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Canopy Photosynthesis: From Basics to Applications pp 61–100Cite as

Modeling Leaf Gas Exchange

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Part of the book series: Advances in Photosynthesis and Respiration ((AIPH,volume 42))

Summary

Leaves are photosynthetic organs that absorb light and convert the photon energy of light to chemical energy for use in CO2 assimilation. Here we review how CO2 assimilation rates vary, depending on environmental factors and among leaves. Net CO2 assimilation is a balance between the carboxylation of ribulose 1,5-bisphosphate (RuBP) catalyzed by ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) and the release of CO2 by photorespiration and mitochondrial respiration. The steady-state biochemical model of CO2 assimilation considers photosynthetic metabolism as a composite of two processes, namely, RuBP carboxylation and regeneration. The former, modeled based on the Rubisco kinetics, is limited mainly by CO2 supply, whereas the latter is assumed to be limited by the rate of photon absorption at low light and by its use in electron transport at high light. CO2 concentration at the assimilation sites in chloroplasts depends on the stomatal and mesophyll conductances for CO2 diffusion. Both these conductances are sensitive to environmental variables, but no mechanistic models of environmental responses for these conductances are available. Various empirical models have been developed and combined with the biochemical photosynthesis model allowing for expression of CO2 assimilation rates as a function of environmental variables.

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Abbreviations

A :

Net CO2 assimilation rate

A c :

RuBP -saturated A

A j :

RuBP -limited A

A max :

Maximum A (photosynthetic capacity)

A t :

A limited by TP use

ABA :

Abscisic acid

AOX :

Cyanide-resistant alternative oxidase

B n :

Net anabolic NADH supply

C a C c , C f and C i :

CO2 partial pressures at air, chloroplast , leaf surface and intercellular space respectively

c p :

Heat capacity of air

CP:

Cytochrome pathway

D :

Leaf-to-air vapor pressure deficit

D 0 :

Fitted parameter for D in Eq. 3.29

d min :

Minimum ion diffusion rate in modeling stomatal conductance in Eq. 3.36

E :

Evapotranspiration rate

E a :

Activation energy

E o :

Activation energy of the respiratory pathway

E oref :

Activation energy at the reference temperature

f i and f ref :

Value of f at T = ∞ and the reference temperature

f m :

Fraction of photorespiratory NADH that remains in mitochondria

F v /F m :

Maximum dark-adapted quantum yield for photosystem I I

f Ψv :

Sensitivity of stomata to leaf water potential

FADH2 :

Flavin adenine dinucleotide

g :

Total diffusion conductance for CO2 from air to chloroplasts

g 0 and g 1 :

Fitted parameters for stomatal conductance

g b g m , and g s :

Conductances for CO2 diffusion across boundary layer, mesophyll and stomata

g bh :

Leaf boundary layer for heat conductance

g c :

Leaf CO2 conductance including stomatal and boundary layer conductance s

g wc :

Leaf water vapor conductance including stomatal and boundary layer conductance s

H :

Sensible heat flux

H d :

Energy of deactivation

h r :

Relative humidity at the leaf surface

HT:

High temperature

I :

Photosynthetically active photon flux density (irradiance )

J :

Electron transport rate

J max :

Maximum J

K c and K o :

Michaelis–Menten constant s for carboxylation and oxygenation

k cat :

Rubisco turnover rate (rate of CO2 fixation per Rubisco active site)

K tot :

Hydraulic conductance for the whole plant

LMA:

Leaf mass per area

LT:

Low temperature

m :

Empirical constant in Eq. 3.32

M :

Plant dry mass

MDH:

Malate dehydrogenase

NADH m :

Fraction of photorespiratory NADH that remains in mitochondria

NDs:

Type II NAD(P)H dehydrogenases

O :

O2 partial pressure in chloroplasts

OAA:

Oxaloacetic acid

OPPP:

Oxidative pentose phosphate pathway

P g and P e :

Pressure potentials of the guard cells and the bulk epidermal cells

P p :

Export rate of triose phosphate

PDH:

Pyruvate dehydrogenase complex

PEPCase:

Phosphoenolpyruvate carboxylase

PGA :

3-phosphoglycerate

PQ:

Plastoquinone

PSII :

Photosystem I I

Q 10 :

Ratio of the process rate at a reference temperature + 10 K to the rate at the reference temperature

r :

Total resistance to CO2 diffusion from air to chloroplasts

r b , r m , and r s :

Resistances to CO2 diffusion at boundary layer, mesophyll, and stomata

r wb and r ws :

Resistances to water vapor at boundary layer and stomata

R :

Universal gas constant

R C :

Rate of non-photorespiratory CO2 release

R d :

Respiration rate in the light

R g :

Growth respiration coefficient

R m :

Maintenance respiration rate

R n :

Respiration rate in the dark

R N :

Net radiation

R O :

Rate of non-photorespiratory O2 consumption

R PR :

Photorespiratory rate

R ref :

Respiration rate at the reference temperature

R min :

Minimum R d

RGR :

Relative growth rate

Rubisco :

Ribulose-1,5-bisphosphate carboxylase/oxygenase

RuBP :

Ribulose 1,5-bisphosphate

S c :

Chloroplast surface area

S c/o :

Relative specificity of Rubisco (specificity factor)

S V :

Sensitivity parameter in Eq. 3.38

SA and SD:

Starch-accumulating and deficient species

TP:

Triose-phosphate

T l and T a :

Leaf and air temperatures

T k :

Temperature in Kelvin

T opt :

Optimal temperature

T ref :

Reference temperature

T sky :

Temperature of the sky

UCP:

Uncoupling protein

UQ:

Ubiquinone

V by :

Rate of carbon flow to CO2 as a by-product of flows into anabolic products

V c :

Rate of carboxylation

V cat :

Rate of CO2 release due to catabolic substrate oxidation

V cmax :

Maximum V c

V o :

Rate of oxygenation

V opc :

CO2 release rate from substrate oxidation via cytosolic OPPP

V omax :

Maximum V o

V opp :

CO2 release rate from substrate oxidation via chloroplastic OPPP

V px :

Photo-reductant export rate

VPD :

Vapor pressure deficit

W a and W i :

Water vapor pressure s in air and intercellular air space

α:

Leaf absorptance

α S and α IR :

Shortwave and infrared absorptance s

β a :

Empirical constant in Eq. 3.34

γ and ξ:

Empirical constants in Eq. 3.36

δ :

Coefficient to describe dynamic response of E o to temperature

δ w :

Thickness of mesophyll cell wall

ΔS :

Entropy term

ε:

Leaf long-wave emissivity

φ :

Ratio of RuBP oxygenation to carboxylation rates

ϕ x :

Initial slopes of the light response curve

Γ:

CO2 compensation point of CO2 assimilation

Γ*:

CO2 compensation point of CO2 assimilation in the absence of R d

λ :

Heat of vaporization

λ s :

Marginal water cost of carbon gain

π a :

Osmotic potential of apoplastic water near the stomatal guard cells

π e :

Osmotic potential of epidermis cells

π g :

Osmotic potential in the guard cells

θ x :

Curvature factors in Eqs. 3.11, 3.12 and 3.16

ρ :

Shortwave reflectance of the surroundings

σ :

Stefan–Boltzmann constant

τ :

Scale R o to a daily rate

τ a :

ATP concentration in the guard cells

υ :

Slope of light response of R d

χ :

Hydraulic conductance between the bulk epidermis and stomatal guard cells

ψ e :

Water potential of epidermis cell

ψ g :

Water potential of guard cell

ψref :

Reference potential

ψ s :

Soil water potential

ψ V :

Bulk leaf water potential

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Acknowledgements

We thank Ülo Niinemets for the valuable comments. This work was supported by Grants-in-Aid for Scientific Research on Innovative Areas (Nos. 21114001, 21114007, 21114009), by KAKENHI (Nos. 20677001, 25291095 and 25660113) and by CREST, JST, Japan.

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  1. Graduate School of Life Sciences, Tohoku University, Sendai, 980-8578, Japan

    Kouki Hikosaka

  2. CREST, JST, Tokyo, Japan

    Kouki Hikosaka

  3. School of Life Sciences, Tokyo University of Pharmacy and Life Sciences, 1432-1, Horinouchi, Hachioji, Tokyo, 192-0392, Japan

    Ko Noguchi

  4. School of Science, The University of Tokyo, Hongo, Bunkyo-ku, Tokyo, 113-0033, Japan

    Ichiro Terashima

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  1. Graduate School of Life Sciences, Tohoku University, Sendai, Japan

    Kouki Hikosaka

  2. Environmental Sciences, Estonian University of Life Sciences Institute of Agricultural and, Tartu, Estonia

    Ülo Niinemets

  3. Crop and Weed Ecology, Centre for Crop Systems Analysis, Wageningen University, Wageningen, Gelderland, The Netherlands

    Niels P.R. Anten

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Hikosaka, K., Noguchi, K., Terashima, I. (2016). Modeling Leaf Gas Exchange. In: Hikosaka, K., Niinemets, Ü., Anten, N. (eds) Canopy Photosynthesis: From Basics to Applications. Advances in Photosynthesis and Respiration, vol 42. Springer, Dordrecht. https://doi.org/10.1007/978-94-017-7291-4_3

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