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The Journal of Neurophysiology Vol. 88 No. 3 September 2002, pp. 1097-1118
Copyright ©2002 by the American Physiological Society
Neurological Sciences Institute, Oregon Health & Science University, Portland, Oregon 97006
Peterka, R. J.
Sensorimotor Integration in Human Postural Control. J. Neurophysiol. 88: 1097-1118, 2002. It is
generally accepted that human bipedal upright stance is achieved by
feedback mechanisms that generate an appropriate corrective torque
based on body-sway motion detected primarily by visual, vestibular, and
proprioceptive sensory systems. Because orientation information from
the various senses is not always available (eyes closed) or accurate
(compliant support surface), the postural control system must somehow
adjust to maintain stance in a wide variety of environmental
conditions. This is the sensorimotor integration problem that we
investigated by evoking anterior-posterior (AP) body sway using
pseudorandom rotation of the visual surround and/or support surface
(amplitudes 0.5-8°) in both normal subjects and subjects with severe
bilateral vestibular loss (VL). AP rotation of body center-of-mass
(COM) was measured in response to six conditions offering different
combinations of available sensory information. Stimulus-response data
were analyzed using spectral analysis to compute transfer functions and
coherence functions over a frequency range from 0.017 to 2.23 Hz.
Stimulus-response data were quite linear for any given condition and
amplitude. However, overall behavior in normal subjects was nonlinear
because gain decreased and phase functions sometimes changed with
increasing stimulus amplitude. "Sensory channel reweighting" could
account for this nonlinear behavior with subjects showing increasing
reliance on vestibular cues as stimulus amplitudes increased. VL
subjects could not perform this reweighting, and their
stimulus-response behavior remained quite linear. Transfer function
curve fits based on a simple feedback control model provided estimates
of postural stiffness, damping, and feedback time delay. There were
only small changes in these parameters with increasing visual stimulus
amplitude. However, stiffness increased as much as 60% with increasing
support surface amplitude. To maintain postural stability and avoid
resonant behavior, an increase in stiffness should be accompanied by a corresponding increase in damping. Increased damping was achieved primarily by decreasing the apparent time delay of feedback control rather than by changing the damping coefficient (i.e., corrective torque related to body-sway velocity). In normal subjects, stiffness and damping were highly correlated with body mass and moment of inertia, with stiffness always about 1/3 larger than necessary to
resist the destabilizing torque due to gravity. The stiffness parameter
in some VL subjects was larger compared with normal subjects,
suggesting that they may use increased stiffness to help compensate for
their loss. Overall results show that the simple act of standing
quietly depends on a remarkably complex sensorimotor control system.
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