Hans Mohr Peter Schopfer Plant Physiology

Hans Mohr Peter Schopfer

Plant Physiology Translated by Gudrun and David W Lawlor

With 698 Figures and 144 Tables

Springer

Professor Dr. HANS MOHR Professor Dr. PETER SCHOPFER

Biologisches In~titut II der UniversiHit Lehrstuhl fUr Botanik Schanzlestraf3e 1 79104 Freiburg

Translated by:

GUDRUN LAWLOR South Bank University Wandsworth Road London SW8 2JZ, UK

Dr. DAVID W LAWLOR Biochemistry & Physiology Department Rothamsted Experimental Station Harpenden, Herts AL5 2JQ, UK

Title of the original German edition: Hans Mohr, Peter Schopfer Pflanzenphysiologie, 4. Auflage (Springer-Lehrbuch)

ISBN-13: 978-3-642-08196-5 e-ISBN-13: 978-3-642-97570-7 DOl: 10.1007/978-3-642-97570-7

Library of Congress Cataloging-in-Publication Data. Mohr, Hans, 1930- [Lehrbuch der Pflanzenphysiolo­gie. English] Plant physiology! Hans Mohr, Peter Schopfer; [translated by Gudrun and David W. Lawlor]. p. cm. Includes bibliographical references (p.) and index. ISBN 3-540-58016-6 (Berlin). - ISBN 0-387-58016-6 (New York) 1. Plant physiology. I. Schopfer, Peter, 1938-. II. Title. QK711.2.M6413 1994 581.1-dc20 94-21765

This work is subject to copyright. All rights are 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 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 permissions for use mnst always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law.

© Springer-Verlag Berlin Heidelberg 1995 Softcover reprint of the hardcover 1 st Edition 1995

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.

Translators' Preface

Translation of the fourth edition of this well-established, comprehensive and detailed plant physiology textbook will, we believe, provide English-speaking plant scientists with a valuable source for teaching and learning, particularly for the later stages of the first degree and above.

We have striven to retain the clarity and subtlety of the German text and to provide a translation of technical terms and concepts which does not grate on the English ear nor upset American readers.

We thank Prof. Dr. H. Mohr, Mrs. E. Janssen and particularly Prof. Dr. P. Schopfer for their help and extremely valuable and constructive criticism of drafts of the translation. Also, Springer-Verlag has been very patient and helpful. Responsibility for the quality of the English text, however, rests with us.

Harpenden, Autumn 1994 GUDRUN and DAVID W LAWLOR

Contents

1 Setting the Aims in Physiology . Concept of Water Potential in the Water Relations of Cells 45 Concepts of Physiology. 1

Chemical Potential of Ions 51 Heterogeneity of Physiology 1 Membrane Potential 52 Limitations of Reductionism 2 Energetics of Biochemical Reactions 54

Laws in Biology 3 Phosphate Transfer Universal Laws in Physiology 3 and Phosphorylation Potential 57 System Theory 4 Redox Systems and Redox Potential 57 Important Concepts 5 Further Reading . 62 Biochemistry and Physiology 5

Further Reading . 6 5 The Cell as a Metabolic System 63

2 Theoretical Basis of Physiology 7 Biological Catalysis 63 Metabolic Compartmentation of the Cell 68

Principles of Scientific Studies . 7 Transport Mechanisms in Biomembranes 69 Reference Units 7 ATP Synthesis by Energy-Transforming The Principle of Causality in Physiology 8 Biomembranes 73 Single-Factor Analysis 8 Uptake of Materials by the Cell . 75 Multifactor Analysis 9 Principles of Metabolic Regulation 81 The Problem of Complexity. 11 Further Reading . 85 Formulation of Laws. 14 Characters and Variability 16 Presentation of Data . 18 6 The Cell as a Dividing System . 87 The Problem of Extrapolation. 19 Regulation of the Rate of Mitosis 90 Further Reading . 19 Determination of the Division Plane 91

Cell Cycle and Cell Differentiation. 92

3 The Cell as a Morphological System 21 Further Reading . 92

The Meristematic Plant Cell 21 The Mature Plant Cell 31 7 The Cell as a Polar System 93 The Woody Plant Cell 34 Significance of Cell Polarity . 94 Further Reading . 37 Induction of Polarity by Light 95

Induction of Polarity by Polarised Light 95 Polarity and the Bioelectrical Field 96 4 The Cell as an Energetic System . 39 Polarity and Signalling Substances 97

The First Law of Thermodynamics 39 Further Reading . 97 The Second Law of Thermodynamics 39 The Cell as an Open System,

8 The Cell as a Growing System . 99 Dynamic Equilibrium 41 Chemical Potential . 42 Biophysical Basis of Cell Growth 99 Chemical Potential of Water 43 Growth and Cell-Wall Structure . 101

VIII Contents

Extension Growth of Multicellular Organs 106 Regulation of Extension Growth 108 Further Reading. . . . . . . . . 109

9 The Cell as an Oscillatory System .. 111

The Original Observation: Daily Leaf Movements of Bean Plants 111

Further Selected Phenomena Illustrating Circadian Rhythms . 112

Selected Experiments to Analyse Endogenous Rhythms 116

Endogenous Rhythm as a System Characteristic 119

Further Reading. . . . . . . . 120

10 The Cell as a Gene-Physiological System. 121

Chromatin and Chromosomes . RNA of the Cell . Protein Synthesis ..... . Specific Inhibitors . . . . . . Folding and Assembly of Proteins Sorting Proteins . . . . . . . Regulation of Gene Expression Further Reading. . . . . . .

121 124 126 127 128 129 129 132

11 Intracellular Morphogenesis 133

Intracellular Communication . 133 Morphogenesis of Mitochondria . 136 Morphogenesis of Plastids 138 Morphogenesis of Peroxisomes (Microbodies) 144 Further Reading. . . . . . . . . . . . . 147

12 Photosynthesis as a Chloroplast Function 149

Photosynthesis as Energy Conversion 149 Energy Conversion in the Chloroplast 152 Pigment Systems of Red

and Blue-Green Algae . . . . . 166 Photosynthetic Electron Transport 168 Protection Mechanisms Against

Photo-Oxidative Destruction of the Photosynthetic Apparatus 173

Mechanism of Photophosphorylation 175 Biochemical Processes 176 Anoxygenic Photosynthesis

of Phototrophic Bacteria 184 Further Reading. . . . . 185

13 Respiratory Metabolism . . . . . .. 187

Catabolic Energy Production . . . . .. 187 Respiratory Metabolism of Carbohydrates 188 Photorespiration. . . . . . . . 198 Mobilisation of Storage Material

in Storage Tissues . . . . . . 203 Regulation of Respiratory Gas Exchange. 210 Regulatory Interaction Between Synthesis

and Breakdown of Carbohydrates 222 Further Reading. . . . . . . . . .. 223

14 The Leaf as a Photosynthetic System. 225

Measurement of the Rate of Photosynthesis 226 Gross and Net Photosynthesis. . . .. 227 Limiting Factors of Net Photosynthesis 229 Ability of Leaves to Adapt

Photosynthetically . . 232 Temperature Dependence

of Net Photosynthesis 234 Influence of Oxygen on Net Photosynthesis. 236 Regulation of CO2 Exchapge by Stomata 237 Further Reading ............. 243

15 C4 Plants and CAM Plants 245

C4 Syndrome . . . . . . . . 245 C4 Dicarboxylate Cycle. . . . 249 Ecological Aspects of the C4 Syndrome 251 CAM, an Alternative to C4 Photosynthesis 253 Isotope Discrimination During CO 2 Fixation 256 Further Reading. . . . . . . . . . .. 257

16 Metabolism of Water and Inorganic Ions 259

Water ........ . Mineral Nutrition of Plants . . Essential Microelements Function of Nutrient Elements

in Metabolism. . . . . . . Salt Excretion in Halophytes . Sequestration of Heavy Metals

by Phytochelatin . Further Reading. . . . . . .

17 Ecological Cycles of Materials and Energy ..... .

Carbon and Oxygen Cycles Nitrogen Cycle .....

259 261 262

263 265

266 267

269

269 271

Energy Flux. . . Further Reading .

18 Biosynthetic Metabolism . . . .

Primary and Secondary Metabolism Shikimate Pathway Biogenesis of Chlorophyll . Further Reading. . . . .

273 274

275

275 281 282 284

19 Physiology of Development 285

Basic Aspects . 285 Growth. . . . . . . . . . 293 Differentiation. . . . . . . 301 Pattern Formation and Morphogenesis. 310 Tumour Formation in Plants . 321 Morphogenesis of Acetabularia 324 Further Reading. . . . . . . 330

20 Physiology of Sexuality . . . . . . . . 333

Five Case Studies of Gametogenesis and the Effects of Pheromones in Lower Plants . . . . . . . . 333

Fertilisation in Flowering Plants. . 340 Maternal and Paternal Inheritance. 343 Male Sterility . . 343 Further Reading. . . . . . . . . 343

21 Photomorphogenesis . 345

Action Spectra 345 Pigments . . . 346 Phytochrome . 347 Mode of Action of Phytochrome

in Photomorphogenesis. . . . 352 Four Case Studies on the Effects

of Phytochrome . . . . . . . 356 Cooperation Between Photosensors 364 A Positive UV-B Effect: Synthesis of Flavone

Glycosides in Cell Suspension Cultures 367 Photomorphogenesis of Fungi. 368 Further Reading. . . . . . . . . .. 373

22 Development of Chloroplasts . .

Regulation of Chlorophyll Synthesis Formation of Holocomplexes ...

375

375 376

Contents IX

Regulation of Matrix Enzymes The Plastid Factor . . . . . . Formation of the Apparatus of Nitrate

Assimilation During the Development of Chloroplasts

Further Reading. . . . . . . . . . .

23 Physiology of Hormone Action . . .

Overview of the Structure and Function

378 379

379 381

383

of Phytohormones . . . . . . . 386 Six Case Studies of the Physiology

of Hormone Action 399 Further Reading. . . . . . . . . 408

24 Ripening and Germination of Reproductive and Distributive Organs . . 409

Development of the Ripe Seed 409 Germination of Ripened Seeds 412 Regulation of Gene Expression

During Embryo Development 418 Regulation of Fruit Development

by the Seed . . . . . . . . . 419 Bud Dormancy and Bud Germination 419 Further Reading. . . . . . . . . . 422

25 Flower Formation and Photo-and Thermoperiodism . . . .

Flower Induction and Morphogenesis Flower Formation and Florigen . . Photoperiodism . . . . . . . . . Grafting Experiments and Florigen Flower Formation and Gibberellin Phytochrome and Photoperiodism . Photoperiodism and Circadian Rhythm Photoperiodic Phenomena Independent

of Flower Formation. . . . Significance of Photoperiodism Thermoperiodism Vernalisation Further Reading .

26 Physiology of Senescence

Monocarpic Plants. . . . . Aging of Leaves in Perennial Plants Further Reading . . . . . . . . .

423

423 424 426 428 430 431 432

432 434 434 435 436

437

437 438 442

X Contents

27 Physiology of Regeneration and Transplantation . . . . 443

Results from Organ Cultures . 443 A Technical Note: Tissue Cultures 445 Proof of the Omnipotence of Specialised Cells 445 Parasexual Hybridisation . . . 450 Wound Healing . . . . . . . 452 Cooperation of Several Factors

During Regeneration. . 453 Regeneration Experiments

with Flower Formation. 454 Intracellular Regeneration 454 Transplantation . 455 Chimaeras 456 Further Reading . 458

28 Effects of Ionising Radiation

Exciting and Ionising Radiation Types of Ionising Radiation. . Process of Ionisation . . . . . Some Thoughts on the Target Theory Effect of Ionising Radiation

on Cell Components . . . . . . . Repair of Radiation Damage in DNA Effect of Ionising Radiation

on Higher Level Organisation in Cells Further Reading. . . . . . . .

29 Physiology of Xylem Transport

The Transport Pathway from Perirhizal Zone into Root Vessels

Root System and Root Hairs Two Plant Transport Systems Nitrogen in the Xylem Sap . Transpiration and Ion Supply Water Balance ... . Pathways ..... . Classical Experiments Transpiration . . . . Analogue Model for Water Transport

in a Plant ........... . Distribution of Water Potential in a Tree Guttation and Root Pressure Further Reading. . . . . . . . . . .

459

459 459 459 461

462 463

464 466

467

467 469 469 471 473 473 474 476 478

482 484 485 486

30 Physiology of Phloem Transport

Basic Aspects . . . Pathways ........... . Transport Molecules . . . . . . . Mechanism of Sieve Tube Transport Further Reading. . . . .

31 Physiology of Movement

Free Movement of Plants Between Sites Phototropism . . . . . . . . Gravitropism ....... . Further Processes of Movement Further Reading. . . . . . .

32 Physiology of Stress Resistance

487

487 489 491 492 495

497

497 504 516 524 537

539

Water Stress. . . . 540 Temperature Stress. . . . . . 544 Light and UV Stress . . . . . 551 Biogenic Stress (Plant Diseases) 558 Further Reading. . . . . . . 565

33 Physiology of Crop Production 567

Basic Aspects . . . . . . . . 567 Formation of Storage Material 570 Production Factors ..... 571 Principles of Yield Production. 571 Optimum of a Production Factor: Nitrogen 573 Suppression of Yield Antagonists:

Herbicides . . . . . . . . . . 579 Synthetic Growth Retardants . . . 583 Improvement of Hereditary Factors 584 Gene Technology . . . . . . . . 586 Case Study: Antibody Production in Plants 588 Further Reading. . . . . . . . . . 589

Appendix: Physical Quantities, Units, Conversion Factors and Constants 591

References 595

Subject Index . . . . . . . . . . . . . . 603