What is the contribution of voluntary and reflex processes to sensorimotor control of balance?

Abstract

The contribution to balance of spinal and transcortical processes including the long-latency reflex is well known. The control of balance has been modelled previously as a continuous, state feedback controller representing, long-latency reflexes. However, the contribution of slower, variable delay processes has not been quantified. Compared with fixed delay processes (spinal, transcortical), we hypothesize that variable delay processes provide the largest contribution to balance and are sensitive to historical context as well as current states. Twenty-two healthy participants used a myoelectric control signal from their leg muscles to maintain balance of their own body while strapped to an actuated, inverted pendulum. We study the myoelectric control signal (u) in relation to the independent disturbance (d) comprising paired, discrete perturbations of varying inter-stimulus-interval (ISI). We fit the closed loop response, u from d, using one linear and two non-linear non-parametric (many parameter) models. Model M1 (ARX) is a generalized, high-order linear-time-invariant (LTI) process with fixed delay. Model M1 is equivalent to any parametric, closed-loop, continuous, linear-time-invariant (LTI), state feedback model. Model M2, a single non-linear process (fixed delay, time-varying amplitude), adds an optimized response amplitude to each stimulus. Model M3, two non-linear processes (one fixed delay, one variable delay, each of time-varying amplitude), add a second process of optimized delay and optimized response amplitude to each stimulus. At short ISI, the myoelectric control signals deviated systematically both from the fixed delay LTI process (M1), and also from the fixed delay, time-varying amplitude process (M2) and not from the two-process model (M3). Analysis of M3 (all fixed delay and variable delay response amplitudes) showed the variable (compared with fixed) delay process 1) made the largest contribution to the response, 2) exhibited refractoriness (increased delay related to short ISI) and 3) was sensitive to stimulus history (stimulus direction 2 relative to stimulus 1). For this whole-body balance task and for these impulsive stimuli, non-linear processes at variable delay are central to control of balance. Compared with fixed delay processes (spinal, transcortical), variable delay processes provided the largest contribution to balance and were sensitive to historical context as well as current states.

The ‘Postural Rhythm’ of the Ground Reaction Force during Upright Stance and Its Conversion to Body Sway—The Effect of Vision, Support Surface and Adaptation to Repeated Trials

“…Maintenance of upright stance occurs despite imprecise and poorly anticipated sensory feedback and uncertain knowledge of the effects of the motor action onto the support surface, so that neither reflex responses nor predetermined muscle synergies would explain coping with critical conditions [201][202][203][204]. This requires the brain to comply with the fortuitous modifications of the sensorimotor feedback [205], where modulation of the receiving sensory cortex by the active motor cortex might play a role [206] (Enomoto et al, 2001). Partial functional disruption of selected cortical areas by transcranial stimulation disturbs body balance, clearly showing that the brain is involved in stance control [207,208].…”