Photosynthesis II: Photosynthetic Carbon Metabolism and Related Processes

ISBN-13: 9783642672446 / Angielski / Miękka / 2011 / 578 str.

M. Gibbs; E. Latzko

Opis

M. GIBBS and E. LATZKO In the preface to his Experiments upon Vegetables, INGEN-Housz wrote in 1779: "The discovery of Dr. PRIESTLEY that plants have a power of correcting bad air . . . shows . . . that the air, spoiled and rendered noxious to animals by their breath ing in it, serves to plants as a kind of nourishment. " INGEN-Housz then described his own experiments in which he established that plants absorb this "nourishment" more actively in brighter sunlight. By the turn of the eighteenth century, the "nourishment" was recognized to be CO . Photosynthetic CO2 assimilation, the 2 major subject of this encyclopedia volume, had been discovered. How plants assimilate the CO was a question several successive generations 2 of investigators were unable to answer; scientific endeavor is not a discipline in which it is easy to "put the cart before the horse." The horse, in this case, was the acquisition of radioactive isotopes of carbon, especially 14c. The cart which followed contained the Calvin cycle, formulated by CALVIN, BENSON and BASSHAM in the early 1950's after (a) their detection of glycerate-3-P as the first stable product of CO fixation, (b) their discovery, and that by HORECKER 2 and RACKER, of the COz-fixing enzyme RuBP carboxylase, and (c) the reports by GIBBS and by ARNON of an enzyme (NADP-linked GAP dehydrogenase) capable of using the reducing power made available from sunlight (via photo synthetic electron transport) to reduce the glycerate-3-P to the level of sugars."

Spis treści

I. Introduction.- II. CO2 Assimilation.- II A. The Reductive Pentose Phosphate Cycle.- 1. The Reductive Pentose Phosphate Cycle and Its Regulation.- A. Introduction.- B. The Reductive Pentose Phosphate Cycle.- I. The Cyclic Path of Carbon Dioxide Fixation and Reduction.- II. Individual Reactions of the RPP Cycle.- III. Stoichiometry and Energetics.- C. Utilization of the Products of the RPP Cycle.- I. Starch Synthesis.- II. Triose Phosphate Export.- III. Glycolate Formation.- D. Mapping the RPP Cycle.- I. Early History.- II. First Products of CO2 Fixation.- III. Sugar Phosphates.- IV. Studies of Light-Dark and High-Low CO2 Transients.- V. Discovery of Enzymes of the RPP Cycle.- E. Metabolic Regulation of the RPP Cycle.- I. In Vivo Kinetic Steady-State Studies with Labeled Substrates.- II. Light-Dark Regulation.- III. Regulation of the RPP Cycle During Photosynthesis.- F. Concluding Remarks.- References.- 2. The Isolation of Intact Leaf Cells, Protoplasts and Chloroplasts.- A. Introduction.- B. Isolation of Plant Leaf Cells and Protoplasts.- I. Mechanical Methods.- II. Enzymic Methods.- III. Cell and Protoplast Isolation from C3 and C4 Grasses.- C. Isolation of Intact Chloroplasts.- I. Plant Material and Media.- II. Isolation Methods.- III. Chloroplast Isolation from Other Plants.- References.- 3. Studies with the Reconstituted Chloroplast System.- A. Reconstituted Chloroplast Systems.- I. Introduction.- II. Definition.- III. Methods of Preparation.- IV. Activities Achieved.- V. Advantages and Drawbacks.- B. Factors Affecting the Activity of Partial Reaction Sequences in Reconstituted Chloroplast Systems.- I. The Light Reactions.- II. The Conversion of 3-Phosphoglycerate to Triose Phosphate.- III. The Conversion of Triose Phosphate to Pentose Phosphate.- IV. The Conversion of Ribulose-5-Phosphate to Ribulose-1, 5-Bisphosphate.- V. The Fixation of Carbon Dioxide.- VI. Autocatalysis.- C. Reconstituted Chloroplast Systems and the Regulation of Photosynthesis.- I. The Role of Magnesium, pH and Reductants.- II. The Role of the ATP/ADP Ratio.- References.- 4. Autotrophic Carbon Dioxide Assimilation in Prokaryotic Microorganisms.- A. Introduction.- B. Principles of Autotrophic Carbon Dioxide Assimilation in Prokaryotic Cells.- C. The Pathway of Carbon Dioxide Assimilation in Green Sulfur Bacteria.- D. The Pathway of Carbon Dioxide Assimilation in Purple Bacteria.- E. The Pathway of Carbon Dioxide Assimilation in Blue-Green Algae.- F. The Pathway of Carbon Dioxide Assimilation in Chemolithotrophic Bacteria.- G. Regulatory Aspects of Carbon Dioxide Assimilation in Prokaryotic Microorganisms.- References.- 5. Light-Enhanced Dark CO2 Fixation.- A. Light-Enhanced Dark CO2 Fixation in C3 Plants.- I. Introduction.- II. Capacity for Light-Enhanced Dark CO2 Fixation.- III. Products.- IV. RuBP, NADPH, and ATP Levels.- V. Fate of PGA.- VI. Higher Plants.- B. Light-Enhanced Dark CO2 Fixation in C4 Plants.- References.- II B. The C4 and Crassulacean Acid Metabolism Pathways.- 6. The C4 Pathway and Its Regulation.- A. Discovery of C4 Photosynthesis.- B. Kranz Leaf Anatomy.- I. Variations in Leaf Anatomy.- C. Environmental Regulation of C4 Photosynthesis.- I. Light Intensity.- II. CO2 Concentration.- III. O2 Concentration.- IV. Temperature.- V. Water.- VI. Salinity.- VII. Nitrogen Supply.- D. Biochemical Schemes for the C4 Pathway.- E. Regulation via Enzymatic and Metabolic Compartmentation into Leaf Cell Types.- F. Efficiency of C4 Leaf Photosynthesis.- I. CO2 Pools.- II. CO2 Trapping.- III. CO2 Fixation by Bundle Sheath Cells.- G. C3–C4 Intermediate Plants.- H. Criteria for the Presence of C4 Photosynthesis.- I. Conclusions in the Regulation of C4 Photosynthesis in Leaves.- References.- 7. C4 Metabolism in Isolated Cells and Protoplasts.- A. Introduction.- B. Three Groups of C4 Plants.- C. Localization of Enzymes of C4 Metabolism in C4 Plants.- I. Intercellular Localization.- II. Intracellular Localization.- D. Criteria for Intactness of Cellular Preparations.- I. Mesophyll Preparations.- II. Bundle Sheath Preparations.- E. Variations in C4 Metabolism.- I. Mesophyll Cells of C4 Plants.- II. Bundle Sheath Cells of C4 Plants.- F. Energetics in C4 Metabolism.- G. Future Studies on C4 Metabolism with Cells and Protoplasts.- I. Transport Studies.- II. Screening for Inhibitors of C4 Photosynthesis.- References.- 8. The Flow of Carbon in Crassulacean Acid Metabolism (CAM).- A. Introduction.- B. Basic Phenomena of CAM.- C. The Metabolic Sequences of CAM.- I. The Flow of Carbon During the Night.- II. The Flow of Carbon During the Day.- D. Regulation of Carbon Flow in CAM.- E. Conclusions.- References.- 9. CAM: Rhythms of Enzyme Capacity and Activity as Adaptive Mechanisms.- A. Introduction.- B. Endogenous Versus Nonendogenous Rhythms: A Necessary Distinction.- C. Enzyme Capacity and Enzyme Activity: Two Distinct Levels of Control.- D. Rhythms Connected with CAM.- I. Components of the Malate Rhythm.- II. CO2 Uptake and CO2 Output.- III. PEP Carboxylase.- IV. Malate Dehydrogenase.- V. Malic Enzyme (NADP).- VI. Aspartate Aminotransferase.- VII. Enzymes of the Glycolytic Pathway.- VIII. Tricarboxylic Acid Cycle.- E. Working Hypothesis and Models.- I. Seasonal Adaptation.- II. Timing CAM.- References.- 10. ?13C as an Indicator of Carboxylation Reactions.- A. Introduction.- B. Carbon Isotope Fractionation and Its Measurement.- C. Variation in ?13C Values Between Plants.- I. Discrimination Caused by the Photosynthetic Pathway.- II. Variation in ?13C Values Between Plant Varieties and Species.- III. Variation in ?13C Values Within a Plant.- IV. Fractionation Associated with Carboxylation Enzymes.- V. Compartmental Organisation and Isotope Fractionation.- VI. Respiration.- D. Environmental Effects on the ?13C Value of Plants.- I. Temperature.- II. Carbon Dioxide Concentration.- III. Oxygen Concentration Effects on Discrimination.- IV. Light Level.- V. Availability of Water.- VI. Salinity and Carbon Isotope Fractionation.- E. Implications of Variation in ?13C Values Among Plant Species.- I. Natural Products.- II. Paleoecology.- III. Physiology — Plant, Animal, and Human.- F. Conclusions.- References.- II C. Factors Influencing CO2 Assimilation.- 11. Interactions Between Photosynthesis and Respiration in Higher Plants.- A. Introduction.- I. The Relevance of Photosynthetic and Respiratory Interactions.- B. Physiological Observations.- I. Plants with C3-Type Photosynthesis.- II. Plants with C4-Type Photosynthesis.- C. Biochemical Observations.- I. Plants with C3-Type Photosynthesis.- II. Plants with C4-Type Photosynthesis.- D. General Conclusions.- References.- 12. The Interaction of Respiration and Photosynthesis in Microalgae.- A. Introduction.- B. The Kok Effect.- C. Electron Transport Mechanisms for the Kok Effect.- I. General Considerations.- II. Prokaryotes.- III. Eukaryotes.- D. The Interaction of Oxygen with the Photosynthetic Electron Transport Chain.- E. Metabolically Mediated Control of Oxygen Uptake.- F. Synopsis.- References.- 13. Effect of Light Quality on Carbon Metabolism.- A. Introduction.- B. Principal Effects of Blue and Red Light on Carbon Metabolism.- C. Specific Features of Blue Light Action on Carbon Metabolism.- I. In the Absence of Photosynthesis.- II. In the Presence of Photosynthesis.- D. Direct Regulation of Certain Enzymes by Blue Light in Vitro and Its Possible Realization in Vivo.- E. Long-Term Effects of Light Quality on Biosyntheses and Chloroplast Organization.- F. Conclusion.- References.- 14. Photoassimilation of Organic Compounds.- A. Introduction.- B. Definitions.- C. Pathways and Products of Photometabolism.- D. Photo assimilation of Acetate.- E. Photo assimilation of Glucose.- References.- 15. Biochemical Basis of Ecological Adaptation.- A. Introduction.- B. Biochemical Variations in C3 Plants.- C. Biochemical Adaptation of CAM and C4 Plants.- I. Adaptive Value of C4 Metabolism.- D. Induced Variations in Carbon Fixation Pathways.- I. Effects of Age, CO2-Concentration, and Nitrogen Nutrition.- II. Effect of NaCl.- E. Concluding Remark.- References.- II D. Regulation and Properties of Enzymes of Photosynthetic Carbon Metabolism.- 16. Light-Dependent Changes of Stromal H+ and Mg2+ Concentrations Controlling CO2 Fixation.- A. Background.- B. Measurement of the pH in the Stroma and the Thylakoid Space of Intact Spinach Chloroplasts.- C. pH Dependence of CO2-Fixation.- D. Measurement of the Stromal Mg2+ Concentration in Intact Spinach Chloroplasts.- E. Mg2+ Dependence of CO2 Fixation.- F. Concluding Remarks.- References.- 17. Ribulose-1,5-Bisphosphate Carboxylase.- A. Fraction-1-Protein and RuBP Carboxylase.- B. Molecular Structure of RuBP Carboxylase.- C. Reaction Mechanism and Regulation.- D. RuBP Oxygenase.- E. Biosynthesis of RuBP Carboxylase.- References.- 18. Carbonic Anhydrase.- A. Introduction.- B. Characterization.- I. Histochemical and Other Detection.- II. Occurrence.- III. Location.- IV. Levels of Activity.- V. Isolation and Purification.- VI. Enzymic Parameters.- C. Function.- I. Chloroplast Envelope Membrane Permease.- II. Carbonic Anhydrase — RuBP Carboxylase Complex.- III. Proton Source, Buffering Capacity and Ionic Flux Regulation.- References.- 19. Enzymes of the Reductive Pentose Phosphate Cycle.- A. Introduction.- B. Characteristics of Regulatory Enzymes.- C. Activities and Location of Calvin Cycle Enzymes.- D. Glycerate-3-P Kinase.- E. Glyceraldehyde-3-P Dehydrogenase.- F. Triose-P Isomerase and Aldolase.- G. Fructosebisphosphatase and Sedoheptulosebisphosphatase.- H. Transketolase, Pentose-P Epimerase, and Pentose-P Isomerase.- I. Ribulose-5-P Kinase.- J. Concluding Remarks.- References.- 20. Enzymes of C4 Metabolism.- A. Introduction.- B. Isolation of Enzymes from Tissues of C4 Plants.- C. Carboxylation — PEP Carboxylase.- I. General Characteristics.- II. Physical Properties and Kinetics.- III. Regulation, Activation and Inhibition.- D. Formation of C4 Acids by Reduction and Transamination.- I. Reduction.- II. Transamination.- E. Decarboxylation.- I. NADP-Malic Enzyme (E.C.1.1.1.40).- II. NAD-Malic Enzyme (E.C.1.1.1.39).- III. PEP Carboxykinase (E.C.4.1.1.49).- F. Substrate Regeneration.- I. Pyruvate Pi Dikinase (E.C.2.7.9.1).- II. Alanine Aminotransferase (E.C.2.6.1.2).- G. Summary.- References.- 21. Enzymes of Crassulacean Acid Metabolism.- A. Introduction.- B. Enzymes of Starch Metabolism.- C. Glycolytic Enzymes.- D. Gluconeogenic Enzymes.- E. Carboxylating Enzymes.- I. The Formation of Malate.- II. The Photosynthetic Fixation of CO2.- F. Decarboxylating Enzymes.- I. The Decarboxylation of Malate.- II. The Decarboxylation of Oxaloacetate.- G. Respiratory Enzymes.- H. Conclusion.- References.- 22. Interaction Between Photochemistry and Activity of Enzymes.- A. Introduction.- B. Light-Mediated Modulation.- I. Occurrence.- II. Metabolic Significance.- III. Mechanism.- IV. Special Cases.- C. Dark Modulation.- D. Thylakoid-Generated Effectors.- I. pH.- II. Mg2+.- III. Energy Charge.- E. Conclusion.- References.- II E. Metabolism of Primary Products of Photosynthesis.- 23. Metabolism of Starch in Leaves.- A. Introduction.- B. Starch Biosynthesis.- I. Reactions Involved in Starch Biosynthesis.- II. The Predominant Pathway of Starch Synthesis.- III. Regulation of Starch Biosynthesis.- IV. Properties of the ADPglucose: 1,4-?-D-glucan 4-? Glucosyltransferase.- V. ?-1,4-Glucan: ?-1,4-Glucan 6-Glycosyl Transferase (Branching or Q Enzyme).- VI. Remaining Problems in Starch Synthesis.- C. Starch Degradation.- I. Reactions Involved in Starch Degradation.- II. Degradation of Intact Granules in Vitro.- III. Starch Degradation in Vivo: Germinating Seeds.- IV. Starch Degradation in Vivo: Leaves.- References.- 24. The Enzymology of Sucrose Synthesis in Leaves.- A. Introduction.- B. Physiological Relationships of Sucrose in Leaves.- C. Enzymology.- I. Sucrose Phosphate Synthetase E.C.2.4.1.14 and Sucrose Synthetase E.C.2.4.1.13.- II. Sucrose Phosphatase (E.C.3.1.3.24).- III. UDPglucose Pyrophosphorylase (E.C.2.7.7.9).- IV. Sucrose Phosphorylase (E.C.2.4.1.7).- V. Invertase (E.C.3.2.1.26).- VI. Enzyme Control Mechanisms.- D. Intracellular and Intercellular Site of Sucrose Synthesis in Leaves.- I. Chloroplast or Cytoplasm?.- II. Intercellular Localization of Sucrose Synthesis in C4 Plants.- III. Intercellular Distribution Between Cells Containing Chlorophyll and Vascular Tissue.- References.- II F. Glycolic Acid and Photorespiration.- 25. Glycolate Synthesis.- A. Introduction: Glycolate Formation, Photorespiration and the Warburg Effect.- B. Environmental Factors Affecting Glycolate Synthesis.- C. Mechanisms of Glycolate Formation.- I. Reductive Glycolate Formation.- II. Oxidative Glycolate Synthesis.- D. Photosynthetic Glycolate Formation in Vivo; Which Reaction Predominates?.- I. Glycolate Synthesis by C3 Plants.- II. Glycolate Formation by C4 Plants.- E. Conclusion: The Inhibition of Glycolate Formation by Some Common Metabolites — an Open Question.- References.- 26. Glycolate Metabolism by Higher Plants and Algae.- A. Introduction.- B. Glycolate Biosynthesis.- I. Properties of Ribulose-P2 Carboxylase/Oxygenase for Phosphoglycolate Biosynthesis.- II. Phosphoglycolate Phosphatase and Phosphoglycerate Phosphatase.- C. Glycolate Pathway.- I. Pathways in Peroxisomes.- II. Mitochondrial Interconversion of Glycine and Serine.- D. O2 and CO2 Exchange and Energy Balance.- I. Sites of O2 Uptake and CO2 Release in the Glycolate Pathway.- II. O2 Uptake During Photosynthesis.- III. Energy Balance.- E. Leaf Peroxisomal Membrane and Transport.- F. Glycerate and Sucrose from Glycolate.- G. The Glycolate Pathway in Algae.- I. Introduction.- II. Glycolate Excretion.- III. Glycolate Dehydrogenase.- IV. Glycerate-Serine Pathway in Algae.- References.- 27. Photorespiration: Studies with Whole Tissues.- A. Discovery of Photorespiration.- B. Assays of Photorespiration in Leaves.- I. Post-Illumination CO2 Outburst.- II. Inhibition of Net CO2 Assimilation by Oxygen.- III. CO2 and 14CO2 Efflux in CO2-Free Air.- IV. Short-Time Uptake of 14CO2 and 12CO2.- V. The Magnitude of Photorespiration in Leaves.- C. Photorespiration in Algae and Submerged Aquatic Plants.- D. Photorespiration in Callus, Isolated Plant Cells, and Protoplasts.- E. The Control of Photorespiration.- I. The Energetics and Possible Origins of Photorespiration.- II. Biochemical Inhibition of Glycolate Oxidation.- III. Biochemical Inhibition of Glycolate Synthesis.- IV. The Metabolic Regulation of Photorespiration.- References.- 28. Photorespiration: Comparison Between C3 and C4 Plants.- A. Introduction.- B. Terminology and Perception.- C. Measurement of Photorespiration.- D. Characteristics of Photorespiration in C3 Plants.- I. Rates of Photorespiration.- II. The Post-Illumination Burst.- III. Compensation Point.- IV. Effect of CO2 Concentration.- V. Effect of O2.- VI. Effect of Temperature.- VII. Interaction of Oxygen, Carbon Dioxide, and Temperature.- VIII. Effect of Light Intensity.- IX. The Glycolate Pathway.- E. Photo respiration in C4 Plants.- I. Photorespiration as CO2 Evolution.- II. Photorespiration as Oxygen Uptake.- III. Photorespiration in C4 Plants — Indirect Evidence.- IV. Evidence Against Photorespiration in C4 Plants.- F. Concluding Remarks.- References.- III. Ferredoxin-Linked Reactions.- 1. Transhydrogenase.- A. Introduction and Definitions.- B. Soluble Flavoproteins with Transhydrogenase Activity.- I. Bacterial Enzymes.- II. Ferredoxin-NAD(P)+ Reductases.- C. Membrane-Bound (Particulate) Transhydrogenases.- I. Mitochondrial and Bacterial Transhydrogenases; General Aspects.- II. Transhydrogenase of Photosynthetic Bacteria.- References.- 2. Oxygen Activation and Superoxide Dismutase in Chloroplasts.- A. Introduction.- B. Principles of Oxygen Activation.- C. Superoxide Anion and Superoxide Dismutase.- I. Dismutation of the Superoxide Anion (O.-2); Superoxide Dismutase in Plants.- II. Superoxide Dismutase in Chloroplasts.- III. Monovalent Oxygen Reduction in Chloroplasts.- D. Determination of the Products of Oxygen Reduction.- E. Possible Functions of Reduced Oxygen Species in Chloroplasts.- I. Desaturation of Fatty Acids.- II. Hydroxylation of Aromatic Compounds.- III. Photorespiration.- IV. Ethylene Formation.- F. Conclusions.- References.- 3. Ferredoxin-Linked Carbon Dioxide Fixation in Photosynthetic Bacteria.- A. Introduction.- B. Ferredoxin-Linked Carboxylation Reactions.- I. Synthesis of Pyruvate.- II. Synthesis of ?-Ketoglutarate.- III. Synthesis of ?-Ketobutyrate.- IV. Synthesis of Phenylpyruvate.- V. Synthesis of ?-Ketoisovalerate.- VI. Synthesis of Formate.- C. The Reductive Carboxylic Acid Cycle.- D. Concluding Remarks.- References.- 4. Reduction of Nitrate and Nitrite.- A. Introduction.- B. Reduction of Nitrate to Nitrite.- I. Assimilatory Nitrate Reductase of Eukaryotes.- II. Assimilatory Nitrate Reduction in Prokaryotes.- C. Reduction of Nitrite to Ammonia.- I. Nitrite Reductase of Photosynthetic Cells.- II. Nitrite Reductase of Nonphotosynthetic Cells.- D. Control of Nitrate Reduction.- I. Synthesis and Degradation of Enzymes.- II. Utilization of Nitrate.- III. Reversible Inactivation of Nitrate Reductase.- IV. Localization of Enzymes and Effect of Light and Carbohydrate on Nitrate Utilization.- V. Conclusions.- References.- 5. Photosynthetic Ammonia Assimilation.- A. Introduction.- B. Photosynthetic Ammonia Assimilation in Intact Organisms.- C. Localization of Enzymes Involved in Ammonia Assimilation.- I. Glutamate Dehydrogenase.- II. Glutamine Synthetase.- III. Glutamate Synthase (GOGAT).- D. Photosynthetic Ammonia Assimilation in Isolated Intact Chloroplasts.- E. Photorespiratory Ammonia Evolution and Reassimilation.- F. Conclusions.- References.- 6. N2 Fixation and Photosynthesis in Microorganisms.- A. Introduction.- B. Distribution of Nitrogenase Among Photosynthetic Prokaryotes.- C. Oxygen Sensitivity and Protection of Algal Nitrogenase.- D. Requirements for an Active Nitrogenase.- E. The Provision of Reductant and ATP in Photosynthetic Prokaryotes.- I. Electron Donation.- II. The Production of ATP.- F. The Nitrogen-Fixing System of Heterocysts of Anabaena cylindrica.- G. Nitrogenase and Its Possible Regulation by Glutamine Synthetase.- References.- 7. Symbiotic N2 Fixation and Its Relationship to Photosynthetic Carbon Fixation in Higher Plants.- A. Introduction.- B. Relationship of N2 Fixation to Carbon Assimilation.- I. Nitrogenase.- II. ATP and Reductant.- III. Ammonia Assimilation.- IV. Photosynthate as the Limiting Factor.- V. Translocation and Partitioning of Nitrogen and Carbon Assimilates.- References.- 8. Photosynthetic Assimilation of Sulfur Compounds.- A. Introduction.- B. Observations with Whole Organisms.- C. Observations with Isolated Organelles.- D. Cell-Free Systems.- I. Sulfate Activation and Degradation of Active Sulfate.- II. Transfer of Sulfate from Sulfonucleotides for Further Reduction.- III. Reduction to the Level of Sulfide.- IV. Biosynthesis of Cysteine.- E. Regulation of Assimilatory Sulfate Reduction in Photosynthetic Organisms.- References.- 9. Hydrogen Metabolism.- A. Introduction.- B. Hydrogenase.- I. Occurrence of Hydrogenase in Photosynthetic Cells.- II. Adaptation and Deadaptation.- III. Cell-Free Preparations of Hydrogenase.- C. Evolution of H2.- I. Dark Evolution of H2.- II. H2 Photoevolution.- III. H2 Evolution by Blue-Green Algae.- D. Consumption of H2.- I. Dark Absorption of H2.- II. Photoreduction.- References.- Author Index.