Cellular energetics during exercise.

Abstract

How is the muscle fiber designed to accomplish the diversity of tasks performed by striated muscle? Basically, a common contractile mechanism and a similar organization of metabolism in striated muscles are used to generate a wide spectrum of speeds and durations of contraction. The speed of contraction ranges from manyfold within an animal to over a hundred-fold between animals, owing to variation in the intrinsic velocities of the myosin isoforms. Recruiting fibers that contain the myosin isoform that contracts at the appropriate velocity varies the speed of locomotion at minimal cost. The magnitude and duration of the energy supply required to meet this contractile demand depends on the size of the cellular energy buffer and the capacities of the metabolic pathways. The faster the contractile speed, the larger the PCr pool and the greater the glycolytic capacity to meet a high rate of ATP use. Slower-contracting fibers have a smaller buffer for the short term, but an increased oxidative capacity for continuous energy supply to maintain energy balance over the long term. In general, fibers trade contractile speed for duration of performance, but a number of exceptions exist where rapid contractions are maintained for extended periods. In the face of this heterogeneity of properties, common features are found that assure an energy balance. The PCr/ATP buffer system offers a simple mechanism of feedback control of energy supply despite the wide range of high-energy phosphate concentrations and oxidative capacities found in skeletal muscle. An oxygen balance system also appears to be present in the terminal structures of the respiratory system, the capillaries, and mitochondria. Despite the diversity of these structures, a rather constant ratio of oxygen delivery capacity to mitochondrial oxidative capacity is found in vertebrate striated muscles. Finally, a critical feature of muscle energy balance that remains unresolved is (are) the mechanism(s) controlling mitochondrial respiration in heart. Feedback control appears to account for linking ATP supply to demand in skeletal muscle, but the mechanisms governing respiratory control in heart are still under vigorous investigation. Thus, the links between contractile demand and oxidative phosphorylation are still unresolved in this tissue, which may indicate that a key element is missing in our understanding of the cellular energetics of exercise.