Biochemical Models of Leaf Photosynthesis

By S von Caemmerer
Biochemical Models of Leaf Photosynthesis

Increasing concerns of global climate change have stimulated research interests in all aspects of carbon exchange. This has restored interest in leaf photosynthetic models to predict and assess changes in photosynthetic CO2 assimilation in different environments.

This is a comprehensive presentation of the most widely used models of steady-state photosynthesis by an author who is a world authority. Treatments of CO3, CO4 and intermediate pathways of photosynthesis in relation to environment have been update to include work on antisense transgenic plants. It will be a standard reference for the formal analysis of photosynthetic metabolism in vivo by advanced students and researchers.

  1. Page vii
    Preface
  2. Page x
    Acknowledgments
  3. Page 1
    1. The kinetics and regulation of rubisco
    1. Page 1
      Introduction
    2. Page 2
      Kinetics of fully activated rubisco
      1. Page 2
        Definition of kinetic constants
      2. Page 3
        Kinetics at saturating RuBP
      3. Page 4
        Relative specificity, Sc/o
      4. Page 6
        Rubisco kinetic constants in vitro and in vivo
      5. Page 10
        RuBP-limited rate of carboxylation
      6. Page 11
        High rubisco site concentrations in the chloroplast
      7. Page 14
        The binding of phosphorylated compounds and other ligands
    3. Page 16
      Activation of rubisco
      1. Page 17
        Activation of rubisco in the absence of RuBP
      2. Page 17
        Activation of rubisco in the presence of RuBP
      3. Page 18
        The carbamylation ratio of rubisco
      4. Page 19
        Rubisco activase
        1. Page 20
          A model of activase action
        2. Page 21
          A hyperbolic response of carbamylation to activase activity
        3. Page 22
          Binding of other ligands
        4. Page 22
          An unknown function of activase
    4. Page 24
      Derivations
      1. Page 24
        Derivation of rate equations for activated enzyme
      2. Page 25
        The rate equations at high concentration of enzyme sites
      3. Page 26
        Derivation of rate equations of activation
      4. Page 27
        Carbamylated enzyme sites as a fraction of total enzyme sites
      5. Page 28
        Ligands other than RuBP
  4. Page 29
    2. Modelling C3 photosynthesis
    1. Page 29
      Introduction
    2. Page 29
      A few simplifying assumptions
    3. Page 30
      Stoichiometry of C3 photosynthesis
      1. Page 30
        Carbon stoichiometry of the PCR and PCO cycles
      2. Page 31
        The carbon path from PGA to RuBP
      3. Page 32
        ATP and NADPH consumption in the PCR and PCO cycles
      4. Page 32
        Thylakoid reactions
        1. Page 32
          Production of NADPH and ATP
        2. Page 34
          Light dependence of electron transport rate
      5. Page 35
        Respiration
    4. Page 35
      Rate equations for CO2 assimilation
      1. Page 35
        RuBP-saturated CO2 assimilation rate
      2. Page 36
        RuBP-limited CO2 assimilation rate
      3. Page 38
        Export-limited CO2 assimilation rate
      4. Page 39
        Summary of rate equations
    5. Page 39
      CO2 partial pressure in the chloroplast
    6. Page 42
      Parameterization of the Model
      1. Page 42
        Values at 25°C
      2. Page 44
        Temperature dependencies
        1. Page 44
          Rubisco and Rd
        2. Page 45
          Temperature dependence of Jmax
    7. Page 47
      The CO2 compensation point
      1. Page 48
        Γ in the absence of Rd
      2. Page 48
        Γ in the presence of Rd
      3. Page 49
        Light and temperature dependence
      4. Page 50
        Measurements of Γ*
    8. Page 51
      CO2 response curves
      1. Page 52
        CO2 response curves in transgenic tobacco with impaired photosynthesis
      2. Page 54
        CO2 assimilation rate at different O2 partial pressures
      3. Page 55
        CO2 assimilation rate at different irradiances
      4. Page 56
        CO2 assimilation rate at different temperatures
      5. Page 57
        The initial slope of the CO2 response curve, dA/dCi
    9. Page 58
      Light response curves
      1. Page 58
        Dependence on O2 and CO2 partial pressures
      2. Page 60
        Quantum yield
    10. Page 62
      Temperature responses
    11. Page 62
      Does the activation state of rubisco need to be incorporated into models of CO2 assimilation?
    12. Page 65
      Long-term effect of environment on photosynthesis
      1. Page 65
        Vcmax and Jmax and the transition from rubisco to electron transport limitation
      2. Page 68
        Effect of growth temperature
      3. Page 68
        Growth at elevated CO2
    13. Page 70
      Engineering a better rubisco
  5. Page 72
    3. Chlorophyll fluorescence and oxygen exchange during C3 photosynthesis
    1. Page 72
      Introduction
    2. Page 72
      Chlorophyll fluorescence and chloroplast electron transport
      1. Page 73
        Calculating electron transport rate from chlorophyll fluorescence
      2. Page 74
        Estimating electron transport rate, JA, from CO2 assimilation rate
      3. Page 76
        Relationship between Jf/(absI) and Fs/Fm
      4. Page 78
        Calculation of mesophyll conductance to CO2 diffusion
      5. Page 79
        Estimation of rubisco oxygenation rate from Jf and JA
    3. Page 80
      Oxygen exchange during C3 photosynthesis
      1. Page 80
        Introduction
      2. Page 81
        Basic equations
        1. Page 81
          Total oxygen evolution
        2. Page 82
          Total oxygen uptake
        3. Page 83
          The Mehler ascorbate peroxidase (MAP) reaction
        4. Page 85
          Net O2 and CO2 exchange
        5. Page 86
          Estimation of rubisco carboxylation and oxygenation rates
      3. Page 87
        The CO2 dependence of O2 exchange
      4. Page 88
        The O2 dependence of O2 exchange
      5. Page 89
        The O2 exchange at the compensation point
      6. Page 90
        The temperature dependence of O2 exchange
  6. Page 91
    4. Modelling C4 photosynthesis
    1. Page 91
      Introduction
    2. Page 91
      Basic model equations
      1. Page 94
        Equations for enzyme-limited photosynthesis
        1. Page 94
          CO2 assimilation rate in the bundle sheath
        2. Page 94
          Bundle-sheath CO2 partial pressure
        3. Page 94
          Bundle-sheath O2 partial pressure
        4. Page 95
          The rate of PEP carboxylation
        5. Page 95
          Quadratic expression for the enzyme-limited CO2 assimilation rate
      2. Page 96
        Light- and electron-transport-limited photosynthesis
        1. Page 96
          Rates of ATP and NADPH consumption
        2. Page 97
          Partitioning of electron transport rate between C3 and C4 cycles
        3. Page 98
          Light dependence of electron transport rate
        4. Page 98
          Quadratic expression for electron-transport-limited CO2 assimilation rate
      3. Page 99
        Summary of equations
    3. Page 99
      Analysis of the model
      1. Page 99
        Parameterization of the model
      2. Page 101
        The model at high irradiance
        1. Page 101
          PEP carboxylase and rubisco activity
        2. Page 102
          Bundle-sheath conductance
        3. Page 103
          Variation of bundle-sheath conductance, PEP carboxylase activity and rubisco with leaf age
        4. Page 105
          CO2 response curves
        5. Page 109
          Oxygen sensitivity of C4 photosynthesis
        6. Page 110
          CO2 compensation point
        7. Page 113
          CO2 diffusion from intercellular air spaces to the mesophyll cytosol
      3. Page 116
        CO2 fixation at limiting light
        1. Page 116
          Optimal partition of electron transport
        2. Page 118
          Leakiness
        3. Page 119
          Quantum yield
      4. Page 120
        Modelling different decarboxylation types
        1. Page 120
          Bundle-sheath conductance
        2. Page 121
          O2 evolution in the bundle sheath
        3. Page 122
          Energy requirements
  7. Page 123
    5. Models of C3-C4 intermediate photosynthesis
    1. Page 123
      Introduction
    2. Page 124
      Basic model equations
    3. Page 125
      Equations for photosynthesis at high irradiance
      1. Page 125
        Rubisco-limited CO2 assimilation rate
      2. Page 126
        Net O2 evolution in the bundle sheath
      3. Page 126
        Quadratic expression for CO2 assimilation rate
    4. Page 127
      Parameterization and analysis of the model at high irradiance
      1. Page 127
        Kinetic Constants
      2. Page 127
        Respiration
      3. Page 128
        Partitioning of photorespiration
      4. Page 128
        Bundle-sheath conductance
    5. Page 129
      The glycine shuttle
      1. Page 130
        Bundle-sheath rubisco activity and conductance
      2. Page 131
        CO2 compensation point
      3. Page 132
        CO2 and O2 responses of CO2 assimilation rate
    6. Page 133
      The contribution of C4 photosynthesis
      1. Page 134
        CO2 compensation point
      2. Page 135
        CO2 and O2 responses of CO2 assimilation
    7. Page 136
      Energy requirements of C3-C4 photosynthesis
      1. Page 136
        Rates of ATP and NADP consumption
      2. Page 137
        Electron transport rate
      3. Page 137
        Light-dependent CO2 assimilation rate
    8. Page 139
      Conclusion
  8. Page 140
    6. Concluding remarks
  9. Page 152
    List of Symbols
  10. Page 145
    References

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