Intralimb compensation strategy depends on the nature of joint perturbation in human hopping

Due to the well-described spring-mass dynamics of bouncing gaits, human hopping is a tractable model for elucidating basic neuromuscular compensation principles. We tested whether subjects would employ a multi-joint or single-joint response to stabilize leg stiffness while wearing a spring-loaded ankle-foot orthosis (AFO) that applied localized resistive and assistive torques to the ankle. We analyzed kinematics and kinetics data from nine subjects hopping in place on one leg, at three frequencies (2.2, 2.4, and 2.8Hz) and three orthosis conditions (freely articulating AFO, AFO with plantarflexion resistance, and AFO with plantarflexion assistance). Leg stiffness was invariant across AFO conditions, however, compensation strategy depended upon the nature of the applied load. Biological ankle stiffness increased in response to a resistive load at twice the rate that it decreased with an assitive load. Ankle adjustments alone fully compensated for an assistive load with no net change in combined (biological plus applied) total ankle stiffness (p > or =0.133). In contrast, a resistive load resulted in a 7.4-9.0% increase in total ankle stiffness across frequencies and a concomitant 10-15% increase in knee joint stiffness at each frequency (p< or =0.037). The increased knee joint stiffness in response to resistive ankle load allowed subjects to maintain a more flexed knee at mid-stance, which attenuated the effect of the increased total ankle joint stiffness to preserve leg stiffness and whole limb biomechanical performance. Our findings suggest humans maintain invariant leg stiffness in bouncing gaits through different intralimb compensation strategies that are specific to the nature of the joint loading.