Energetics of shortening muscles in twitches and tetanic contractions. II. Force-determined shortening heat.

The extra heat liberation accompanying muscular shortening, the force-determined shortening heat, is defined as the difference between the heat produced when shortening occurs and that produced in an isometric contraction developing the same amount of force and performing the same amount of internal work. Based on this definition, the initial energy production in twitches and tetanic contractions (E) is given by E = A + f (P, t) + alpha(F)x + W, where A is the activation heat, f(P, t), the tension-related heat (a heat production associated with the development and maintenance of tension), alpha(F)x, the force-determined shortening heat, and W, the external work. It is demonstrated that this equation accurately accounts for the time-course of heat evolution and the total initial energy production in both twitches and tetani at 0 degrees C. The force-determined shortening heat is liberated, during shortening, in direct proportion to (a) the distance shortened, and (b) the force against which shortening occurs. The normalized value of the force-determined shortening heat coefficient, alpha(F)/P(o), is the same in both the twitch and the tetanus. Finally, this formulation of the muscle's energy production also accounts for the total energy production in afterload isotonic twitches at 20 degrees C, where a Fenn effect is not demonstrable.