Metabolic regulation by pH gradients. Inhibition of photosynthesis by indirect proton transfer across the chloroplast envelope.

Anions of several weak acids inhibited photosynthesis in isolated spinach chloroplasts. Inhibition was drastic at low pH and weak or absent at high pH. Glyoxylate was particularly effective and inhibition decreased in the order: glyoxylate, nitrite, glycerate, formate, hydroxypyruvate, glycolate, propionate, acetate, pyruvate. These anions operated as indirect proton shuttles across the chloroplast envelope. They compensated active proton fluxes into the medium, minimized gradients in proton activity across the chloroplast envelope, and so prevented light-dependent stroma alkalization. This caused inhibition of sugar bisphosphatases which are known to be pH-regulated. At concentrations that caused potosynthesis inhibition, the proton shuttles were not effective in decreasing the proton gradient across the thylakoids. Some anions also inhibited fructose-bisphosphatase directly, when present at concentratins higher than needed for photosynthesis inhibition.