Donald L. Smith. Chantal Hamel (Eds.) Physiology and Processes

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1 Donald L. Smith. Chantal Hamel (Eds.) Physiology and Processes Crop Yield

2 Springer Berlin Heidelberg New York Barcelona Budapest Hong Kong London Milan Paris Singapore Tokyo

3 Donald L. Smith. Chantal Hamel (Eds.) Crop Yield Physiology and Processes With 103 Figures Springer

4 Professor DONALD L. SMITH Plant Science Department Professor CHANTAL HAMEL Natural Resourch Department Mc Gill University Macdonald Campus 21, III Lakeshore Road Ste. Anne de Bellevue QC, Canada H9X 3V9 ISBN Springer-Verlag Berlin Heidelberg New York Library of Congress Cataloging-in-Publication Data Crop yield: physiology and processes I Donald L. Smith, Chantal Hamel (eds.). p. cm. Includes bibliographical references and index. ISBN (hardcover: alk. paper) 1. Crops - Physiology. 2. Crop yields. I. Smith, Donald L., II. Hamel, Chantal, SB1l2.5.C '58 - dc21 This work is subject to copyright. All rights reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9,1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. Springer-Verlag Berlin Heidelberg 1999 The use of general descriptive names, 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. Product liability: The publisher cannot guarantee the accuracy of any information about dosage and application thereof contained in this book. In every individual case the user must check such information by consulting the relevant literature. Typesetting: Best-set Typesetter Ltd., Hong Kong Cover Design: Design & Production GmbH, Heidelberg SPIN / Printed on acid-free paper

5 Preface This book has been prepared for those seeking a better understanding of the functioning of crop plants, particularly the processes that lead to the generation of products valued by human beings. The contributors, who are among the world's foremost experts on the important crops upon which humanity depends for food or fibre, address the relevant processes for their specific crop. Currently, the world population is continuing to increase. It is projected to plateau around the middle of the next century, and while there is considerable controversy regarding the population level when this plateau is achieved, most estimates are in the area of At present, there are about people in the world who do not have secure access to food. Over the last 50 years various aspects of agricultural research have been combined to increase the output of world crops approximately 2.5-fold. Given the need to feed the increasing population, and to provide better access, it is predicted that during the next 50 years the agricultural research community must repeat this achievement. Molecular genetics provides us with a wealth of information regarding a wide range of organisms. Currently, genome sequencing of new organisms is being completed at a rate of about one organism every 2 weeks. Recently, sequencing of the major cereal crops, wheat, rice and maize, has been undertaken. It seems certain that before long all of the world's major crops will be sequenced. The next great challenge will be to understand how all the proteins coded for by the genes in the genome combine to make up the physiology of each crop species. This begins at the cellular level, and moves to the whole plant and ecophysiology level to have its greatest impact. We are in the golden age of biology, and we will soon be able to acquire all the information at any level of biological organization. This will be at the genome level initially, progressing soon to the proteome and physiology levels. This is, and will continue to be, a time of great challenge and great opportunity for those of us working in agricultural research. Thus, physiology will play a key role in the coming decades.

6 Physiological Control of Growth and Yield in Wheat: Analysis and Synthesis R.K.M.Hay 1 Introduction Components of Yield Wheat Development Leaf, Spikelet and Floret Initiation at the Mainstem Apex Leaf Appearance and Expansion; Tiller Development Quantitative Aspects of Development: Canopy Generation Germination and Crop Emergence Leaf Appearance Final Leaf Number Leaf Expansion Leaf Senescence Tillers Leaf Area Index Leaf (Green) Area Index and the Interception of PAR Radiation-Use Efficiency Photosynthetic Efficiency of Single Leaves and Canopies Radiation-Use Efficiency of Wheat Canopies Variation in Radiation-Use Efficiency: Stress and Ontogeny Quantitative Aspects of Development: Generation of Grain Population Density Simulation of Organ Survival Simulation of Grain Yield and its Components Conclusions References Growth and Development of Oat with Special Reference to Source-Sink Interaction and Productivity P. Peltonen-Sainio 1 Oat as a Subject for Crop Physiological Studies Phenostages and Phenophases of Oat with Agricultural Interest Phenostages Phenophases... 44

7 VIII 3 Canopy Architecture and Function in Relation to Dry-Matter Production Early Canopy Closure and Achievement of Optimum LAI During Formation of Yield Potential Maintaining Efficient Interception of PAR During Grain Growth Vegetative Organs as Competing Sinks During Formation and Realization of Yield Potential Root Growth and Function: The Hidden Component of Crop Physiology Tillers: Waste or Economic Reservoirs of Assimilates? Stem Elongation and Effects of Plant Height on Assimilate Competition Grain Filling as a Final Process Affecting Realization of Yield Potential Increasing Grain-Filling Rate Increasing Harvest Index? The Need for Description of Oat Ideotypes? References Barley: Physiology of Yield D.L. Smith, M. Dijak, P. Bulman, B.L. Ma, and C. Hamel 1 Apex Development and Contribution to Yield Inflorescence Development Structure of the Barley Spike Inflorescence Differentiation Factors Affecting Apical Development Yield Components Tillering and Tiller Contribution to Grain Yield Tillering Tillering and Yield Tillering: Hormonal Control and Plant Growth Regulators Grain Filling Carbon Metabolism Photosynthesis and Radiation Use Efficiency Water Use Efficiency Harvest Index Dry Matter Partitioning Nitrogen Metabolism Nitrogen Uptake Nitrate and Nitrite Reductase Activity Nitrogen Assimilation Senescence Amino Acid Translocation... 85

8 IX 5.6 Pattern of N Uptake Grain Nitrogen Protein Synthesis and Amino Acid Composition Contribution of Preanthesis Carbon and Nitrogen to Yield Retranslocation of Carbon During Grain Filling Retranslocation of Nitrogen During Grain Filling Phosphorus Uptake and Mycorrhizal Associations Potassium Uptake Plant Lodging and Growth Regulators Conclusions References Rice A.R. Sharma and D.P. Singh Introduction.... Growth and Development.... Vegetative Growth.... Reproductive Growth.... Photosynthesis and Dry-Matter Production.... Leaf Photosynthetic Rate.... Leaf Orientation.... Leaf Area.... Varietal Differences.... Environmental Control of Growth and Yield.... Rainfall.... Drought.... Submergence.... Adaptation to Prolonged Submergence.... Submergence Tolerance.... Temperature.... Light.... CO2 Concentration.... Relative Humidity and Wind Velocity.... Yield Determining Processes..... Storage of Assimilates.... Determination of Yield Capacity.... Partitioning and Translocation of Assimilates.... Contribution of Pre- and Post-Flowering Carbohydrates to Grain Yield.... Source-Sink Relationship.... Analysis of Yield Components.... Spikelet Number.... Percentage of Filled Spikelets.... Sterility

9 x Grain Weight Response to Cultural Factors Method of Crop Establishment Nitrogen Fertilization Improving Yield Potential Estimation of Yield Potential Physiological Traits for Yield Improvement Increasing Biomass Production Increasing Harvest Index Improving Sink Capacity Improving Ripening Percentage Yield Potential and Nitrogen Requirement New Plant Types Plant Type for Irrigated Ecosystems Reduced Tillering and Large Panicles Grain Size and High-Density Grains Canopy and Leaf Characteristics Short and Stiff Culm Crop Growth Duration Plant Types for Rain-Fed Ecosystems Present Status of New-Plant Type Development Hybrid Rice Biotechnological Approaches Future Directions References Physiology of Maize M. Tollenaar and L.M. Dwyer 1 Introduction History, Production, and Utilization Taxonomy and Morphology Agronomy and Physiology Phenology Phasic Development Relative Maturity and Rate of Development Dry Matter Accumulation Incident Solar Radiation Absorption of Solar Irradiance Leaf Area Index Canopy Extinction Coefficient Radiation-Use Efficiency and Total Dry Matter Accumulation Dry Matter Partitioning

10 Xl 4.1 General Source-Sink Relationships Source Strength Export from Leaves Temporary Storage Competing Demands for Nitrogen Genetic Improvement Appendix References Leaf Expansion and Phenological Development: Key Determinants of Sunflower Plasticity, Growth and Yield V.O. Sadras and N. Trapani 1 Introduction Phenotypic Plasticity Yield Responses to Winter Sowing Yield Responses to Plant Population Leaf Area Leaf Area, Light Interception and Crop Growth Leaf Area Components Responses of Leaf Expansion to Water Availability Growth and Water Relations of Plants in Controlled Environments Leaf Expansion in Field-Grown Sunflower Effects of Nitrogen on Growth and Development Effects of Nitrogen on Leaf Area Leaf Expansion Rate as Affected by Nitrogen Availability: Comparison with Rate of Photosynthesis Phenological Development Phenological Pattern and yield Physiological Basis and Modelling of Phenological Development Sowing to Emergence Emergence to Anthesis Anthesis to Physiological Maturity Conclusions References Cotton: Factors Associated with Assimilation Capacity, Flower Production, Boll Set, and Yield J.J. Heitholt 1 Introduction General Characteristics of Yield Relative Importance of Yield Components

11 XII 2 Root and Canopy Development Morphological Developmental Stages Solar Radiation Interception Canopy Architecture COz-Assimilation Capacity Canopy CO 2-Exchange Rate Single-Leaf CO 2-Exchange Rate Fruiting-Form CO 2-Exchange Rate Fruiting-Form Formation Floral Bud (Square) and Boll Retention Physiology of Boll Retention/Shed Flower Production and Yield Fruit Loss and Yield Compensation Importance of Fruiting Positions Carbohydrate Metabolism and Utilization Leaf Assimilate Physiology Fiber Assimilate Physiology Environmental Effects Temperature Nutrition CO 2- Enrichment and Yield Monitoring and Management Growth Regulators Crop Termination Breeding and Yield Physiology Research Challenges Treatment and Genotype Comparisons Management Intensity Summary Glossary of Cotton Yield Physiology Terms References Jute P. Palit 1 Introduction Seed: Viability and Germination Seedling Growth and Initial Field Operations Root System Leaf Area, Light Interception and Canopy Photosynthesis CO 2 Metabolism and Carbon-Use Efficiency Assimilate Translocation and Partitioning Photoperiod and Flowering Improvement of Jute Productivity: Problems and Prospects References

12 XIII Sugarcane S.E. Lingle 1 Introduction Economic Importance Botanical Description Anatomy and Morphology Growth and Yield Dynamics Stand Establishment Growth and Sugar Accumulation Sucrose Synthesis and Partitioning Photosynthesis Translocation and Partitioning Stem Anatomy and Sucrose Storage Sucrose Metabolism Conclusion: Prospects for Increasing Sucrose Yield References Sugar Beet T.R. Thomas 1 Introduction Crop Establishment and Early Development Seedling Establishment Identifying Seed Vigour Improving Seed Performance Manipulating Seed Production Leaf Growth and Development Fibrous Root Development Root Growth in the Soil Root-to-Shoot Ratios Growth and Development of the Storage Root Anatomical Development of the Storage Root Sucrose Accumulation Photosynthesis and Growth Carbon Fixation in the Leaf Light and Photosynthesis Modification of Photosynthetic Efficiency... 32l 7 Water Relations and Stress Control of Water Uptake Stress Tolerance Assimilate Partitioning Assimilate Movement in the Plant Assimilate Storage in the Root Modifying Sucrose Storage Capacity...,

13 XIV 9 Crop Growth and Sucrose Accumulation Sugar Beet and Climate Change Sugarbeet in the Future References Potato J. Vos 1 Introduction The Basic Plan of the Potato Plant Physiological Age and Growth Vigour Interrelations Between Numbers of Component Plant Parts The Process of Tuber Formation Rate and Duration of Development Processes Radiation Interception, Growth Rate and Total Dry Matter Production Temperature and the Rate of Growth Processes Dry Matter Distribution Nitrogen Nutrition and Potato Production Drought and Potato Production Second Growth Tuber Size Distribution References Cowpea A.E. Hall Introduction Grain Yield in Optimal Environments.... Grain Yield Under Drought.... Photoperiod and Temperature Effects on Phenology and Responses to Stress.... Conclusions References Soybean [Glycine max (1.) Merr.] Physiology and Symbiotic Dinitrogen Fixation F. Zhang and D.L. Smith 1 Origin and Introduction Plant Development Germination and Vegetative Stages Reproductive Stage Plant Physiology

14 xv 3.1 Carbon Assimilation and Metabolism Water Requirement and Water Stress Mineral Nutrition and Metabolism Symbiotic Dinitrogen Fixation Dinitrogen Fixation Nodule Formation Recognition Between Symbiotic Partners Biochemistry and Physiology General Genetic Characteristics Factors Affecting Soybean Dinitrogen Fixation References Physiological Control of Growth and Yield in White Clover J.R. Caradus and M.J.M. Hay 1 Growth Form of White Clover Clonal Growth Exceptions to Clonal Growth Plagiotropic Habit Flowering Physiological Control of Growth Environmental Effects Temperature Light Nutrients Nitrogen Fixation Moisture Physical Impedance Defoliation Biotic Influences Ozone and Ultraviolet Light Carbon Dioxide Concentration Correlative Effects Age of Phytomer Presence of Root, Leaf, Flower or Branch at a Phytomer Genotype References Physiological Control of Alfalfa Growth and Yield J.J. Volenec 1 Introduction Alfalfa Yield Component Analysis Factors Influencing Alfalfa Yield Components Alfalfa Persistence Mass per Shoot

15 XVI 3.3 Shoots per Plant Conclusions and Future Prospects References Physiological Control of Forage Grass Yield and Growth A.R. Overman and D.M. Wilson 1 Introduction Dynamic Models Probability Extended Probability Phenomenological Seasonal Models Logistic Extended Logistic Multiple Logistic Extended Multiple Logistic Harvest Interval and Water Availability Examples Warm-Season Perennial Warm-Season Annual Cool-Season Annual Harvest Interval Summary References Subject Index

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