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PLANT PHYSIOLOGY Molecular, Biochemical, and Physiological Fundamentals of Metabolism and Development
PLANT PHYSIOLOGY
Molecular, Biochemical, and Physiological Fundamentals of Metabolism and Development
PLANT PHYSIOLOGY
Molecular, Biochemical, and Physiological Fundamentals of Metabolism and Development
Dieter Hess
Springer-Verlag Berlin • Heidelberg· New York 1975
DIETER HESS
Professor, Botanische En twicklungsphysiologie Universitaet Hohenheim Postfach 106 D-7000, Stuttgart 70 West Germany
Library of Congress Cataloging in Publication Data
Hess, Dieter. Plant Physiology
Translation of Pflanzenphysiologie. Bibliography: p. 1. Plant physiology. I. Title
QK711.2.H4713 581.1 73-22278
All rights reserved.
No part of this book may be translated or reproduced in any form without written permission from Springer-Verlag.
© 1975 by Springer- Verlag New York Inc.
ISBN-13: 978-3-540-06643-9 e-ISBN-13: 978-3-642-80813-5 DOl: 10.1007/978-3-642-80813-5
To my mother
PREFACE
In recent years, molecular biology has infiltrated into all branches of botany. This is particularly true of plant physiology. This book attempts to provide an introduction to the metabolic and developmental physiology of higher plants from a molecular biological point of view. Starting from the heterocatalytic function of DNA the first ten chapters deal with metabolism; development is presented in the last nine, starting from the autocatalytic functions of DNA and including certain topics oriented more toward metabolic physiology. Both fields of plant physiology are so closely linked that an in tegrated presen tation of this kind seemed not only possible but desirable.
In contrast to other accounts, an attempt has been made to give equal weight to metabolism and development. In particular, the so-called "secondary" plant materials, which are of considerable interest to the pharmacist, the nutrition technologist, the plant breeder, and the agriculturalist, as well as to the biologist, are treated sufficiently. It is obvious that the wealth of material made an illustrative style of presentation necessary.
The book is intended for beginners, and so it has had, in part, to be simplified. Even so it has not been possible to write it without mentioning hypotheses that anticipate much more research. The beginner ought also to learn how working hypotheses are first postulated on the basis of certain facts and then must either be proved or refuted.
In this connection it may be asked how much the "beginner" is expected to know. He should be provided with a good textbook of general botany as well as an introduction to biochemistry. The latter is all the more necessary because of the material that is treated in every textbook of biochemistry, such as biological oxidation, only the basic concepts are presented here. In compensation for this the metabolic processes that are specific for higher plants have been given more than usual emphasis.
Apart from brief references, an outline of methodology had to be omitted in the interests of the size and price of this book. Information of this nature can, however, be found in any biochemically oriented textbook and there are also abridged introductory texts, e.g. E. S. Lenhoff's Tools of Biology.
VII
viii Preface
The author hopes that his book will be of use not only to students of biology and related disciplines at the beginning of their studies but also to teachers at the high school and grammar school level. They may wish to keep themselves informed of recent developments in the field of plant physiology and also to take this or that fact into account in teaching.
The author wishes to thank his publisher, Herrn Roland Ulmer, and his staff for their kind cooperation and Herrn Ekkehart Yolk for his care in preparing the figures. Thanks are also due to his wife and daughter who showed full understanding of the many hours of "overtime" the book entailed.
It is a real pleasure to thank Dr. Derek Jarvis for the English translation and for his efforts in reading the galleys and finishing the index. Should the book be well accepted in the English speaking countries, it will be due to his interest and cooperation.
The German editions of this book received a warm reception from the reading public. It is the author's hope that the same may prove true of the English edition.
Stuttgart-Hohenheim Dieter Hess
CONTENTS
Preface ....................................................... vii
CONTROL OF CHARACTER FORMATION BY NUCLEIC ACIDS 1
A. The Chemical Constitution of the Nucleic Acids. . . . . . . . . . . I 1. The Building Blocks of the Nucleic Acids. . . . . . . . . . . . . . 1 2. Nucleosides, Nucleotides, Polynucleotides. . . . . . . . . . . . . . 4 3. The Watson-Crick Model of DNA..................... 5
B. Direct Evidence for the Role of the Nucleic Acids as Carriers of Genetic Information. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 1. Transformation...................................... 7 2. Transfection ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
C. The Heterocatalytic Function of DN A: Transcription and Translation. .. ......................................... 10 1. The Concept of Molecular Genetics. . . . . . . . . . . . . . . . . . . 10 2. The Genetic Code. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . II 3. Transcription........................................ 12 4. Translation.......................................... 14 5. Antimetabolites of Transcription and Translation. . . . . . . 20 6. Evidence for mRNA in Higher Plants. . . . . . . . . . . . . . . . . 24 7. Transcription and Translation in a Cell-Free System. . . . . 29 8. One Gene-One Polypeptide. . . . . . . . . . . . . . . . . . . . . . . . . . . 30
PHOTOSYNTHESIS 35
A. Division of Photosynthesis into Primary and Secondary Processes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
B. Primary Processes of Photosynthesis. . . . . . . . . . . . . . . . . . . . . . 36 I. Electron Transport Chains. . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 2. Redox Systems in the Primary Processes of Photosynthesis 38 3. Pigment Systems I and II of Photosynthesis. . . . . . . . . . . . 43 4. Primary Processes of Photosynthesis. . . . . . . . . . . . . . . . . . . 46 5. Quantum Yield of Photosynthesis. . . . . . . . . . . . . . . . . . . . . 48
ix
x Contents
C. Secondary Processes of Photosynthesis. . . . . . . . . . . . . . . . . . . . 49 1. The CO2 Acceptor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 2. The Connection with the Primary Processes. . . . . . . . . . . . 50 3. The Calvin Cycle.................................... 51 4. The C4 dicarboxylic Acid Pathway. . . . . . . . . . . . . . . . . . . . . . 53
D. The Chloroplast: Site of Photosynthesis. . . . . . . . . . . . . . . . . . . 54
CARBOHYDRATES 57
A. Monosaccharides. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 l. Phosphorylation (Kinases) .. . . . . . . . . . . . . . . . . . . . . . . . . . . 57 2. Intramolecular Migration of Phosphate (Mutases). . . . . . . . 58 3. Sugar Nucleotides (UDPG). . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 4. Inversion of an OH Group (Epimerases). . . . . . . . . . . . . . . . 59 5. Control of the Equilibrium Between Aldoses and Ketoses
(Isomerases). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 6. Oxidative Degradation of 1 C Atom (Hexose-pentose
Transition) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 7. The Pentose Phosphate Cycle. . . . . . . . . . . . . . . . . . . . . . . . . 62
B. Oligosaccharides and Polysaccharides .... . . . . . . . . . . . . . . . . . 63 1. Glycosides.......................................... 63 2. Oligosaccharides..................................... 64 3. Polysaccharides...................................... 67
BIOLOGICAL OXIDATION 74
A. Glycosis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
B. Oxidative Decarboxylstion of Pyruvate, Formation of Active Acetate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
C. Citric Acid Cycle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
D. The Respiratory Chain. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
E. Mitochondria as Power Plants. . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
FATS 89
A. Chemical Constitution of the Fatty Acids. . . . . . . . . . . . . . . . . 89
B. Biosynthesis of the Fatty Acids. . . . . . . . . . . . . . . . . . . . . . . . . . 90 1. Formation of Malonyl CoA . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 2. Fatty Acid Synthesis Proper. . . . . . . . . . . . . . . . . . . . .. . . .. 91
C. Biosynthesis of the Neutral Fats. . . . . . . . . . . . . . . . . . . . . . . . . 93
Contents xi
D. Degradation of the Fats. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 1. ,B-Oxidation......................................... 95 2. a-Oxidation......................................... 96
E. The Glyoxylate Cycle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
TERPENOIDS 99
A. Chemical Constitution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
B. Secondary Plan t Substances. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 100
C. Volatile Oils. . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 101
D. Biosynthesis (General). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 102
E. Biosyn thesis (Particular) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 104 1. Monoterpenes....................................... 104 2. Sesquiterpenes....................................... 105 3. Triterpenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 106 4. Diterpenes.......................................... 110 5. Tetraterpenes: Carotenoids .... . . . . . . . . . . . . . . . . . . . . . .. III 6. Polyterpenes........................................ 115
PHENOLS 117
A. Chemical Constitution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 117
B. Biosynthesis (General). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 118 1. The Shikimic Acid Pathway ........................... 118 2. The Acetate-Malonate Pathway. . . . . . . . . . . . . . . . . . . . . .. 120 3. Precursors and Intermediates. . . . . . . . . . . . . . . . . . . . . . . .. 120
C. Biosynthesis (Particular) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 121 1. Cinnamic Acids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 121 2. Coumarins.......................................... 122 3. Lignin.............................................. 124 4. Phenol Carboxylic Acids and Simple Phenols. . . . . . . . . .. 128 5. Flavan Derivatives. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 129 6. Flower Pigmentation. . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . .. 136
AMINO ACIDS 138
A. The Reduction of Nitrogen. . . . . . . . . . . . . . . . . . . .. . . . . . . . .. 138
B. Reductive Amination .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 140
C. The Formation of Glutamine. . . . . . . . . . . . . . . . . . . . . . . . . . .. 140
D. Transamination ............ ,............................ 141
E. The Origin of the C Skeleton of the Amino Acids. . . . . . . . .. 142
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ALKALOIDS 144
A. Derivatives of the Aliphatic Amino Acids, Ornithine and Lysine ................................................ , 146 1. Quinolizidine Alkaloids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 146 2. Nicotiana Alkaloids and Nicotinic Acid. . . . . . . . . . . . . . .. 147 3. Tropane Alkaloids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 149
B. Derivatives of the Aromatic Amino Acids, Phenlalanine and Tyrosine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 150 1. Amaryllidaceae Alkaloids and Colchicine... ............ 150 2. Betacyanins and Betaxanthins. . . . . . . . . . . . . . . . . . . . . . . .. 152 3. Isoquinoline Alkaloids (Benzylisoquinoline Alkaloids) . . .. 153
C. Derivative of the Amino Acid Tryptophan: Indole Alkaloids and Derivatives......................................... 154
D. Purine Alkaloids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 156
E. Biochemical Systematics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 157
A.
B.
PORPHYRINS
CELL DIVISION
Development- Growth and Differen tiation
Cell Division .......................................... . 1. The Mitotic Cycle .................................. . 2. The Autocatalytic Function of ON A: Replication ...... . 3. Plant Tumors: Crown Galls .......................... .
DIFFERENTIAL GENE ACTIVITY AS PRINCIPLE OF
160
163
163
163 163 165 169
DIFFERENTIATION 172
A. Totipotency. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 172
B. Differential Gene Activity: The Phenomenon. . . . . . . . . . . .. 174 1. RNA Synthesis on Giant Chromosomes. . . . . . . . . . . . . . .. 175 2. Phase-specific mRNA ............................... 176 3. Phase- and Tissue-specific Protein Patterns. . . . . . . . . . .. 178
REGULATION 181
A. States of Activity of the Gene............................ 181
Contents xiii
B. Regulation: Point of Departure. . . . . . . . . . . . . . . . . . . . . . . . . .. 182
C. Regulation by Internal Factors........................... 183 1. Intracellular Regulation.............................. 183 2. Intercellular Regulation: Phytohormones............... 194
D. Regulation by External Factors. . . . . . . . . . . . . . . . . . . . . . . . . .. 219 1. Temperature........................................ 219 2. Light ............................................... 220
POLARITY AND UNEQUAL CELL DIVISION AS FUNDAMENTALS OF DIFFERENTIATION 225
A. Polarity. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 225
B. Unequal Cell Division. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 227
A.
B.
1. Development of Stomata. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 228 2. Root Hair Formation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 228 3. Pollen Mitosis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 229
CELL ELONGATION 231
The Phenomenon 231
The Process of Elongation Within a Cell. . . . . . . . . . . . . . . . .. 232 1. The Suction Pressure Equation of the Cell. . . . . . . . . . . . .. 232 2. The Stages of Cell Elongation. . . . . . . . . . . . . . . . . . . . . . . .. 234
C. Regulation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235 1. Adjustmen t of the Equilibrium Between Division and
Elongation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 235 2. Regulation by IAA ................................ ,. 235
THE FORMATION OF SEEDS AND FRUITS 241
A. Complex Developmental Processes and Their Regulation 241
B. Formation of Seeds and Fruit. . . . . . . . . . . . . . . . . . . . . . . . . . .. 242 1. The Process of Formation. . . . . . . . . . . . . . . . . . . . . . . . . . .. 242 2. Regulation.......................................... 244
GERMINATION 250
A. Dormancy. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 250 I. Incomplete Embryos ............. , . . . . . . . . . . . . . . . . . .. 250 2. Maturation by Drying. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 250
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3. Impermeability to Water and/or Gases. . . . . . . . . . . . . . . .. 251 4. Inhibitors........................................... 251
B. Conditions for Germination. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 252 1. Water.............................................. 253 2. Oxygen............................................. 253 3. Temperature........................................ 253 4. Light............................................... 256
C. Mobilization of Reserve Materials. . . . . . . . . . . . . . . . . . . . . . .. 256
D. Assembly of the Photosynthetic Apparatus. . . . . . . . . . . . . . .. 257
E. Regulation of Germination by Photohormones .... . . . . . . .. 258
F. Regulation of Germination and Evolution. . . . . . . . . . . . . . . .. 259
THE VASCULAR SYSTEM 263
A. The Elements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 264
B. Differentiation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 264
C. Function. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 268 1. Transport in Both Directions. . . . . . . . . . . . . . . . . . . . . . . . .. 268 2. Transport of the Xylem . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 269 3. Transport in the Phloem. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 277
FLOWER FORMATION 286
A. Definitions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . .. 286
B. Temperature and Flower Induction: Vernalization. . . . . . . . .. 287 1. Petkus Rye. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 288 2. Henbane (Hyoscyamus Niger) . . . . . . . . . . . . . . . . . . . . . . . . .. 290 3. Streptocarpus Wendlandii. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 292 4. A Hypothesis Concerning Vernalization. . . . . . . . . . . . . . .. 296
C. Length of Day and Flower Induction: Photoperiodism...... 297 1. Long and Short Day Plants, Neutral Day Plants. . . . . . . .. 298 2. Analysis of Photoperiodism in Flower Induction. . . . . . .. 298 3. Photoperiodism in Flower Induction as a Sign of
Adaptation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 307 4. Light and Circadian Rhythms. . . . . . . . . . . . . . . . . . . . . . . .. 308
BIBLIOGRAPHY. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . .. 313 SOURCES OF ILLUSTRATION ......... ,.......................... 319 INDEX...................................................... 322
PLANT PHYSIOLOGY
Molecular, Biochemical, and Physiological Fundamentals of Metabolism and Development
