A POSSIBLE RATE-LIMITING FUNCTION OF CHLOROPLAST HEXOSEMONOPHOSPHATE ISOMERASE IN STARCH SYNTHESIS OF LEAVES

Levels of fructose 6-phosphate and glucose 6-phosphate were measured in chloroplasts which had been isolated non-aqueously from leaves of various plants. A large decrease in the ratio of glucose 6-phosphate to fructose 6-phosphate in the light indicated considerable displacement of the hexosephosphate isomerase reaction from equilibrium in leaves of spinach and red beet which were photosynthesizing at high rates. The decrease in the ratio of glucose 6-phosphate to fructose 6-phosphate was correlated with an increase in the chloroplastic level of 3-phosphoglyceric acid, which proved to be a competitive inhibitor of chloroplast hexosephosphate isomerase. Other metabolites, especially the product of the reaction, glucose 6-phosphate, and ions in concentrations as present in the stroma under natural conditions, cause a further reduction in the rate of the forward reaction of the hexosemonophosphate isomerase. When the concentration of O2 in air was decreased from 21 to 2%, both the rate of leaf photosynthesis and the ratio of glucose 6-phosphate to fructose 6-phosphate increased; the concentration of 3-phosphogylceric acid and starch synthesis decreased. The results are explained in terms of activation of ADPglucose pyrophosphorylase and of inhibition of hexosephosphate isomerase by 3-phosphogylceric acid. Hexosephosphate isomerase appears to assume a rate-limiting function in starch synthesis in the light when ADP glucose pyrophosphorylase is activated.