The role of intersegmental dynamics during rapid limb oscillations.

The interactive dynamic effects of muscular, inertial and gravitational moments on rapid, multi-segmented limb oscillations were studied. Using three-segment, rigid-body equations of motion, hip, knee and ankle intersegmental dynamics were calculated for the steady-state cycles of the paw-shake response in adult spinal cats. Hindlimb trajectories were filmed to obtain segmental kinematics, and myopotentials of flexors and extensors at each of the three joints were recorded synchronously with the ciné film. The segmental oscillations that emerged during the paw-shake response were a consequence of an interplay between active and passive musculotendinous forces, inertial forces, and gravity. During steady-state oscillations, the amplitudes of joint excursions, peak angular velocities, and peak angular accelerations increased monotonically and significantly in magnitude from the proximal joint (hip) to the most distal joint (ankle). In contrast to these kinematic relationships, the maximal values of net moments at the hip and knee were equal in magnitude, but of significantly lower magnitude than the large net moment at the ankle joint. At both the ankle and the knee, the flexor and extensor muscle moments were equal, but at the hip the magnitude of the peak flexor muscle moment was significantly greater than the extensor muscle moment. Muscle moments at the hip not only acted to counterbalance accelerations of the more distal segments, but also acted to maintain the postural orientation of the hindlimb. Large muscle moments at the knee functioned to counterbalance the large inertial moments generated by the large angular accelerations of the paw. At the ankle, the muscle moments dominated the generation of the paw accelerations. At the ankle and the knee, muscle moments controlled limb dynamics by slowing and reversing joint motions, and the active muscle forces contributing to ankle and knee moments were derived from lengthening of active musculotendinous units. In contrast to the more distal joints, the active muscles crossing the hip predominantly shortened as a result of the interplay among inertial forces and gravitational moments. The muscle function and kinetic data explain key features of the complex interactions that occur between central control mechanisms and multi-segmented, oscillating limb segments during the paw-shake response.