Advances in understanding the energetics of muscle contraction.

Muscle energetics encompasses the relationships between mechanical performance and the biochemical and thermal changes that occur during muscular activity. The biochemical reactions that underpin contraction are described and the way in which these are manifest in experimental recordings, as initial and recovery heat, is illustrated. Energy use during contraction can be partitioned into that related to cross-bridge force generation and that associated with activation by Ca2+. Activation processes account for 25-45% of ATP turnover in an isometric contraction, varying amongst muscles. Muscle energy use during contraction depends on the nature of the contraction. When shortening muscles produce less force than when contracting isometrically but use energy at a greater rate. These characteristics reflect more rapid cross-bridge cycling when shortening. When lengthening, muscles produce more force than in an isometric contraction but use energy at a lower rate. In that case, cross-bridges cycle but via a pathway in which ATP splitting is not completed. Shortening muscles convert part of the free energy available from ATP hydrolysis into work with the remainder appearing as heat. In the most efficient muscle studied, that of a tortoise, cross-bridges convert a maximum of 47% of the available energy into work. In most other muscles, only 20-30% of the free energy from ATP hydrolysis is converted into work.