Kinetic modeling of the photosynthetic electron transport chain

A simulation model of the photosynthetic electron transport chain operating under steady state conditions is presented. The model enables the calculation of (1) the rates of electron transport and transmembrane proton translocation, (2) the proton/electron stoichiometry, (3) the number of electrons stored in the different redox centers and (4) the stationary transmembrane pH difference. Light intensity and proton permeability of the thylakoid membrane are varied in order to compare the predictions of the model with experimental data. The routes of electron transport and proton translocation are simulated by two coupled arithmetic loops. The first one represents the sequence of reaction steps making up the linear electron transport chain and the Q-cycle. This loop yields the electron flow rate and the proton/electron ratio. The second loop balances the H+ fluxes and yields the internal H+ concentration. The bifurcation of the electron transport pathways at the stage of plastoquinol oxidation is obligatory. The first electron enters always the linear branch and is transferred to photosystem I. The electron of the remaining semiquinone can enter the Q-cycle or, alternatively, the semiquinone can be lost from the cytochrome b6f complex. The competition between these two reactions explains the experimentally observed variability of the proton/electron ratio. We also investigated additional model variants, where the variation of the proton/electron stoichiometry is attributed to other loss reactions within the cytochrome b6f complex. However, the semiquinone detachment seems to be the best candidate for a satisfactory description of the experimental data. Additional calculations were done in order to assess the effects of the movement of the Rieske protein on linear electron transport; it was found that this conformational change does not limit the electron transport rate, if it occurs with a time constant of at least 1000 s(-1).