Metabolic regulation of pH in plant cells: role of cytoplasmic pH in defense reaction and secondary metabolism.

A new biochemical pH-stat hypothesis that revised the classic hypothesis is presented to understand the metabolic regulation of intracellular pH in plant cells. Alternative pathway glycolysis, alternative pathway respiration and malate-derived lactic and alcoholic fermentation (alternative pathway fermentation), all unique to plants, are integrated into a regulatory mechanism of pH in the cytoplasm. Its uniqueness to plant kingdom is discussed from the evolutionary viewpoint: it is suggested that when the ancestors of extant terrestrial plants expanded their habitat from oceans to freshwater, they abandoned a "sodium system" and adopted a "proton system" for nutrient uptake. Validity of the new hypothesis is examined with available data on a secondary active transport, anoxia and other experimental evidence. The hypothesis predicts that biotic and abiotic stress-induced cytoplasmic acidification triggers synthesis of phytoalexins and other secondary metabolites. Possible roles of cyanide-resistant alternative pathway respiration in the secondary metabolite production, metabolic switching between primary and secondary metabolisms, and defense reactions are proposed.