Active stiffness of the ankle in response to inertial and elastic loads.

Effective stiffness of the musculoskeletal system was examined as a function of the characteristics of an external load. Thirteen healthy subjects provided active contraction of the ankle plantarflexion musculature in a neutral ankle posture to support an external load. Musculoskeletal stiffness was computed from kinetic data recorded in response to dorsiflexion/plantarflexion perturbations. Ankle dynamics were recorded while supporting external loads of 19 and 38 kg with and without antagonistic co-contraction. External loads were applied using pure gravitational mass. In separate trials external loads were applied from stretch of steel springs in parallel with the plantarflexion musculature that also provided added parallel stiffness to the system. Adding external stiffness of 4.9 and 8.1 kN/m surprisingly failed to significantly change the stiffness of the ankle-plus-spring system. This suggests contributions from intrinsic muscle stiffness and reflex stiffness declined in response to added external stiffness. This could not be explained by load magnitudes, ankle postures, or co-activation as these were similar between the inertial and elastic loading conditions. However, non-linear parametric analyses suggest mean intrinsic stiffness of 35.5 kN/m and reflex gain of 11.6 kN/m with a constant reflex delay of 70 ms accurately described the empirical results. The phase response between the mechanical dynamics of the musculoskeletal system and delayed neuromotor feedback combine to provide robust control of system behavior.