Limits and constraints in the scaling of oxidative and glycolytic enzymes in homeotherms

It is now empirically well established that basal and maximum rates of O2 uptake in homeotherms scale approximately to the 0.75 power; log–log plots of mass-specific metabolic rates versus body mass yield slopes of −0.20 to 0.25. Recent studies of 10 mammalian species and 1 hummingbird species indicate that marker enzymes of mitochondrial metabolism (citrate synthase, for example) scale inversely with body mass. Hummingbirds and shrews are near the upper limit in the degree to which the oxidative capacity of heart and skeletal muscles can be elevated; further increases in mitochondrial volume densities would sacrifice myofilament or sarcoplasmic reticulum volume densities. Whales weighing about 105 kg may be near the limit at the opposite extreme because their mass-specific resting metabolic rates are predicted to be approaching those of hypometabolic ectotherms. In contrast to oxidative enzyme scaling patterns, enzymes normally operative in muscle anaerobic glycolysis, such as lactate dehydrogenase, scale directly with body mass. Hummingbirds and shrews are considered to have reduced muscle lactate dehydrogenase levels near a lower limit commensurate with buffering of cytosolic redox, a distinctly aerobic lactate dehydrogenase function. How much anaerobic glycolytic potential can be packed into muscle cells in the largest mammals is unknown; this upper limit appears to be set by a compromise between myofilament volume densities and the combined volume densities of glycogen granules, intracellular buffering components, and glycolytic enzymes.