Potential roles of force cues in human stance control.

Human stance is inherently unstable. A small deviation from upright body orientation is enough to yield a gravitational component in the ankle joint torque, which tends to accelerate the body further away from upright ('gravitational torque'; magnitude is related to body-space lean angle). Therefore, to maintain a given body lean position, a corresponding compensatory torque must be generated. It is well known that subjects use kinematic sensory information on body-space lean from the vestibular system for this purpose. Less is known about kinetic cues from force/torque receptors. Previous work indicated that they are involved in compensating external contact forces such as a pull or push having impact on the body. In this study, we hypothesized that they play, in addition, a role when the vestibular estimate of the gravitational torque becomes erroneous. Reasons may be sudden changes in body mass, for instance by a load, or an impairment of the vestibular system. To test this hypothesis, we mimicked load effects on the gravitational torque in normal subjects and in patients with chronic bilateral vestibular loss (VL) with eyes closed. We added/subtracted extra torque to the gravitational torque by applying an external contact force (via cable winches and a body harness). The extra torque was referenced to body-space lean, using different proportionality factors. We investigated how it affected body-space lean responses that we evoked using sinusoidal tilts of the support surface (motion platform) with different amplitudes and frequencies (normals +/-1 degrees, +/-2 degrees, and +/-4 degrees at 0.05, 0.1, 0.2, and 0.4 Hz; patients +/-1 degrees and +/-2 degrees at 0.05 and 0.1 Hz). We found that added/subtracted extra torque scales the lean response in a systematic way, leading to increase/decrease in lean excursion. Expressing the responses in terms of gain and phase curves, we compared the experimental findings to predictions obtained from a recently published sensory feedback model. For the trials in which the extra torque tended to endanger stance control, predictions in normals were better when the model included force cues than without these cues. This supports our notion that force cues provide an automatic 'gravitational load compensation' upon changes in body mass in normals. The findings in the patients support our notion that the presumed force cue mechanism provides furthermore vestibular loss compensation. Patients showed a body-space stabilization that cannot be explained by ankle angle proprioception, but must involve graviception, most likely by force cues. Our findings suggest that force cues contribute considerably to the redundancy and robustness of the human stance control system.