Bioenergetics of lactic acid bacteria: cytoplasmic pH and osmotolerance

Abstract

Lactic acid bacteria maintain a cytoplasm that is more alkaline than the medium, but whose pH decreases as the medium is acidified during growth and fermentation. Streptococci generally acidify the cytoplasm from approximately pH 7.6 to 5.7 (external pH 4.5) before growth and then fermentation cease. The internal enzyme machinery of these anaerobic fermenters thus tolerates a fairly wide range in internal proton concentration. Lactobacilli tolerate a significantly more acidic cytoplasmic pH of 4.4 (external pH 3.5). However, when the cytoplasmic pH decreases below a threshold pH, which depends on the organism cellular functions are inhibited. Fermentation end-products, such as organic acids or alcohols, exert their deleterious effects by bringing about acidification of the cytoplasm below the permissible pH. Organic acids, which act as protonophores, or solvents, which perturb membrane phospholipids, at high concentrations increase the inward leak of H+ so that H+ efflux is not rapid enough to alkalinize the cytoplasm. The membrane pH gradient is thus dissipated. A specific strain of Lactobacillus acidophilus has been found to be unusually osmotolerant. The osmoresistance is due to the cells' capacity to accumulate glycine betaine by a transport carrier that is activated, but not induced, by high medium osmotic pressure.