Factors regulating the slow electrogenic phase in green algae and higher plants.

The electrochromic rise in the millisecond range, corrected for the subsequent decay, has been studied in Chlorella cells and spinach chloroplasts. The half-time of the electrochromic rise in the millisecond range is independent of the redox states of the components in the electrogenic loop. It is also independent of pH in the absence of a pH gradient, but it is dependent upon the transmembrane electric field and the pH gradient. The limiting step of the overall process is thus the electrogenic reaction itself. The amplitude of the electrochromic rise in the millisecond range is controlled by two classes of factors: the redox state of its reactants and the free energy available in the overall process including the electrogenic reaction. This free energy depends upon delta pH and transmembrane electric field. A fast reduction of cytochrome f+ (and thus probably of the Rieske protein) by Photosystem II is still possible when the electrogenic reaction is impeded for energetic reasons. The kinetic behavior of the electrogenic reaction could be interpreted by the following reaction: UH2int + FeS+ + X2 + Hext in equilibrium U + 2H+int + FeS + X2H where FeS is the Rieske protein; X2 would be a new protein (observed in Bouges-Bocquet, B. (1980) FEBS Lett. 117, 54-58); UH2 which is likely a special quinone, would release its two protons inside the thylakoids and X2 would use protons from the external medium.