Ecophysiology of Photosynthesis

A: Molecular and Physiological Control and Limitations.- 1 Dynamics in Photosystem II Structure and Function.- 1.1 Introduction.- 1.2 Function of Photosystem II.- 1.3 Structure of Photosystem II.- 1.4 Dynamics in the D1 Protein in Rapid Turnover and Stress-Enhanced Photoinhibition.- 1.5 Photoinhibition and Environmental Stress.- 1.6 Regulation of Photosystem II by Phosphorylation.- 1.7 Conclusions.- References.- 2 Regulation of Photosynthetic Light Energy Capture, Conversion, and Dissipation in Leaves of Higher Plants.- 2.1 Introduction.- 2.2 The Concept of Excess Photon Flux Density.- 2.3 Regulation of Light Interception.- 2.3.1 Changes in Leaf Orientation.- 2.3.2 Changes in Leaf Reflectance.- 2.3.3 Chloroplast Movements.- 2.3.4 Changes in Chlorophyll Content and Photosynthetic Capacity.- 2.4 Regulation of Energy Dissipation.- 2.4.1 Dissipation in Metabolic Processes.- 2.4.2 Efficiency of Photochemical Energy Conversion and Extent of Nonradiative Energy Dissipation.- 2.4.3 Nonradiative Energy Dissipation and the Xanthophyll Cycle.- 2.4.4 Mechanism of Nonradiative Dissipation.- 2.5 Conclusions.- References.- 3 Chlorophyll Fluorescence as a Nonintrusive Indicator for Rapid Assessment of In Vivo Photosynthesis.- 3.1 Introduction.- 3.2 Indicator Function of Chlorophyll Fluorescence.- 3.3 Rapid Fluorescence Induction Kinetics.- 3.4 Slow Fluorescence Induction Kinetics and Fluorescence Quenching Under Steady-State Conditions.- 3.5 The Saturation Pulse Method.- 3.6 Quantum Yield and Rate Determination by Fluorescence Measurements.- 3.7 Fluorescence as an Indicator of Nonassimilatory Electron Flow.- 3.8 In Situ Measurements of ?F/Fm? and of Relative Electron Transport Rate.- 3.9 Yield Limitation and Excessive Photon Flux Density.- 3.10 Conclusions.- References.- 4 Higher Plant Respiration and Its Relationships to Photosynthesis.- 4.1 Introduction.- 4.2 Pathways and Controls of Respiration.- 4.2.1 Unique Properties of Plant Respiration and Mitochondrial Metabolism.- 4.2.2 Control of Respiration Rate.- 4.2.3 Energy Conservation During Plant Respiration.- 4.2.4 Respiration Rate and Carbohydrate Level.- 4.3 Respiration in Photosynthesizing Leaves.- 4.4 Photorespiration and Mitochondrial Metabolism.- 4.4.1 Oxidation of Photorespiratory NADH by the Respiratory Chain.- 4.4.2 Oxidation of Photorespiratory NADH via Substrate Shuttles.- 4.5 Daytime Photosynthesis and Nighttime Respiration.- 4.5.1 Light Level.- 4.5.2 CO2 Concentration.- 4.6 Photosynthesis and Root Respiration.- 4.7 Conclusions.- References.- 5 Apoplastic and Symplastic Proton Concentrations and Their Significance for Metabolism.- 5.1 Introduction.- 5.2 Definitions.- 5.2.1 The pH Concept.- 5.2.2 The Buffer Concept.- 5.2.3 Techniques to Determine Intra- and Intercellular pH.- 5.3 Cellular pH.- 5.3.1 The Apoplastic pH.- 5.3.2 The Symplastic pH.- 5.4 Conclusions.- References.- 6 The Significance of Assimilatory Starch for Growth in Arabidopsis thaliana Wild-Type and Starchless Mutants.- 6.1 Introduction.- 6.2 The Metabolic Pathway of Assimilatory Starch Formation and the Use of Mutants to Circumvent Chloroplast Starch Formation.- 6.3 The Diurnal Starch Turnover.- 6.4 Significance of Leaf Starch for Growth.- 6.4.1 Effects of Leaf Starch on Biomass Formation.- 6.4.2 Effects of Leaf Starch on Regulation of Shoot/Root Ratios.- 6.5 The Carbon Balance.- 6.6 Conclusions.- References.- 7 Photosynthesis, Storage, and Allocation.- 7.1 Introduction.- 7.2 The Impact of Photosynthesis on Growth, Storage, and Biomass Allocation in Transgenic Tobacco.- 7.2.1 Photosynthesis and Growth.- 7.2.2 Photosynthesis and Biomass Allocation.- 7.2.3 Carbon and Nitrogen Storage in Relation to Photosynthesis.- 7.2.4 The Tobacco System: Conclusions.- 7.3 Allocation in Relation to Shoot and Root Activity.- 7.3.1 Resource, Growth, and Allocation.- 7.3.2 Photosynthesis, Specific Absorption Rate, and Allocation.- 7.3.3 The Radish System: Conclusions.- 7.4 Storage as Related to Resource Availability.- 7.5 Conclusions.- References.- 8 Gas Exchange and Growth.- 8.1 Introduction.- 8.2 How Plants Grow.- 8.3 Photosynthesis and Growth Rates.- 8.4 The Importance of Allocation.- 8.5 Do Growth Rates Influence Carbon Assimilation?.- 8.6 Light Interception by Canopies and Plant Productivity.- 8.7 Phenology and Rates of Growth and Photosynthesis.- 8.8 Environmental Stresses Change the Relationship Between Photosynthesis and Growth.- 8.8.1 Water Deficits.- 8.8.2 Nitrogen Abundance.- 8.8.3 Temperature Effects.- 8.9 Conclusions.- Appendix: List of Symbols and Definitions.- References.- B: Responses of Photosynthesis to Environmental Factors.- 9 Internal Coordination of Plant Responses to Drought and Evaporational Demand.- 9.1 Introduction.- 9.2 Environmental and Plant-Internal Influences on Transpiration.- 9.3 Root-Leaf Signals Under Moisture Shortage Contribute to Drought Avoidance Responses of Leaves.- 9.4 Leaf Anatomy, Canopy Structure, and Stomatal Function.- 9.5 Xylem Conductivity and Leaf Conductance.- 9.6 Conclusions.- References.- 10 As to the Mode of Action of the Guard Cells in Dry Air.- 10.1 Introduction.- 10.2 Two Seminal Experiments.- 10.3 Some Relevant Observations.- 10.3.1 On Stomatal Mechanics.- 10.3.2 Signals and Responses.- 10.3.3 Hydrology of the Epidermis.- 10.4 Hypothesis.- 10.4.1 Feedback.- 10.4.2 Of Bubbles and Balloons.- 10.4.3 Piers and Vaults.- 10.5 Conclusions.- References.- 11 Direct Observations of Stomatal Movements.- 11.1 Introduction.- 11.2 The Methodical Approach.- 11.3 General Aspects.- 11.4 Stomatal Responses.- 11.4.1 Air-Humidity Response.- 11.4.2 Response to Changing CO2 Concentrations of the Air.- 11.4.3 Response to Heat.- 11.4.4 The Transient Phase and Other Pecularities of the Stomatal Response.- 11.5 Conclusions.- References.- 12 Carbon Gain in Relation to Water Use: Photosynthesis in Mangroves.- 12.1 Introduction.- 12.2 Water Relations: Why Be Conservative?.- 12.3 Implications of Conservative Water Use for Plant Function.- 12.4 Implications of Conservative Water Use for Display and Properties of Leaves.- 12.5 Coping with Excessive Light: Another By-Product of Conservative Water Use.- 12.6 Into the Future: Coping with Global Increase In Atmospheric CO2 Concentration.- References.- 13 Photosynthesis as a Tool for Indicating Temperature Stress Events.- 13.1 Introduction.- 13.2 Development of Temperature Stress and Characteristic Responses of Photosynthesis.- 13.3 Use of Photosynthetic Responses for Determining Heat Tolerance.- 13.4. Photosynthetic Function as a Criterion for Screening Chilling Susceptibility.- 13.5 Assay and Analysis of Freezing Events by Monitoring Photosynthesis.- 13.6 Conclusions.- References.- 14 Air Pollution, Photosynthesis and Forest Decline: Interactions and Consequences.- 14.1 Introduction.- 14.2 Sites of Interaction of Air Pollutants with Plants.- 14.3 The Magnitude of Fluxes into Leaves.- 14.4 Toxicity.- 14.5 Detoxification.- 14.5.1 The Path of Air Pollutants.- 14.5.2 The Fate of Nitrogen Oxides.- 14.5.3 The Fate of Ozone.- 14.5.4 The Fate of SO2.- 14.5.5 Acid-Dependent Cation Requirements.- 14.5.6 Interactions Between Different Air Pollutants.- 14.5.7 Interactions with Climatic Conditions.- 14.6 Tolerance Limits.- 14.7 Conclusions.- References.- C: Plant Performance in the Field.- 15 Photosynthesis in Aquatic Plants.- 15.1 Introduction.- 15.2 Definition of the Aquatic Habitat.- 15.3 The Diversity of Aquatic Plants.- 15.4 Contribution of Aquatic Plants to Global Net Primary Productivity.- 15.5 Photon Absorption and Use by Aquatic Plants.- 15.6 Inorganic Carbon Acquisition by Aquatic Plants: When Does It Limit Net Productivity?.- 15.7 Water Relations of Intertidal Aquatic Plants in Relation to Photosynthesis.- 15.8 Conclusions.- References.- 16 Photosynthesis in Poikilohydric Plants: A Comparison of Lichens and Bryophytes.- 16.1 Introduction.- 16.2 CO2 Exchange of Lichens and Bryophytes.- 16.2.1 Net Photosynthetic Rates.- 16.2.2 Compensation Points and Photorespiration.- 16.2.3 Dark Respiration Rates.- 16.2.4 Lichens and Bryophytes as Shade Plants.- 16.2.5 Thallus Water Content and Photosynthesis.- 16.2.6 Environmental CO2 Concentration.- 16.3 Plant Morphology and Photosynthesis.- 16.3.1 Bryophytes.- 16.3.2 Lichens.- 16.4 Water Location and Transport.- 16.4.1 Bryophytes.- 16.4.2 Lichens.- 16.5 An Upper Limit for Photosynthetic Rate?.- 16.6 Lichens and Bryophytes as Early Land Plants?.- 16.7 Conclusions.- References.- 17 The Consequences of Sunflecks for Photosynthesis and Growth of Forest Understory Plants.- 17.1 Introduction.- 17.2 Sunflecks in Forest Understories.- 17.3 Mechanisms Regulating the Utilization of Sunflecks.- 17.4 Photosynthesis in Natural Sunfleck Pegimes.- 17.5 The Significance of Sunflecks to Annual Carbon Gain.- 17.6 Consequences of Sunflecks for Growth and Reproduction.- 17.7 Conclusions.- References.- 18 Variation in Gas Exchange Characteristics Among Desert Plants.- 18.1 Introduction.- 18.2 Species Distribution Gradients in the Desert.- 18.3 Variation in Moisture and Temperature as Selective Forces for Photosynthetic Variation.- 18.3.1 Predictability of Precipitation.- 18.3.2 Drought Duration.- 18.3.3 Predictability of Temperature.- 18.4 Gas Exchange Patterns Among Life-Forms.- 18.4.1 Photosynthetic Pathway Distribution Among Life-Forms.- 18.4.2 Environment and Life-Form Distribution.- 18.5 Longevity and Gas Exchange.- 18.5.1 Water Use in Relation to Carbon Gain.- 18.5.2 Gas Exchange Flux Versus Set Point.- 18.5.3 Carbon Isotope Discrimination as a Measure of Intercellular Carbon Dioxide Concentration.- 18.5.4 Intercellular CO2 and Life History in C3 Plants.- 18.6 Integrating Gas Exchange Across Complex Environmental Gradients.- 18.6.1 Evaporative Gradients.- 18.6.2 Utilization of Summer Moisture Inputs.- 18.7 Conclusions.- References.- 19 Deuterium Content in Organic Material of Hosts and Their Parasites.- 19.1 Introduction.- 19.2 The Relative Deuterium Content in the Host and Parasitic Organic Material in Different Kinds of Parasite Performance.- 19.2.1 Isotope Contents of Galls.- 19.2.2 Isotope Contents of Holoparasites and Their Host Plants.- 19.2.3 Isotope Contents of Mistletoes (Hemiparasites) and Their Hosts.- 19.3 What Are the Reasons for the Isotope Discriminations?.- 19.3.1 ?13C.- 19.3.2 ?D.- 19.4 Conclusions.- References.- 20 Photosynthesis of Vascular Plants: Assessing Canopy Photosynthesis by Means of Simulation Models.- 20.1 Introduction.- 20.2 General Structure of Canopy Photosynthesis Models.- 20.3 The Simple Case: Single-Species Homogeneous Canopies.- 20.3.1 General Model Description.- 20.3.2 Model Validation.- 20.3.3 Case Study: How Do Different Parts of the Canopy Contribute to Total Canopy Photosynthesis?.- 20.4 Multispecies Homogeneous Canopies.- 20.4.1 Description of the Model Extensions.- 20.4.2 Case Study: Symmetric Competition.- 20.4.3 Case Study: Asymmetric Competition.- 20.5 Canopies with Nonhomogeneous Structure: Radiation Fluxes in Three Dimensions.- 20.5.1 Step 1: The Case of Single Plants.- 20.5.2 Step 2: Scaling Up from Single Plants to Plant Neighborhoods.- 20.6 Conclusions.- References.- 21 Effects of Phenology, Physiology, and Gradients in Community Composition, Structure, and Microclimate on Tundra Ecosystem CO2 Exchange.- 21.1 "Phenomenological" or "Aggregate" Models of Ecosystem CO2 Flux.- 21.2 Concept and General Structure of the Stand Model GAS-FLUX.- 21.3 Structural Inputs to GAS-FLUX Along Water Gradients in Tundra.- 21.4 Ecophysiological Inputs to GAS-FLUX Along Water Gradients in Tundra.- 21.4.1 CO2 Exchange of Vascular Plant Species of Differing Growth Forms.- 21.4.2 CO2 Exchange of Poikilohydric Plants.- 21.4.3 CO2 Exchange of the Soil.- 21.5 Simulations of Ecosystem CO2 Exchange.- 21.5.1 Diurnal Course of Gas Exchange of Major Tundra Structural Components.- 21.5.2 Environmental Effects on Diurnal CO2 Exchange and Aggregate Formulations.- 21.6 Conclusions: Future Directions of GAS-FLUX Development.- References.- D: Global Aspects of Photosynthesis.- 22 Leaf Diffusive Conductances in the Major Vegetation Types of the Globe.- 22.1 The Significance of Leaf Conductances in Vegetation Modeling.- 22.2 Constraints of Utilizing Leaf Conductances in Vegetation Modeling.- 22.3 How Was the Data Set Compiled?.- 22.3.1 Definition of Maximum Leaf Conductance.- 22.3.2 Definition of Minimum Leaf Conductance for Water Vapor.- 22.3.3 Definition of Stomatal Response Functions.- 22.4 Selection of Vegetation Types.- 22.5 Maximum Leaf Diffusive Conductances in Important Vegetation Types.- 22.6 Maximum Leaf Diffusive Conductances and Maximum Rate of Leaf Photosynthesis.- 22.7 Minimum Leaf Diffusive Conductances.- 22.8 Stomatal Responses in the Field.- 22.8.1 Long-Term Trends and Seasonal Changes.- 22.8.2 Short-Term and Diurnal Changes.- 22.9 Conclusions and Recommendations for Further Research.- References.- 23 Predictions and Measurements of the Maximum Photosynthetic Rate, Amax, at the Global Scale.- 23.1 Introduction.- 23.2 Philosophy.- 23.3. Experimental Evidence for the Soil N Supply Constraint on Amax.- 23.3.1 Introduction.- 23.3.2 Experimental Detail.- 23.3.3 Results.- 23.3.4 Discussion.- 23.4 Modeling Amax at the Global Scale.- 23.4.1 Introduction.- 23.4.2 Method of Predicting Amax from Soil C.- 23.4.3 Validating Amax Predictions.- 23.4.4 Predicting Amax from Soil C and Soil N.- 23.5 Global Predictions and Tests of Soil-Based Amax.- 23.6 Conclusions: Gobai Scale Maps of Observed and Predicted Amax.- Appendix: References for Global Measurements of Amax.- References.- 24 Remote Sensing of Terrestrial Photosynthesis.- 24.1 Remote Sensing, from the Leaf of the Globe.- 24.1.1 A Range of Approaches.- 24.2 Models: from Radiance to CO2 Exchange.- 24.3 Remote Sensing of Photosynthetic Capacity.- 24.3.1 Absorbed Radiation.- 24.3.2 Photosynthetic Pigments.- 24.3.3 Other Compounds.- 24.4 Remote Sensing of Physiological Status.- 24.4.1 Fluorescence.- 24.4.2 Xanthophyll Pigments.- 24.4.3 Canopy Temperature.- 24.5 Remote Sensing of Environmental Factors.- 24.6 Conclusions.- References.- 25 Are C4 Pathway Plants Threatened by Global Climatic Change?.- 25.1 Introduction.- 25.2 Low Atmospheric CO2 Concentrations and Evolution of C4 Pathway Photosynthesis.- 25.3 Physiological Flexibility in C4 Plants Under High CO2 Concentrations.- 25.3.1 Coordination of Metabolism.- 25.3.2 Leakage of CO2 from the Bundle Sheath.- 25.3.3 Translocation of Carbohydrate.- 25.3.4 Water Use Efficiency.- 25.3.5 Nitrogen Use Efficiency.- 25.4 Growth and Competition Between C4 and C3 Plants Under Elevated CO2.- 25.5 Present Distributions and Diversity of C4 Plants.- 25.6 Future Distributions of C4 Plants.- 25.7 Conclusion.- References.- E: Perspectives in Ecophysiological Research of Photosynthesis.- 26 Overview: Perspectives in Ecophysiological Research of Photosynthesis.- 26.1 Introduction: A Historic Perspective.- 26.2 Methodology.- 26.3 The Molecular and Biochemical Venue of Photosynthetic Ecophysiology.- 26.4 Balancing Photosynthesis and Transpiration.- 26.5 Photosynthetic Performance of Different Plant Groups.- 26.6 Photosynthesis and Global Climate Change: Making Global Predictions.- 26.7 Where Will Ecophysiology of Photosynthesis Venture in the Coming Decade? We Offer Some Thoughts.- References.- Species Index.