Modelling (18)O2 and (16)O2 unidirectional fluxes in plants. IV: role of conductance and laws of its regulation in C3 plants.

Numerous studies focus on the measurement of conductances for CO2 transfer in plants and especially on their regulatory effects on photosynthesis. Measurement accuracy is strongly dependent on the model used and on the knowledge of the flow of photochemical energy generated by light in chloroplasts. The only accurate and precise method to quantify the linear electron flux (responsible for the production of reductive energy) is the direct measurement of O2 evolution, by (18)O2 labelling and mass spectrometry. The sharing of this energy between the carboxylation (P) and the oxygenation of photorespiration (PR) depends on the plant specificity factor (Sp) and on the corresponding atmospheric concentrations of CO2 and O2 (André, 2013). The concept of plant specificity factor simplifies the equations of the model. It gives a new expression of the effect of the conductance (g) between atmosphere and chloroplasts. Its quantitative effect on photosynthesis is easy to understand because it intervenes in the ratio of the plant specificity factor (Sp) to the specificity of Rubisco (Sr). Using this 'simple' model with the data of (18)O2 experiments, the calculation of conductance variations in response to CO2 and light was carried out. The good fitting of experimental data of O2 and CO2 exchanges confirms the validity of the simple model. The calculation of conductance variation during the increase of external CO2 concentration reveals a linear law of regulation between external and internal CO2 concentrations. During CO2 variations, the effects of g regulation tend to maintain a higher level of oxygenation (PR) in expense of a better carboxylation (P). Contrary to CO2, the variation of O2 creates a negative feedback effect compatible with a stabilization of atmospheric O2. The regulation of g amplifies this result. The effect of light in combination with CO2 is more complex. Below 800μmolquantam(-2)s(-1) the ratio PR/P is maintained unchangeable in expense of carboxylation efficiency. Above that irradiance value, PR/P increases dramatically. It appears that the saturation curves of photosynthesis under high light could be simply due to the regulation by the conductance g and not by any biochemical or biophysical limitation. In conclusion, the regulatory effect of conductance operates in a way that it preserves the rate of photorespiration. This confirms a positive and protective role of photorespiration at the biochemical, whole plant and atmosphere levels. Since the effects of photorespiration are linked to the properties of Rubisco, they add new arguments for a co-evolution of plant and atmosphere, including the evolution of CO2 conductance.