CHAPTER 1 Introduction ......................................... 1
Heredity and Environmental Regulation
of Growth ....................................................... 1
Physiological Regulation of Growth .............................. 2
Some Important Physiological Processes and Conditions ........ 3
Complexity of Physiological Processes ........................ 3
Problems of Foresters, Horticulturists, and Arborists ........... 3
Physiology in Relation to Present and Future Problems ........ 4
Summary ......................................................... 6
General References .............................................. 6
CHAPTER 2 The Woody Plant Body ................................. 9
Introduction .................................................... 9
Crown Form ..................................................... 10
Variations in Crown Form .................................... 10
Stem Form ...................................................... 11
Vegetative Organs and Tissues .................................. 12
Leaves ......................................................... 12
Angiosperms ................................................. 13
Variations in Size and Structure of Leaves .................. 15
Gymnosperms ................................................. 17
Stems .......................................................... 19
Sapwood and Heartwood ....................................... 19
Xylem Increments and Annual Rings ........................... 20
Earlywood and Latewood ...................................... 21
Phloem Increments ........................................... 22
Wood Structure of Gymnosperms .................................. 23
Axial Elements .............................................. 24
Horizontal Elements ......................................... 25
Wood Structure of Angiosperms .................................. 25
Axial Elements .............................................. 26
Horizontal Elements ......................................... 27
Bark ........................................................... 27
Roots .......................................................... 28
Adventitious Roots .......................................... 30
Root Tips ................................................... 30
Root Hairs .................................................. 31
Suberized and Unsuberized Roots ............................. 32
Mycorrhizas ................................................. 33
Reproductive Structures ........................................ 35
Angiosperms ................................................. 35
Gymnosperms ................................................. 36
Summary ........................................................ 37
General References ............................................. 38
CHAPTER 3 Vegetative Growth ................................... 39
Introduction ................................................... 39
Cell and Tissue Growth ......................................... 40
Dormancy .................................................... 42
Dormancy Concepts ........................................... 42
Hormonal Influences on Bud Dormancy ............................ 44
Shoot Growth ................................................ 45
Bud Characteristics ......................................... 46
Dormant and Adventitious Buds ............................... 46
Hormonal Influences on Shoot Growth ......................... 47
Leaf Growth ................................................. 48
Seasonal Leaf Growth Characteristics ........................ 49
Leaf Area Index ............................................. 50
Shoot Growth Types and Patterns ................................ 50
Determinate and Indeterminate Shoots ........................ 50
Epicormic Shoots ............................................ 50
Preformed and Neoformed Shoots .............................. 51
Recurrently Flushing Shoots ................................. 51
Abnormal Late-Season Shoots ................................. 52
Apical Dominance ............................................ 53
Maximum Height .............................................. 54
Shoot Growth in the Tropics .................................... 54
Cambial Growth ................................................. 55
Cell Division in the Cambium ................................ 55
Production of Xylem and Phloem .............................. 55
Time of Growth Initiation and Amounts of Xylem and Phloem
Produced ................................................. 56
Differentiation of Cambial Derivatives ...................... 56
Increase in Cell Size .................................... 58
Hormonal Influences on Cambial Growth ....................... 58
Cell Wall Thickening ........................................ 62
Loss of Protoplasts ......................................... 62
Formation and Development of Rays ........................... 63
Expansion of the Cambium .................................... 63
Variations in Growth Increments ............................. 63
Seasonal Duration of Cambial Growth ......................... 64
Anomalous Cambial Growth .................................... 64
Sapwood and Heartwood Formation ............................. 64
Wounding and Wound Healing .................................. 67
Root Growth .................................................... 68
Root Elongation ............................................. 69
Rate of Root growth ......................................... 70
Seasonal Variations ......................................... 70
Cambial Growth in Roots ..................................... 71
Shedding of Plant Parts ........................................ 72
Leaves ...................................................... 73
Branches .................................................... 77
Bark ........................................................ 78
Roots ....................................................... 79
Measurement and Analysis of Growth ............................. 80
Analysis of Growth .......................................... 80
Relative Growth Rates ....................................... 81
Allometric Formula and the Allometric Coefficient ........... 81
Net Assimilation Rate and Other Growth Parameters ........... 82
Limitations of Traditional Growth Analysis for Woody
Plants ................................................... 84
Summary ........................................................ 85
General References ............................................. 86
CHAPTER 4 Reproductive Growth ................................. 87
Introduction ................................................... 87
Reciprocal Relations between Vegetative and Reproductive
Growth ................................................... 88
Sexual Reproduction in Angiosperms ............................. 88
Flowering Periodicity ....................................... 88
Pollination ................................................. 90
Fruit Set ................................................... 90
Fertilization ............................................... 91
Postfertilization Development ............................... 91
Polyembryony ................................................ 92
Apomixis .................................................... 92
Parthenocarpy ............................................... 92
Growth of Fruits ............................................ 92
Fruit Ripening .............................................. 93
Sexual Reproduction in Gymnosperms ............................. 96
Cone Initiation and Development ............................. 96
Polyembryony ................................................ 99
Parthenocarpy ............................................... 99
Duration and Timing of Cone Development ..................... 99
Increase in Size and Dry Weight of Cones and Seeds ......... 100
Maturation of Seeds ........................................... 102
Abscission of Reproductive Structures ......................... 103
Abscission and Crop Yield .................................. 103
Summary ....................................................... 105
General References ............................................ 106
CHAPTER 5 Photosynthesis ..................................... 107
Introduction .................................................. 107
Chloroplast Development and Structure ......................... 108
Pigments ................................................... 109
Proteins ................................................... 110
Membrane Systems ........................................... 110
The Photosynthetic Mechanism .................................. 110
Light Reactions ............................................ 110
Photochemistry ............................................. 111
Electron Transport ......................................... 112
NADP+ Reduction ............................................ 112
Photophosphorylation ....................................... 112
Photoinhibition ............................................ 113
Dark Reactions ............................................. 116
Carbon Dioxide Uptake by Photosynthetic Tissues ............... 119
Carbon and Oxygen Isotope Discrimination During
Photosynthesis ............................................. 121
Variations in Rates of Photosynthesis ......................... 122
Species and Genetic Variations ............................. 123
Photosynthesis and Productivity ............................ 124
Diurnal Variations ......................................... 126
Seasonal Variations ........................................ 128
Environmental Factors ......................................... 132
Light Intensity ............................................ 132
Air Temperature ............................................ 140
Soil Temperature ........................................... 142
Carbon Dioxide ............................................. 144
Water Supply .................................................. 147
Soil Drying and Photosynthesis ............................. 147
Humidity ................................................... 151
Flooding ................................................... 152
Mineral Nutrition .......................................... 152
Salinity ................................................... 155
Pollution .................................................. 156
Applied Chemicals .......................................... 158
Plant Factors ................................................. 158
Stomatal Characteristics and Capacity of Photosynthetic
Partial Processes ....................................... 159
Source-Sink Relations ...................................... 160
Age of Leaves and of the Plant ............................. 162
Summary ....................................................... 164
General References ............................................ 166
CHAPTER 6 Enzymes, Energetics and Respiration ................ 169
Introduction .................................................. 169
Enzymes and Energetics ........................................ 169
Enzymes .................................................... 169
Energetics ................................................. 172
Respiration ................................................... 173
Biological Oxidations ...................................... 174
ATP ........................................................ 174
Other High-Energy Compounds ................................ 175
Glycolysis and the Krebs Cycle ............................. 175
Electron Transfer and Oxidative Phosphorylation ............ 176
Metabolic Decomposition of Respiration ..................... 176
Other Oxidases ............................................. 177
The Pentose Shunt .......................................... 178
Anaerobic Respiration ...................................... 178
Respiratory Quotient ....................................... 179
Photorespiration ........................................... 179
Respiration of Plants and Plant Parts ......................... 180
Amount of Food Used in Respiration ......................... 180
Respiration of Entire Trees ................................ 180
Respiration of Various Plant Parts ......................... 181
Seasonal Variations ........................................ 185
Scaling of Respiration to the Ecosystem Level Respiration
of Harvested Fruits ..................................... 187
Factors Affecting Respiration ................................. 188
Age and Physiological Condition of Tissues Available
Substrate ............................................... 188
Light ...................................................... 188
Hydration .................................................. 188
Temperature ................................................ 189
Composition of the Atmosphere .............................. 190
Soil Aeration .............................................. 191
Mechanical Stimuli and Injuries ............................ 191
Chemicals .................................................. 192
Air Pollutants ............................................. 193
Assimilation .................................................. 194
Summary ....................................................... 195
General References ............................................ 197
CHAPTER 7 Carbohydrates ...................................... 199
Introduction .................................................. 199
Kinds of Carbohydrates ........................................ 199
Monosaccharides ............................................ 199
Oligosaccharides ........................................... 200
Polysaccharides ............................................ 201
Carbohydrate Transformations .................................. 204
Phosphorylation ............................................ 204
Sucrose .................................................... 205
Starch ..................................................... 205
Uses of Carbohydrates ......................................... 205
Respiration ................................................ 206
Growth ..................................................... 206
Defense .................................................... 207
Leaching ................................................... 207
Exudation .................................................. 208
Accumulation of Carbohydrates ................................. 208
Carbohydrate Distribution .................................. 208
Storage Sites .............................................. 208
Autumn Coloration ............................................. 211
Summary ....................................................... 214
General References ............................................ 215
CHAPTER 8 Lipids, Terpenoids, and Related Substances ......... 217
Introduction .................................................. 217
Lipids ........................................................ 218
Simple Lipids .............................................. 218
Fatty Acids ................................................ 218
Lipid Distribution ......................................... 219
Waxes, Cutin, and Suberin ..................................... 220
Cuticle .................................................... 220
Waxes ...................................................... 220
Cutin and Suberin .......................................... 222
Internal Lipids ............................................... 223
Phospholipids .............................................. 223
Glycolipids ................................................ 224
Membrane Lipids ............................................ 224
Isoprenoids or Terpenoids ..................................... 224
Essential Oils ............................................. 224
Resins ..................................................... 225
Oleoresins ................................................. 226
Monoterpenes ............................................... 227
Carotenoids ................................................ 229
Rubber ..................................................... 229
Related Compounds .......................................... 231
Summary ....................................................... 231
General References ............................................ 232
CHAPTER 9 Nitrogen Metabolism ................................ 233
Introduction .................................................. 233
Distribution and Seasonal Fluctuations of Nitrogen ............ 234
Concentration in Various Tissues ........................... 234
Seasonal Changes in Nitrogen Concentration ................. 235
Changes in Distribution with Age ........................... 238
Important Nitrogen Compounds .................................. 240
Amino Acids ................................................ 240
Amino Acid Synthesis ....................................... 240
Nitrate Reduction .......................................... 240
Ammonia Assimilation ....................................... 242
Transamination ............................................. 242
Peptides ................................................... 242
Amides ..................................................... 242
Proteins ................................................... 243
Nucleic Acids and Related Compounds ........................ 244
Alkaloids .................................................. 244
Nitrogen Requirements ......................................... 244
Sources of Nitrogen ........................................... 246
Nitrogen Fixation .......................................... 247
Release from Litter ........................................ 249
The Nitrogen cycle ............................................ 250
Summary ....................................................... 253
General References ............................................ 253
CHAPTER 10 Mineral Nutrition .................................. 255
Introduction .................................................. 255
Functions of Mineral Nutrients and Effects of Deficiencies .... 256
Nitrogen ................................................... 256
Phosphorus ................................................. 256
Potassium .................................................. 256
Sulfur ..................................................... 257
Calcium .................................................... 257
Magnesium .................................................. 258
Iron ....................................................... 258
Manganese .................................................. 259
Zinc ....................................................... 259
Copper ..................................................... 259
Boron ...................................................... 259
Molybdenum ................................................. 260
Chlorine ................................................... 260
Nickel ..................................................... 260
Other Mineral Nutrients .................................... 260
Accumulation and Distribution of Mineral Nutrients ............ 261
Mineral Cycling ............................................... 262
The Soil Mineral pool ......................................... 262
Atmospheric Deposition ..................................... 262
Leaching from Plants ....................................... 264
Throughfall and Stemflow ................................... 266
Weathering of Rocks and Minerals ........................... 268
Decomposition of Organic Matter ............................ 268
Temperature ................................................ 269
Exudation from Roots ....................................... 270
Losses of Mineral Nutrients From Ecosystems ................... 270
Ecosystem Disturbance ...................................... 270
Temperate Forests .......................................... 270
Tropical Forests ........................................... 274
Leaching from Soil ......................................... 276
Absorption of Mineral Nutrients ............................... 277
Terminology ................................................ 277
Ion Movement in Soil ....................................... 279
The Absorbing Zone ......................................... 279
Factors Affecting Absorption ............................... 280
Absorption by Leaves and Twigs ............................. 283
Summary ....................................................... 284
General References ............................................ 285
CHAPTER 11 Absorption of Water and Ascent of Sap .............. 287
Introduction .................................................. 287
Importance of Water ........................................... 287
Cell Water Relations ....................................... 288
Cell Structure ............................................. 288
Water Status Quantification and Terminology ................ 288
Water Movement ............................................. 290
Measurement of Water Potential and Its Components .......... 291
The Soil-Plant-Atmosphere Continuum ........................ 292
Absorption of Water ........................................... 294
Soil Water ................................................. 294
Concentration and Composition of Soil Solution ............. 298
Soil Aeration .............................................. 298
Soil Temperature ........................................... 299
Absorption through Leaves and Stems ........................ 300
Absorption through Roots ................................... 301
Root Resistance ............................................ 304
Extent and Efficiency of Root Systems ...................... 305
Mycorrhizas and Water Relations ............................ 307
Water Absorption Processes .................................... 308
Osmotically Driven Absorption .............................. 308
Passive Absorption ......................................... 308
Root and Stem Pressures ....................................... 309
Root Pressure .............................................. 309
Guttation .................................................. 309
Maple Sap Flow ............................................. 309
Other Examples of Stem Pressure ............................ 312
Ascent of Sap ................................................. 312
The Water Conducting System ................................... 313
Efficiency of Water Conduction ............................. 315
Air Embolism and Xylem Blockage ............................ 317
Disease .................................................... 321
Summary ....................................................... 321
General References ............................................ 322
CHAPTER 12 Transpiration, Plant Water Balance and Adaptation
to Drought ......................................... 325
Introduction .................................................. 325
The Process of Transpiration 326
Transpiration as a Physical Process ........................... 326
Vapor Concentration Gradient from Leaf to Air .............. 327
Resistances in the Water Vapor Pathway ..................... 328
Factors Affecting Transpiration ............................... 330
Leaf Area .................................................. 330
Root-Shoot Ratio ........................................... 331
Leaf Size and Shape ........................................ 331
Leaf Orientation ........................................... 331
Leaf Surfaces .............................................. 332
Stomata .................................................... 333
Stomatal Control of Transpiration .......................... 338
Interaction of Factors Affecting Transpiration ................ 338
Transpiration Rates ........................................... 340
Water Loss From Plant Stands .................................. 342
Factors Controlling Evapotranspiration ..................... 342
Effects of Changes in Plant Cover .......................... 343
Thinning ................................................... 343
Relative Losses by Evaporation and Transpiration ........... 344
Changes in Species Composition ............................. 345
Methods for Reducing Transpiration ......................... 345
Transpiration Ratio and Water Use Efficiency ............... 346
The Water Balance ............................................. 349
The Dynamics of Plant Water Status ......................... 349
The Absorption Lag ......................................... 350
Internal Competition for Water ............................. 351
Long-Term Variations in Water Content ...................... 352
Seasonal Variations in Water Content ....................... 352
Effects of Water Stress ....................................... 354
Adaptation to Drought ......................................... 355
Drought Avoidance .......................................... 356
Drought Tolerance .......................................... 356
Drought Hardening .......................................... 364
Summary ....................................................... 365
General References ............................................ 366
CHAPTER 13 Plant Hormones and Other Endogenous Growth
Regulators ........................................ 367
Introduction .................................................. 367
Major Classes of Plant Hormones ............................... 367
Auxins ..................................................... 368
Gibberellins ............................................... 368
Cytokinins ................................................. 369
Abscisic Acid (ABA) ........................................ 371
Ethylene ................................................... 371
Other Regulatory Compounds .................................... 373
Brassinosteroids ........................................... 373
Jasmonates ................................................. 374
Salicylic Acid ............................................. 374
Phenolic Compounds ......................................... 374
Polyamines ................................................. 375
Other Compounds ............................................ 375
Mechanisms of Hormone Action .................................. 376
Summary ....................................................... 377
General Reference ............................................. 377
Bibliography .................................................. 379
Index ......................................................... 441
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