A cross-bridge model that is able to explain mechanical and energetic properties of shortening muscle.

The responses of muscle to steady and stepwise shortening are simulated with a model in which actin-myosin cross-bridges cycle through two pathways distinct for the attachment-detachment kinetics and for the proportion of energy converted into work. Small step releases and steady shortening at low velocity (high load) favor the cycle implying approximately 5 nm sliding per cross-bridge interaction and approximately 100/s detachment-reattachment process; large step releases and steady shortening at high velocity (low load) favor the cycle implying approximately 10 nm sliding per cross-bridge interaction and approximately 20/s detachment-reattachment process. The model satisfactorily predicts specific mechanical properties of frog skeletal muscle, such as the rate of regeneration of the working stroke as measured by double-step release experiments and the transition to steady state during multiple step releases (staircase shortening). The rate of energy liberation under different mechanical conditions is correctly reproduced by the model. During steady shortening, the relation of energy liberation rate versus shortening speed attains a maximum (approximately 6 times the isometric rate) for shortening velocities lower than half the maximum velocity of shortening and declines for higher velocities. In addition, the model provides a clue for explaining how, in different muscle types, the higher the isometric maintenance heat, the higher the power output during steady shortening.