Cell physiological aspects of the plasma membrane electrogenic H+ pump

This article deals with cell physiological aspects of the plasma membrane electrogenic proton (H+) pump and emphasizes the contribution of the giant algal cells of the Characeae in elucidating the mechanism of the pump. First, a history of the development of intracellular perfusion techniques in characean internodal cells is described, including preparation of tonoplast-free cells. Then, an outline of the hypothesis of the electrogenic H+ pump proposed by Kitasato is introduced, who prophesied the existence of an electric potential generated by an active H+ efflux. Subsequently, a history of finding ATP as the direct energy source of the electrogenic ion pump is presented. Quantitative agreement between the pump current and the ATP-dependent H+ efflux supports the notion that the ion carried by the electrogenic ion pump is H+. The role of the H+ pump in regulation of the cytosolic pH is discussed. Mechanisms of light-induced potential change through photosynthesis-controlled activation of the H+ pump are discussed in terms of changes in the levels of adenine nucleotides and in modulation of the Km value for the ATP of H+-ATPase. Recent progress in the molecular mechanism of the blue-light-induced activation of the H+-ATPase in guard cells is presented. However, there are cases where H+-ATPase activity is inhibited by blue light, indicating the flexibility of the control mechanisms of H+-ATPase activity. Finally, modulation of H+-pumping or H+-ATPase activities in response to environmental factors, such as anoxia, membrane excitation, osmotic and salt stresses, nutrient deficiencies and aluminum toxicity are described. Discussions are presented on the regulation of the electrogenic H+ pump.