and development (as treated in Volume VI) are such closely complementary topics that it is diffi- cult, and often arbitrary, to treat the one without involving the other. They are related as the two faces of a coin. To this extent, the intro- duction to these problems in Volume VA is equally applicable to Volume VI. While the original plan to deal with the physiology of development has been retained, the problems that arose before it could eventually be carried out merit some mention. The physiology of development is now presented in three parts, and, within the essential design of the treatise, they comprise Volumes VIA, B, and C. This device became necessary as, with the passing years, topics became more complex and some chapters longer. When these problems arose, the decision was made to cover the subject matter as desired by authors rather than to compress it arbitrarily into a prescribed space. Inevitably, the allocation of subjects and chapters to the several parts A, Β and C is in a measure arbitrary—it had to be adapted to their respective lengths, to the time of their completion, as well as to their content. The rationale is as follows. During development, one sees the emergence of form within an inherited, evolved, plan. It is appropriate, therefore, to introduce the problems of development and reproduction morphologically ; this is done in two chapters, one largely concerned with higher plants and the other with lower forms (especially fungi). The fungi have unique properties which render them both useful in physiological research and especially important to man. Part A, therefore, contains four chapters, conveniently grouped under the subtitle Plants and Their Reproduction, and they embrace the various problems that arise as plants, responsive to environmental stimuli, develop a vegetative plant body, produce seeds and fruits or organs of perennation. Part Β is the outgrowth of an original chapter destined to deal with the hormone concept in plant physiology—with its history and present status. Although references to plant hormones and hormonal action occur throughout the treatise, they needed to be synthesized and integrated in a chapter devoted to an overview of plant hormones. But the volume of work and the new classes of active, growth-regulat- ing substances necessitated both a comprehensive restatement of their nature and actions, even as it required more specialists in its comple- tion. But, to conform to the original design and to retain earlier cross ix χ PREFACE references, Part Β, subtitled The Hormones, is organized as a chapter in Volume VI of the treatise. Part C could be called Ontogeny, but because it is largely devoted to angiosperms it is entitled From Seeds to Sexuality. Three concise papers on Germination, on Cell Growth and Cell Development, and on Cell Division in Higher Plants treat these topics from a single point of view, and they focus attention on the problems of early develop- ment in angiosperms. Part VIC is completed by a comprehensive treatment of sexuality in flowering plants. Appropriately, the volume concludes with problems that arise, and points of view that emerge, as development is considered in the light of genetics. With the lapse of time, the topics dealt with in Volume VI are up to date and, in so presenting them, much factual information that also relates to earlier volumes has been included. Thus, the treatise, as the organisms with which it deals, has developed, and in the progress of the whole its parts have been renewed. As always, the merit of the volume rests upon the work of the individual authors; for this and for their forbearance with my prob- lems I am grateful. The understanding of the staff of Academic Press in the face of circumstances which complicated production is gratefully acknowledged. Again, Dr. William J. Dress of the L. H. Bailey Hortorium, Cornell University, relieved me by considering all problems of nomenclature and by preparing the Index of Plant Names. For Volumes VIA, B, and C the Subject Indexes were prepared by Mr. George H. Craven. F. C. STEWARD CONTENTS OF OTHER VOLUMES Volume IA. CELLULAR ORGANIZATION AND RESPIRATION Historical Introduction—F. C. STEWARD The Plant Cell and Its Inclusions—R. BROWN Proteins, Enzymes, and the Mechanism of Enzyme Action—BIRGIT VENNESLAND Cellular Respiration—DAVID R. GODDARD AND WALTER D. BONNER Volume IB. PHOTOSYNTHESIS AND CHEMOSYNTHESIS Energy Storage: Photosynthesis—HANS GAFFRON Chemosynthesis : The Energy Relations of Chemoautotrophic Orga- nisms—MARTIN GIBBS AND JEROME A. SGHIFF Volume II. PLANTS IN RELATION TO WATER AND SOLUTES Cell Membranes : Their Resistance to Penetration and Their Capacity for Transport—RUNAR COLLANDER Water Relations of Cells—T. A. BENNET-CLARK The Water Relations of Stomatal Cells and the Mechanisms of Stomatal Movement—Ο. V. S. HEATH Plants in Relation to Inorganic Salts—F. C. STEWARD AND J. F. SUTCLIFFE Translocation of Organic Solutes—C. A. SWANSON Translocation of Inorganic Solutes—O. BIDDULPH Transpiration and the Water Economy of Plants—PAUL J. KRAMER Volume III. INORGANIC NUTRITION OF PLANTS Mineral Nutrition of Plants in Soils and in Culture Media—C. BOULD AND E.J. HEWITT The Essential Nutrient Elements: Requirements and Interactions in Plants—E.J. HEWITT Inorganic Nutrient Nutrition of Microorganisms—D.J. D. NICHOLAS Modes of Action of the Essential Mineral Elements—ALVIN NASON AND WILLIAM D. MGELROY Biological Nitrogen Fixation—ARTTURI I. VIRTANEN AND JORMA K. MIETTINEN Microbial Activities of Soil as They Affect Plant Nutrition—J. H. QUASTEL xii CONTENTS OF OTHER VOLUMES Volume IVA. METABOLISM: ORGANIC NUTRITION AND NITROGEN METABOLISM Introduction: The Concept of Metabolism—F. C. STEWARD Photosynthesis (Carbon Assimilation) : Environmental and Metabolic Relationships—MOYER D. THOMAS Micrometeorology and the Physiology of Plants in Their Natural Environment—EDGAR LEMON The Respiration of Plants and Their Organs—E. W. YEMM The Respiration of Bulky Organs—DOROTHY F. FORWARD Metabolism of Nitrogenous Compounds—F. C. STEWARD AND D. J. DURZAN Volume IVB. METABOLISM: INTERMEDIARY METABOLISM AND PATHOLOGY Carbohydrates: Their Role in Plant Metabolism and Nutrition— MARTIN GIBBS Metabolism of the Organic Acids—H. BEEVERS, M. L. STILLER, AND V. S. BUTT The Plant Lipids—V. S. BUTT AND H. BEEVERS The Physiology of Parasitic Disease—E. G. BOLLARD AND R. E. F. MATTHEWS Volume VA. ANALYSIS OF GROWTH: BEHAVIOR OF PLANTS AND THEIR ORGANS The Quantitative Analysis of Growth—F. J. RICHARDS Phyllotaxis: A Problem of Growth and Form—F. J. RICHARDS AND W. W. SCHWABE Tropic, Nastic, and Tactic Responses—NIGEL G. BALL Chemical Plant Growth Regulation—R. L. WAIN AND C. H. FAWCETT Environmental Factors in Regulation of Growth and Development: Ecological Factors—F. W. WENT AND LILLIAN OVERLAND SHEPS Volume VB. ANALYSIS OF GROWTH: THE RESPONSES OF CELLS AND TISSUES IN CULTURE Growth in Organized and Unorganized Systems—H. E. STREET Biochemical Differentiation : The Biosynthetic Potentialities of Grow- ing and Quiescent Tissue—A. D. KRIKORIAN AND F. C. STEWARD Growth and Morphogenesis in Tissue and Free Cell Cultures—F. C. STEWARD, WITH M. O. MAPES AND P. V. AMMIRATO Abnormal Growth in Plants—ARMIN C. BRAUN CONTENTS OF OTHER VOLUMES xiii Volume VIB. PHYSIOLOGY AND DEVELOPMENT: THE HORMONES The Natural Plant Hormones—Κ. V. THIMANN With sections by L. G. PALEG AND C. A. WEST, and F. SKOOG AND R. Y. SCHMITZ Volume VIC. PHYSIOLOGY OF DEVELOPMENT: FROM SEEDS TO SEXUALITY ( Tentative ) Germination—R. BROWN Cell Division in Higher Plants—R. BROWN AND A. F. DYER Cell Growth and Cell Development—R. BROWN Sexuality of Angiosperms—J. HESLOP-HARRISON Developmental Genetics: With Emphasis on Lower Eucaryotic Plants—A. M. SRB Developmental Genetics of Higher Plants: An Outline—J. HESLOP- HARRISON Integration and Organization: Control Mechanisms—F. C. STEWARD PLANT PHYSIOLOGY The Plan of the Treatise The treatise is planned in three main sections, as follows : Section on Cell Physiology and Problems Relating to Water and Solutes The purpose of this section is to present the properties of cells, their energy relations (Volume I), and behavior toward water and solutes with the closely related problems of the movement of solutes within the plant body and the economy of water in plants (Volume II). The underlying theme of Volumes I and II is the basis of plant physiology in cell physiology. Section on Nutrition and Metabolism In this section the detailed facts and knowledge of nutrition and metabolism are presented, first with reference to the need for, and utilization of, inorganic nutrients (Volume III), and second with respect to the processes of organic nutrition (Volume IV). The treat- ment of organic nutrition leads to a reconsideration of photosynthesis and respiration at the level of organs and organisms. Volume IV describes the intermediary metabolism of carbon and nitrogenous compounds and presents a brief comparison of plants in health and in disease. The main theme of Volumes III and IV is the nutrition, organic and inorganic, of plants and the biochemical steps by which these processes are achieved. Section on Growth and Development The purpose of the last section is to present the problems of plant physiology as seen through the analysis of growth and development, mainly with reference to flowering plants. As reconstituted, Volume V now contains an appraisal of the main events of growth as seen through the behavior of plants and their organs (VA) and, later, of cultured cells and tissues (VB). The quantitative interpretation of growth, the effects of environmental factors, the treatment of tropisms and of the effects of many exogenous growth-regulating compounds are all dis- cussed in Volume VA. Volume VB deals, collectively, with various aspects of the normal and abnormal growth of cells and tissues in culture. Thus Volume VI will be concerned with various aspects of development; these will include a morphological approach to de- XV