Ecophysiology of Photosynthesis

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In a world of increasing atmospheric CO2, there is intensified interest in the ecophysiology of photosynthesis and increasing attention is being given to carbon exchange and storage in natural ecosystems. We need to know how much photosynthesis of terrestrial and aquatic vegetation will change as global CO2 increases. Are there major ecosystems, such as the boreal forests, which may become important sinks of CO2 and slow down the effects of anthropogenic CO2 emissions on climate? Will the composition of the vegetation change as a result of CO2 increase?
This volume reviews the progress which has been made in understanding photosynthesis in the past few decades at several levels of integration from the molecular level to canopy, ecosystem and global scales.

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.

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Ecophysiology of Photosynthesis
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