During oxygen limitation in animals, glucose can be fermented via several metabolic pathways varying in energetic efficiency and leading to various end products (such as lactate, alanopine, octopine, succinate, or propionate). Because of opposite p H dependencies of proton production by fermentation and by hydrolysis of adenosine triphosphate formed in the fermentation, the total number of moles of protons generated is always two per mole of the fermentable substrate. However, two and three times more adenosine triphosphate can be turned over per mole of protons produced in succinate and propionate fermentations, respectively, than in lactate fermentation.