A rapid-equilibrium model for the control of the Calvin photosynthesis cycle by cytosolic orthophosphate.

1. A simple model based on rapid-equilibrium assumptions is derived which relates the steady-state activity of the Calvin cycle for photosynthetic carbohydrate formation in C3 plants to the kinetic properties of a single cycle enzyme (fructose bisphosphatase) and of the phosphate translocator which accounts for the export of photosynthate from the chloroplast. Depending on the kinetic interplay of these two catalysts, the model system may exhibit a single or two distinct modes of steady-state operation, or may be unable to reach a steady state. 2. The predictions of the model are analysed with regard to the effect of external orthophosphate on the steady-state rate of photosynthesis in isolated chloroplasts under conditions of saturating light and CO2. Due to the possible existence of two distinct steady states, the model may account for the stimulatory as well as the inhibitory effects of external phosphate observed in experiments with intact chloroplasts. Stability arguments indicate, however, that only the steady-state case corresponding to phosphate inhibition of the rate of photosynthesis could be of physiological interest. 3. It is concluded that chloroplasts under physiological conditions most likely operate in a high-velocity steady state characterized by a negative Calvin cycle flux control coefficient for the phosphate translocator. This means that any factor enhancing the export capacity of the phosphate translocator can be anticipated to decrease the actual steady-state rate of photosynthate export due to a decreased steady-state rate of cyclic photosynthate production.