Simulated Analysis of Influence of Changes in H+-ATPase Activity and Membrane CO2 Conductance on Parameters of Photosynthetic Assimilation in Leaves

Abstract

Photosynthesis is an important process in plants which influences their development and productivity. Many factors can control the efficiency of photosynthesis, including CO₂ conductance of leaf mesophyll, which affects the CO₂ availability for Rubisco. It is known that electrical stress signals can decrease this conductance, and the response is probably caused by inactivation of H⁺-ATPase in the plasma membrane. In the current work, we analyzed the influence of both CO₂ conductance in the plasma membrane, and chloroplast envelopes and H⁺-ATPase activity on photosynthetic CO₂ assimilation, using a two-dimensional mathematical model of photosynthesis in leaves. The model included a description of assimilation on the basis of the Farquhar-von Caemmerer-Berry model, ion transport through the plasma membrane, diffusion of CO₂ in the apoplast, and transport of CO₂ through the plasma membrane and chloroplast envelope. The model showed that the photosynthetic CO₂ assimilation rate was mainly dependent on the plasma membrane and chloroplast envelope conductance; direct influence of the H⁺-ATPase activity (through changes in pH and CO₂/HCO₃^- concentration ratio) on this rate was weak. In contrast, both changes in CO₂ conductance of the plasma membrane and chloroplast envelopes and changes in the H⁺-ATPase activity influenced spatial heterogeneity of the CO₂ assimilation on the leaf surface in the simulated two-dimensional system. These effects were also observed under simultaneous changes in the CO₂ conductance of the plasma membrane and H⁺-ATPase activity. Qualitatively similar influence of changes in the CO₂ conductance of the plasma membrane and chloroplast envelopes, and changes in the H⁺-ATPase activity on photosynthesis were shown for two different densities of stomata in the simulated leaf; however, lowering the density of stomata decreased the assimilation rate and increased the heterogeneity of assimilation. The results of the model analysis clarify the potential influence of H⁺-ATPase inactivation on photosynthesis, and can be the basis for development of new methods for remote sensing of the influence of electrical signals.

Keywords: H+-ATPase; chloroplast envelope CO2 conductance; electrical signals; photosynthetic CO2 assimilation; plasma membrane CO2 conductance; spatial heterogeneity; two-dimensional photosynthetic model.