Spatial and temporal modulation of joint stiffness during multijoint movement.

Joint stiffness measurements during small transient perturbations have suggested that stiffness during movement is different from that observed during posture. These observations are problematic for theories like the classical equilibrium point hypothesis, which suggest that desired trajectories during movement are enforced by joint stiffness. We measured arm impedances during large, slow perturbations to obtain detailed information about the spatial and temporal modulation of stiffness and viscosity during movement. While our measurements of stiffness magnitudes during movement generally agreed with the results of measurements using fast perturbations, they revealed that joint stiffness undergoes stereotyped changes in magnitude and aspect ratio which depend on the direction of movement and show a strong dependence on joint angles. Movement simulations using measured parameters show that the measured modulation of impedance acts as an energy conserving force field to constrain movement. This mechanism allows for a computationally simplified account of the execution of multijoint movement. While our measurements do not rule out a role for afferent feedback in force generation, the observed stereotyped restoring forces can allow a dramatic relaxation of the accuracy requirements for forces generated by other control mechanisms, such as inverse dynamical models.