Plant Physiological Ecology

Plant Physiological Ecology......Page 2
Foreword to Second Edition......Page 5
About the Authors......Page 7
Foreword to First Edition......Page 9
Acknowledgments......Page 11
Abbreviations......Page 12
Contents......Page 16
 1.2 The Roots of Ecophysiology......Page 29
 1.3 Physiological Ecology and the Distribution of Organisms......Page 30
 1.4 Time Scale of Plant Response to Environment......Page 32
 1.5 Conceptual and Experimental Approaches......Page 34
 1.7 The Structure of the Book......Page 35
  2.2.1 The ‘‘Light’’ and ‘‘Dark’’ Reactions of Photosynthesis......Page 38
   2.2.1.1 Absorption of Photons......Page 39
   2.2.1.2 Fate of the Excited Chlorophyll......Page 40
   2.2.1.4 Photosynthetic Carbon Reduction......Page 41
   2.2.1.5 Oxygenation and Photorespiration......Page 42
   2.2.2.1 Demand for CO2-the CO2-Response Curve......Page 43
  Modeling C3 Photosynthesisis......Page 46
  2.2.2.2 Supply of CO2-Stomatal and Boundary Layer Conductances......Page 48
  Fractionation of Carbon Isotopes in Plants......Page 49
  2.3.1 The Light Climate Under a Leaf Canopy......Page 53
   2.3.2.1 The Light-Response Curve of Sun and Shade Leaves......Page 54
   2.3.2.2 Anatomy and Ultrastructure of Sun and Shade Leaves......Page 56
  Carbon--Fixation and Light--Absorption Profiles Inside Leaves......Page 58
  2.3.2.3 Biochemical Differences Between Shade and Sun Leaves......Page 59
  2.3.2.4 The Light-Response Curve of Sun and Shade Leaves Revisited......Page 60
  2.3.2.5 The Regulation of Acclimation......Page 62
   2.3.3.1 Photoinhibition-Protection by Carotenoids of the Xanthophyll Cycle......Page 63
  Chlorophyll Fluorescence......Page 64
  2.3.3.2 Chloroplast Movement in Response to Changes in Irradiance......Page 68
  2.3.4 Responses to Variable Irradiance......Page 69
   2.3.4.2 Light Activation of Rubisco......Page 70
   2.3.4.4 Metabolite Pools in Sun and Shade Leaves......Page 72
  2.4.1 Partitioning Within the Cell......Page 74
  2.4.2 Short-Term Regulation of Photosynthetic Rate by Feedback......Page 75
 2.5 Responses to Availability of Water......Page 78
  The Measurement of Gas Exchange......Page 79
  2.5.1 Regulation of Stomatal Opening......Page 80
  2.5.2 The A-Cc Curve as Affected by Water Stress......Page 81
  2.5.3 Carbon-Isotope Fractionation in Relation to Water-Use Efficiency......Page 83
  2.5.4 Other Sources of Variation in Carbon-Isotope Ratios in C3 Plants......Page 84
  2.6.1 The Photosynthesis-Nitrogen Relationship......Page 85
  2.6.3 Photosynthesis, Nitrogen, and Leaf Life Span......Page 86
  2.7.1 Effects of High Temperatures on Photosynthesis......Page 87
  2.7.2 Effects of Low Temperatures on Photosynthesis......Page 88
 2.8 Effects of Air Pollutants on Photosynthesis......Page 90
  2.9.2 Biochemical and Anatomical Aspects......Page 91
  2.9.3 Intercellular and Intracellular Transport of Metabolites of the C4 Pathway......Page 94
  2.9.4 Photosynthetic Efficiency and Performance at High and Low Temperatures......Page 95
  2.9.5 C3-C4 Intermediates......Page 98
  2.9.6 Evolution and Distribution of C4 Species......Page 100
  2.10.1 Introduction......Page 102
  2.10.2 Physiological, Biochemical, and Anatomical Aspects......Page 103
  2.10.4 Incomplete and Facultative CAM Plants......Page 106
  2.10.5 Distribution and Habitat of CAM Species......Page 107
  2.10.6 Carbon-Isotope Composition of CAM Species......Page 108
  2.11.2 The CO2 Supply in Water......Page 109
  2.11.3 The Use of Bicarbonate by Aquatic Macrophytes......Page 110
  2.11.4 The Use of CO2 from the Sediment......Page 111
  2.11.7 The Role of Aquatic Macrophytes in Carbonate Sedimentation......Page 112
 2.12 Effects of the Rising CO2 Concentration in the Atmosphere......Page 114
  2.12.1 Acclimation of Photosynthesis to Elevated CO2 Concentrations......Page 116
 2.13 Summary: What Can We Gain from Basic Principles and Rates of Single-Leaf Photosynthesis?......Page 117
  2.2.1 The Respiratory Quotient......Page 127
  2.2.3 Mitochondrial Metabolism......Page 129
   2.2.3.1 The Complexes of the Electron-Transport Chain......Page 130
   2.2.3.3 Substrates, Inhibitors, and Uncouplers......Page 131
   2.2.3.4 Respiratory Control......Page 132
   2.2.5.2 ATP Production In Vivo......Page 133
   2.2.6.1 Competition or Overflow?......Page 135
   2.2.6.2 The Intricate Regulation of the Alternative Oxidase......Page 136
  2.3.1 Heat Production......Page 138
  2.3.2 Can We Really Measure the Activity of the Alternative Path?......Page 139
  2.3.3 The Alternative Path as an Energy Overflow......Page 140
  Measuring Oxygen-Isotope Fractionation in Respiration......Page 141
  2.3.5 NADH Oxidation to Oxidize Excess Redox Equivalents from the Chloroplast......Page 143
  2.3.7 A Summary of the Various Ecophysiological Roles of the Alternative Oxidase......Page 144
   2.4.1.2 Fermentation......Page 145
   2.4.1.3 Cytosolic Acidosis......Page 146
   2.4.1.4 Avoiding Hypoxia: Aerenchyma Formation......Page 147
  2.4.2 Salinity and Water Stress......Page 148
  2.4.4 Irradiance......Page 149
  2.4.5 Temperature......Page 153
  2.4.6 Low pH and High Aluminum Concentrations......Page 155
  2.4.7 Partial Pressures of CO2......Page 156
  2.4.8 Effects of Plant Pathogens......Page 157
   2.5.1.1 Root Respiration......Page 158
   2.5.1.2 Respiration of Other Plant Parts......Page 159
   2.5.2.1 Maintenance Respiration......Page 160
   2.5.2.2 Growth Respiration......Page 162
   2.5.2.4 Experimental Evidence......Page 166
 2.6 Plant Respiration: Why Should It Concern Us from an Ecological Point of View?......Page 169
 2.2 Major Transport Compounds in the Phloem: Why Not Glucose?......Page 177
 2.3 Phloem Structure and Function......Page 179
  2.3.2 Minor Vein Anatomy......Page 180
  2.3.3 Sugar Transport against a Concentration Gradient......Page 181
 2.5 Phloem Unloading......Page 183
 2.6 The Transport Problems of Climbing Plants......Page 186
 2.7 Phloem Transport: Where to Move from Here?......Page 187
  3.1.1 The Role of Water in Plant Functioning......Page 189
  3.1.2 Transpiration as an Inevitable Consequence of Photosynthesis......Page 190
 3.3 Water Availability in Soil......Page 191
  The Water Potential of Osmotic Solutes and the Air......Page 192
  Positive and Negative Hydrostatic Pressures......Page 194
  3.3.1 The Field Capacity of Different Soils......Page 195
  3.3.2 Water Movement Toward the Roots......Page 196
  3.3.3 Rooting Profiles as Dependent on Soil Moisture Content......Page 197
  Oxygen and Hydrogen Stable Isotopes......Page 198
  3.3.4 Roots Sense Moisture Gradients and Grow Toward Moist Patches......Page 199
 3.4 Water Relations of Cells......Page 200
  3.4.2 Cell-Wall Elasticity......Page 201
  3.4.3 Osmotic and Elastic Adjustment as Alternative Strategies......Page 203
  3.5.1 The Soil-Plant-Air Continuum......Page 204
  3.5.2 Water in Roots......Page 205
  3.5.3 Water in Stems......Page 209
  Methods to Measure Sap Flow in Intact Plants......Page 210
  3.5.3.1 Can We Measure Negative Xylem Pressures?......Page 211
  3.5.3.2 The Flow of Water in the Xylem......Page 212
  3.5.3.3 Cavitation or Embolism: The Breakage of the Xylem Water Column......Page 214
  3.5.3.4 Can Embolized Conduits Resume Their Function?......Page 217
  3.5.3.5 Trade-off Between Conductance and Safety......Page 218
  3.5.3.6 Transport Capacity of the Xylem and Leaf Area......Page 220
  3.5.3.7 Storage of Water in Stems......Page 221
   3.5.4.1 Effects of Soil Drying on Leaf Conductance......Page 222
   3.5.4.2 The Control of Stomatal Movements and Stomatal Conductance......Page 225
   3.5.4.3 Effects of Vapor Pressure Difference or Transpiration Rate on Stomatal Conductance......Page 227
   3.5.4.5 The Cuticular Conductance and the Boundary Layer Conductance......Page 229
   3.5.4.6 Stomatal Control: A Compromise Between Carbon Gain and Water Loss......Page 230
  3.6.1 Water-Use Efficiency and Carbon-Isotope Discrimination......Page 232
  3.6.2 Leaf Traits That Affect Leaf Temperature and Leaf Water Loss......Page 233
  3.6.3 Water Storage in Leaves......Page 235
 3.7 Water Availability and Growth......Page 236
  3.8.1 Desiccation Avoidance: Annuals and Drought-Deciduous Species......Page 237
  3.8.3 Resurrection Plants......Page 238
 3.9 Winter Water Relations and Freezing Tolerance......Page 240
 3.11 Final Remarks: The Message That Transpires......Page 242
  4.2.1 Short Overview of a Leaf’s Energy Balance......Page 250
  4.2.2 Short-Wave Solar Radiation......Page 251
  4.2.3 Long-Wave Terrestrial Radiation......Page 254
  4.2.4 Convective Heat Transfer......Page 255
  4.2.5 Evaporative Energy Exchange......Page 257
 4.3 Modeling the Effect of Components of the Energy Balance on Leaf Temperature......Page 259
 4.4 A Summary of Hot and Cool Topics......Page 260
  4.2.2 Effects of Ultraviolet Radiation......Page 262
   4.2.2.2 Protection Against UV: Repair or Prevention......Page 263
  4.3.1 How Do Plants Avoid Damage by Free Radicals at Low Temperature?......Page 264
  4.3.3 Are Isoprene and Monoterpene Emissions an Adaptation to High Temperatures?......Page 266
  4.3.4 Chilling Injury and Chilling Tolerance......Page 267
  4.3.5 Carbohydrates and Proteins Conferring Frost Tolerance......Page 268
 4.4 Global Change and Future Crops......Page 269
 5.2 Canopy Water Use......Page 272
  Optimization of Nitrogen Allocation to Leaves in Plants Growing in Dense Canopies......Page 273
 5.3 Canopy CO2 Fluxes......Page 276
 5.4 Canopy Water-Use Efficiency......Page 277
 5.6 Aiming for a Higher Level......Page 278
   6.2.1.1 Nutrient Availability as Dependent on Soil Age......Page 280
   6.2.1.2 Nutrient Supply Rate......Page 282
   6.2.1.3 Nutrient Movement to the Root Surface......Page 284
   6.2.2.1 Increasing the Roots’ Absorptive Surface......Page 287
   6.2.2.2 Transport Proteins: Ion Channels and Carriers......Page 288
    6.2.2.3.1 Response to Nutrient Supply......Page 290
    6.2.2.3.2 Response to Nutrient Demand......Page 292
    6.2.2.3.3 Response to Other Environmental and Biotic Factors......Page 293
   6.2.2.4 Acquisition of Nitrogen......Page 294
    6.2.2.5.1 Plants Can Also Use Some Organic Phosphate Compounds......Page 295
    6.2.2.5.2 Excretion of Phosphate-Solubilizing Compounds......Page 296
    6.2.2.6.1 Changing the Rhizosphere pH......Page 300
    6.2.2.6.2 Excretion of Organic Chelates......Page 301
   6.2.2.7 Rhizosphere Mineralization......Page 304
  Molecular Control of Local Root Proliferation......Page 305
  6.2.3 Sensitivity Analysis of Parameters Involved in Phosphate Acquisition......Page 307
   6.3.1.1 Aluminum Toxicity......Page 309
   6.3.1.3 Aluminum Resistance......Page 312
  6.3.2 Calcareous Soils......Page 313
   6.3.3.1 Why Are the Concentrations of Heavy Metals in Soil High?......Page 314
   6.3.3.2 Using Plants to Clean or Extract Polluted Water and Soil: Phytoremediation and Phytomining......Page 315
   6.3.3.4 Heavy-Metal-Resistant Plants......Page 316
  6.3.4 Saline Soils: An Ever-Increasing Problem in Agriculture......Page 321
   6.3.4.2 Energy-Dependent Salt Exclusion from Roots......Page 322
   6.3.4.4 Transport of Na+ from the Leaves to the Roots and Excretion via Salt Glands......Page 323
  6.3.5 Flooded Soils......Page 326
   6.4.1.1 Tissue Nutrient Concentration......Page 327
   6.4.1.2 Tissue Nutrient Requirement......Page 328
   6.4.2.2 The Mean Residence Time of Nutrients in the Plant......Page 329
   6.4.3.1 Leaching Loss......Page 331
   6.4.3.2 Nutrient Loss by Senescence......Page 332
  6.4.4 Ecosystem Nutrient-Use Efficiency......Page 333
 6.5 Mineral Nutrition: A Vast Array of Adaptations and Acclimations......Page 335
 7.2 Growth of Whole Plants and Individual Organs......Page 346
   7.2.1.2 Plants with High Nutrient Concentrations Can Grow Faster......Page 347
   7.2.2.1 Cell Division and Cell Expansion: The Lockhart Equation......Page 348
   7.2.2.2 Cell-Wall Acidification and Removal of Calcium Reduce Cell-Wall Rigidity......Page 349
  Phytohormones......Page 351
  7.2.2.3 Cell Expansion in Meristems Is Controlled by Cell-Wall Extensibility and Not by Turgor......Page 352
 7.3 The Physiological Basis of Variation in RGR-Plants Grown with Free Access to Nutrients......Page 353
  Phytochrome......Page 354
  7.3.1 SLA Is a Major Factor Associated with Variation in RGR......Page 355
  7.3.3 Anatomical and Chemical Differences Associated with Leaf Mass Density......Page 357
  7.3.5 RGR and the Rate of Leaf Elongation and Leaf Appearance......Page 358
  7.3.7 RGR and Suites of Plant Traits......Page 359
 7.4 Allocation to Storage......Page 360
  7.4.1 The Concept of Storage......Page 361
  7.4.3 Storage and Remobilization in Annuals......Page 362
  7.4.5 Storage in Perennials......Page 363
 7.5 Environmental Influences......Page 365
    7.5.1.1.1 Effects on Growth Rate, Net Assimilation Rate, and Specific Leaf Area......Page 366
    7.5.1.1.2 Adaptations to Shade......Page 367
    7.5.1.1.4 The Role of Phytochrome......Page 368
    7.5.1.1.6 Effects of Total Level of Irradiance......Page 369
   7.5.1.2 Effects of the Photoperiod......Page 370
   7.5.2.2 Changes in the Allocation Pattern......Page 371
  7.5.3 Growth as Affected by Soil Water Potential and Salinity......Page 372
   7.5.3.3 Effects on Root Elongation......Page 373
   7.5.4.1 Cycling of Nitrogen Between Roots and Leaves......Page 374
   7.5.4.2 Hormonal Signals That Travel via the Xylem to the Leaves......Page 375
   7.5.4.4 Integrating Signals from the Leaves and the Roots......Page 376
   7.5.4.6 Nitrogen Allocation to Different Leaves, as Dependent on Incident Irradiance......Page 377
   7.5.5.2 Changes in Root Length and Diameter: A Modification of the Lockhart Equation......Page 379
  7.5.6 Growth as Affected by Soil Flooding......Page 380
   7.5.6.1 The Pivotal Role of Ethylene......Page 381
   7.5.6.2 Effects on Water Uptake and Leaf Growth......Page 382
  7.5.7 Growth as Affected by Submergence......Page 383
   7.5.7.2 Perception of Submergence and Regulation of Shoot Elongation......Page 384
  7.5.8 Growth as Affected by Touch and Wind......Page 385
  7.5.9 Growth as Affected by Elevated Concentrations of CO2 in the Atmosphere......Page 386
  7.6.1 Fast- and Slow-Growing Species......Page 387
  7.6.2 Growth of Inherently Fast- and Slow-Growing Species Under Resource-Limited Conditions......Page 388
   7.6.3.1 Various Hypotheses......Page 389
   7.6.3.2 Selection on RGRmax Itself, or on Traits That Are Associated with RGRmax?......Page 390
   7.6.3.3 An Appraisal of Plant Distribution Requires Information on Ecophysiology......Page 391
 7.7 Growth and Allocation: The Messages About Plant Messages......Page 392
 8.2 Seed Dormancy and Germination......Page 400
  8.2.1 Hard Seed Coats......Page 401
  8.2.2 Germination Inhibitors in the Seed......Page 402
  8.2.4 Other External Chemical Signals......Page 403
  8.2.5 Effects of Light......Page 405
  8.2.6 Effects of Temperature......Page 407
  8.2.7 Physiological Aspects of Dormancy......Page 409
  8.3.1 Seedling Phase......Page 410
  8.3.2 Juvenile Phase......Page 411
   8.3.2.1 Delayed Flowering in Biennials......Page 412
   8.3.2.3 Vegetative Reproduction......Page 413
   8.3.2.4 Delayed Greening During Leaf Development in Tropical Trees......Page 415
   8.3.3.1 Timing by Sensing Daylength: Long-Day and Short-Day Plants......Page 416
   8.3.3.3 Timing by Sensing Temperature: Vernalization......Page 418
   8.3.3.5 Attracting Pollinators......Page 419
   8.3.3.6 The Cost of Flowering......Page 420
  8.3.4 Fruiting......Page 421
  8.4.1 Dispersal Mechanisms......Page 422
 8.5 The Message to Disperse: Perception, Transduction, and Response......Page 423
 9.2 Mycorrhizas......Page 428
  9.2.1 Mycorrhizal Structures: Are They Beneficial for Plant Growth?......Page 429
   9.2.1.1 The Infection Process......Page 433
   9.2.1.2 Mycorrhizal Responsiveness......Page 435
  9.2.2 Nonmycorrhizal Species and Their Interactions with Mycorrhizal Species......Page 437
   9.2.3.1 Mechanisms That Account for Enhanced Phosphate Absorption by Mycorrhizal Plants......Page 438
   9.2.3.2 Suppression of Colonization at High Phosphate Availability......Page 440
  9.2.4 Effects on Nitrogen Nutrition......Page 441
  9.2.5 Effects on the Acquisition of Water......Page 442
  9.2.6 Carbon Costs of the Mycorrhizal Symbiosis......Page 443
  9.2.7 Agricultural and Ecological Perspectives......Page 444
 9.3 Associations with Nitrogen-Fixing Organisms......Page 446
  9.3.1 Symbiotic N2Fixation Is Restricted to a Fairly Limited Number of Plant Species......Page 447
  9.3.3 The Infection Process in the Legume--Rhizobium Association......Page 449
   9.3.3.2 Rhizobial nod Genes......Page 450
   9.3.3.3 Entry of the Bacteria......Page 452
   9.3.3.4 Final Stages of the Establishment of the Symbiosis......Page 453
  9.3.4 Nitrogenase Activity and Synthesis of Organic Nitrogen......Page 454
  9.3.5 Carbon and Energy Metabolism of the Nodules......Page 456
  9.3.6 Quantification of N2 Fixation In Situ......Page 457
  9.3.7 Ecological Aspects of the Nonsymbiotic Association with N2-Fixing Microorganisms......Page 458
  9.3.8 Carbon Costs of the Legume-Rhizobium Symbiosis......Page 459
  9.3.9 Suppression of the Legume-Rhizobium Symbiosis at Low pH and in the Presence of a Large Supply of Combined Nitrogen......Page 460
 9.4 Endosymbionts......Page 461
 9.5 Plant Life Among Microsymbionts......Page 462
 9.2 Allelopathy (Interference Competition)......Page 469
  9.3.1 Defense Against Herbivores......Page 472
  9.3.3 The Arms Race of Plants and Herbivores......Page 475
  9.3.4 How Do Plants Avoid Being Killed by Their Own Poisons?......Page 479
  9.3.5 Secondary Metabolites for Medicines and Crop Protection......Page 481
  9.4.1 Abiotic Factors......Page 484
  9.4.2 Induced Defense and Communication Between Neighboring Plants......Page 486
  9.4.3 Communication Between Plants and Their Bodyguards......Page 488
  9.5.1 Diversion of Resources from Primary Growth......Page 490
  9.5.2 Strategies of Predators......Page 492
 9.6 Detoxification of Xenobiotics by Plants: Phytoremediation......Page 493
 9.7 Secondary Chemicals and Messages That Emerge from This Chapter......Page 496
 9.2 Constitutive Antimicrobial Defense Compounds......Page 502
 9.3 The Plant’s Response to Attack by Microorganisms......Page 504
 9.4 Cross-Talk Between Induced Systemic Resistance and Defense Against Herbivores......Page 508
 9.5 Messages from One Organism to Another......Page 511
 9.1 Introduction......Page 513
  9.2.1 Seed Germination......Page 514
  9.2.2 Haustoria Formation......Page 515
  9.2.3 Effects of the Parasite on Host Development......Page 518
 9.3 Water Relations and Mineral Nutrition......Page 520
 9.4 Carbon Relations......Page 522
 9.5 What Can We Extract from This Chapter?......Page 523
 9.1 Introduction......Page 526
  Plant Ecological Strategies......Page 527
   Leaf Economics Spectrum......Page 528
   The Future......Page 529
 9.3 How Do Plants Perceive the Presence of Neighbors?......Page 530
  9.4.1 Growth Rate and Tissue Turnover......Page 533
  9.4.2 Allocation Pattern, Growth Form, and Tissue Mass Density......Page 534
  9.4.3 Plasticity......Page 535
  9.5.1 Nutrients......Page 537
  9.5.2 Water......Page 538
  9.5.4 Carbon Dioxide......Page 539
  9.6.3 Allelochemical Benefits......Page 542
 9.7 Plant-Microbial Symbiosis......Page 543
 9.8 Succession......Page 545
 9.9 What Do We Gain from This Chapter?......Page 547
 9.2 Structures Associated with the Catching of the Prey and Subsequent Withdrawal of Nutrients from the Prey......Page 553
 9.3 Some Case Studies......Page 556
  9.3.1 Dionaea Muscipula......Page 557
  9.3.2 The Suction Traps of Utricularia......Page 559
  9.3.3 The Tentacles of Drosera......Page 561
  9.3.5 Passive Traps of Genlisea......Page 562
 9.4 The Message to Catch......Page 563
 10.1 Introduction......Page 565
  10.2.1 Species Effects on Litter Quality: Links with Ecological Strategy......Page 566
  10.2.2 Environmental Effects on Decomposition......Page 567
  10.3.1 The Process of Nutrient Release......Page 568
  10.3.2 Effects of Litter Quality on Mineralization......Page 569
  10.3.3 Root Exudation and Rhizosphere Effects......Page 570
 10.4 The End Product of Decomposition......Page 572
  2.1 Scaling from Plants to Ecosystems......Page 575
  2.2 Physiological Basis of Productivity......Page 576
  2.3 Disturbance and Succession......Page 578
  2.4 Photosynthesis and Absorbed Radiation......Page 579
  2.6 The Global Carbon Cycle......Page 581
  3.1 Vegetation Controls over Nutrient Uptake and Loss......Page 583
   4.1.1 Albedo......Page 585
   4.1.2 Surface Roughness and Energy Partitioning......Page 586
   4.2.1 Evapotranspiration and Runoff......Page 587
 5 Moving to a Higher Level: Scaling from Physiology to the Globe......Page 588
Glossary......Page 592
Index......Page 610