Implications of positive feedback in the control of movement

In this paper we review some theoretical aspects of positive feedback in the control of movement. The focus is mainly on new theories regarding the reflexive role of sensory signals from mammalian tendon organ afferents. In static postures these afferents generally mediate negative force feedback. But in locomotion there is evidence of a switch to positive force feedback action. Positive feedback is often associated with instability and oscillation, neither of which occur in normal locomotion. We address this paradox with the use of analytic models of the neuromuscular control system. It is shown that positive force feedback contributes to load compensation and is surprisingly stable because the length-tension properties of mammalian muscle provide automatic gain control. This mechanism can stabilize control even when positive feedback is very strong. The models also show how positive force feedback is stabilized by concomitant negative displacement feedback and, unexpectedly, by delays in the positive feedback pathway. Other examples of positive feedback in animal motor control systems are discussed, including the beta-fusimotor system, which mediates positive feedback of displacement. In general it is seen that positive feedback reduces the sensitivity of the controlled extremities to perturbations of posture and load. We conclude that positive force feedback can provide stable and effective load compensation that complements the action of negative displacement and velocity feedback.