Physiology and Biochemistry of Seeds

Transcription

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2 Physiology and Biochemistry of Seeds in Relation to Germination In Two Volumes 1 J. D. Bewley. M. Black Development, Germination, and Growth Springer-Verlag Berlin Heidelberg GmbH 1978

3 Dr. J. DEREK BEWLEY, Department of Biology, University ofcalgary, Calgary, Alberta, Canada T2N 1 N4 Dr. MICHAEL BLACK, Department of Biology, Queen Elizabeth College, University oflondon, Campden Hill Road, London W8 7 AH, Great Britain With 122 Figures ISBN DOI DOl / ISBN (ebook) Library of Congress Cataloging in Publication Data. Bewley. J. Derek ~. Physiology and biochemistry of seeds in relation to germination. Includes bibliographical references and index. Contents: v.l. v.1. Development, germination, and growth. 1. I. Seeds. 2. Germination. I. 1. Black, Michael, joint author, II. Title. DNLM: 1. I. Seeds~Physiology. 2. Seeds~Growth and development. Qk740 B572p QK661.B '.01' This work is subject to copyright. All AII rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under 54 of the German Copyright Law, where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. by Springer-Verlag Berlin Heidelberg Originally published by Springer-Verlag erlag Berlin Heidelberg New York in 1978 Softcover So/kover reprint of the hardcover 1 st edition 1978 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. 2131/

4 "The true writer will let his wife starve, his children go barefoot, his mother drudge for his living at seventy sooner than work at anything but his book" Slightly amended, from G.B. Shaw's "Man and Superman" (with apologies) Dedicated to Christine Bewley and Marianne Black and their respective children who, despite their initial shock on discovering that they had aspiring writers as husbands and fathers, proved to be pillars of strength and encouragement.

5 Acknowledgements First and foremost we owe a debt to the thousands of scientists who have studied and written about various aspects of seed biology. Many of their contributions have been drawn upon to sustain our efforts but inevitably much work receives no reference, if only because of limitations of space and time. Especial thanks are due to Drs. D.L. Laidman, M. Parker, P.I. Payne, P. Slack and D.L. Smith who allowed us to use their unpublished work and to many others for the use of published photographs and graphs: these sources are indicated with the appropriate figures. We thank Dr. P. Halmer for valuable discussions and critical advice and 1. Pacey for photographic contributions. We also express our appreciation to our respective research students and other colleagues who soldiered on at their laboratory benches while we were immured in the libraries. Queen Elizabeth College, University of London, kindly gave facilities to J.D. Bewley during his sabbatical leave which was generously permitted by the University of Calgary and supported by the British Council, the Department of External Affairs, Government of Canada and the National Research Council of Canada. Thanks to the stamina, conscientiousness and excellent typing of Catherine Wardale our manuscript was transformed into a presentable condition. We sincerely thank her and Gwen Turner, also in London, who helped out when the pressure became too high. The final typescript was produced by Laurie Dowson and Linda Dunne of Calgary for whose efforts we are most grateful.

6 Contents Chapter 1. Introduction The Subject Matter of This Book The Seed.... What is Germination?.... How is Germination Measured? Some Commen ts on Our Sources Plant Names Some Articles of General Interest References Chapter 2. The Structure of Seeds and Their Food Reserves Seed Structure..... The Testa.... Perisperm and Endosperm The Embryo... Food Reserves... Location of Reserves Protein Storage Proteins of Legumes Storage Proteins of Gramineae Protein Bodies Lectins Other Nitrogenous Seed Reserves Phytin... Carbohydrates Starch.... Starch Grains. Sugars... Other Polysaccharides Oils or Fats (" Lipids") Oil Bodies Some Articles of General Interest References Chapter 3. The Legacy of Seed Maturation General Developmental Pattern Filling of the Grain (Kernel) in Cereals.. Source of Assimilates for Starch Formation I I

7 VIII Contents Development of the Starchy Endosperm and Aleurone Layer The Synthesis of Starch Protein and RNA Synthesis in the Developing Endosperm Storage of Phosphate Establishment of Cotyledon Reserves in Dicots Non-Endospermic Legumes Endospermic Legumes Lipid-Storing Seeds Fat (Oil) Synthesis Fatty Acid Synthesis Synthesis of Triglycerides (Triacylglycerols) General Pattern of Oil Formation Development of Oil-Storing Bodies Protein Synthesis in Fat-Storing Seeds Energy Supply in the Maturing Seed The Fate of the Synthesizing Machinery Conservation of the Protein-Synthesizing Apparatus During Drying Hormones and Seed Development Gibberellins Auxins Cytokinins Abscisic Acid The Role of Hormones in the Developing Seed 99 Some Articles of General Interest 101 References Chapter 4. Imbibition, Germination, and Growth l l.4 4.l Uptake of Water Some Theoretical Considerations... Soil Matric Potential and Seed-Soil Contact Seed Germination in Petri Dishes. Kinetics of Water Uptake by Seeds.... Soaking Injury and Solute Leakage.... Radicle Expansion-Cell Elongation or Cell Division? The Control of Germination.. Seedling Development..... Some Articles of General Interest References Chapter 5. Biochemistry of Germination and Growth Respiration - Pathways and Products Requirement for Oxygen by Seeds. Respiration of Germinating Seeds

8 Contents IX The Lag Phase in Respiration - Possible Causes Seeds Without a Lag Phase of Respiration Sources of Substrate for Respiration Prior to Reserve Mobilization Mitochondrial Activity and A TP Synthesis Reducihg Power-the Synthesis and Utilization of Pyridine Nucleotides Special Oxidation Systems Protein and Nucleic Acid Synthesis The Mechanism of Protein Synthesis Protein Synthesis in Imbibing Embryos and Axes Messenger RNA - Conserved, Synthesized, or Both? Ribosomal and Transfer RNA Synthesis RNA and Protein Synthesis in Storage Tissues DNA Synthesis, Germination, and Growth DNA in Storage Tissues Some Articles of General Interest 173 References Chapter 6. Mobilization of Reserves 6.1 Stored Carbohydrate Metabolism General Metabolism of Starch Sucrose Synthesis Mobilization of Stored Carbohydrate Reserves in Cereals The Embryo Reserves The Endosperm Reserves Dissolution of the Endosperm and the Role of the Aleurone Layer The Fate of the Products of Starch Hydrolysis Mobilization of Stored Carbohydrate Reserves in Legumes Non-Endospermic Legumes Endospermic Legumes Mannan-Containing Seeds Other Than Legumes Stored Lipid Metabolism General Metabolism Fat Mobilization in Seeds The Fate of Glycerol and Fatty Acids The Glyoxysome The Synthesis and Degradation of Glyoxysomes Assimilation of the Breakdown Products Stored Protein Metabolism General Metabolism Protein Hydrolysis in Cereals Fate of the Liberated Amino Acids Protein Hydrolysis in Dicots

9 x Fate of the Liberated Amino Acids Proteinase Inhibitors.... Stored Phosphate Metabolism General Metabolism... Phosphate Metabolism in Cereals Phosphate Mobilization in Dicots. Mobilization of Nucleic Acids from the Storage Regions of the Seed.... Patterns of Reserve Mobilization in Seeds-Examples Some Articles of General Interest References Contents Chapter 7. Control Processes in the Mobilization of Stored Reserves Control Processes in Cereals Gibberellin and the Barley Aleurone Layer..... Gibberellins and a-amylase -" The a-amylase Story" Events During the Lag Period.... Membranes, Polysomes, and a-amylase Synthesis and Release of a-amylase a-amylase and Its Messenger RNA-Site of Action of GA? GA and a-amylase: Regulation in the Intact Grain Control of a-amylase Synthesis by the Products of Enzyme Hydrolysis Regulation of Other Hydrolases in the Barley Aleurone Layer GA-Induced Enzymes in Other Cereals 7.2 Control Processes in Other Seeds Control by the Embryo and Embryonic Axis The Mechanism of Axial and Embryonic Control Conclusions and Appraisals.. Some Articles of General Interest References Glossary and Index of English and Botanical Names. 283 Author Index Subject Index

10 Contents of Volume 2 Include: Viability and Longevity Physiological and Biochemical Parameters Dormancy Primary and Secondary Dormancy Embryo Dormancy and Immaturity Seed Coat Effects Light and Phytochrome High and Low Temperature Effects Effects of Chemicals (Gases, Inhibitors, Applied Growth Substances) Metabolism of Dormant Seeds The Breaking of Dormancy Possible Mechanisms of Action of Light, Chilling, After-Ripening, etc. Role of Endogenous Hormones and Hormonal Interactions Hormonal Action at the Molecular Level-Protein Synthesis, Nucleic Acids, Respiration Environmental Aspects of Germination and Dormancy Action and Interaction of Environmental Factors Genetic Aspects Water Stress The Lettuce Seed An Illustration of Dormancy Mechanisms in a Much-Researched Seed Author Index SUbject Index