Bacterial energetics at high pH: what happens to the H+ cycle when the extracellular H+ concentration decreases?

A decrease in the extracellular H+ concentration creates difficulties for membrane-linked energetics in bacteria employing H+ as the coupling ion. At high extracellular pH (pHo), H+ ions pumped from the cell by, say, the respiratory chain, are immediately neutralized by the alkaline extracellular medium. Under such conditions, the only driving force that might compel outer H+ ions to return to cytosol and perform their function is the electric potential difference across the cytoplasmic membrane (delta psi). However, when delta pH in the opposite direction is equal to, e.g., 2 pH units (intracellular pH = 7.5 at pHo = 9.5), delta psi would be so high that the risk of membrane electric breakdown would increase. This is why some bacteria deal with high pH by, for example, replacing H+ by Na+ as the coupling ion rather than by increasing delta psi. It has been shown in several species of bacteria that the alkalinization of the growth medium induces primary Na+ pumps (e.g. Na(+)-motive respiratory chain enzymes and Na+ ATPase). Electrogenic Na+ efflux via these pumps produces an electrochemical Na+ potential difference (delta mu Na+) composed of delta psi and delta pNa+. delta mu Na+ can be used to perform various types of membrane-linked work. The delta psi constituent of delta mu Na+ may maintain electrophoretic influx of H+ such that the alkalinization of cytoplasm is prevented. The latter function may be supported by a mechanism based on the uphill influx of Cl- instead of Na+. This seems to be the case for alkaliphilic and halophilic Natronobacter pharaonis. There is an indication that not only Na+ but also Ca2+ may substitute for H+ in Gleobacter violaceus growing at high pH.