Contributions of anaerobic metabolism to pH regulation in animal tissues: theory.

Proton balance is analysed in relation to the anaerobic and aerobic metabolism of carbohydrates, carbonic acids, amino acids and fat by considering oxidation, carboxylation, decarboxylation and phosphorylation reactions, as well as the influence of ammonium, on the acid-base status of animal tissues. The functional role of the adenylates, phosphagens and inorganic phosphate in acid-base balance is investigated with respect to differences in the physicochemical properties of organic and inorganic phosphates. General principles are established for different anaerobic metabolic pathways in species from several phyla. It is concluded that proton release from the substrate, which is always involved in substrate-level phosphorylations, is essential for the mechanism of ATP formation. Anaerobic metabolism, which is characterized by incomplete oxidation of carbon chains and an accumulation of acidic groups, supports pH regulation in facultative anaerobes by minimizing the amount of accumulated protons. High levels of phosphagens mean high proton absorption during hydrolysis and an increase in the intracellular buffer value. Decarboxylation reactions in catabolic pathways are equivalent to proton consumption. The degradation of carbonic acids during anaerobiosis, therefore, contributes to pH regulation. Release of ammonia or ammonium ions in catabolism is also linked to the buffering of protons originating from the formation of carboxyl groups and net cleavage of ATP. Net disposal of amino groups or ammonium ions by transamination, reductive amination or ion exchange does not change this general picture. The proton, bicarbonate and CO2 turnover in metabolic pathways is discussed with respect to the interrelationships between pH and metabolic regulation.