The role of action reflexes in the damping of mechanical oscillations.

Reflex responses measured during voluntary contraction of the muscle being stretched are known to differ markedly from reflex responses elicited from passive muscle. The term 'action tonic stretch reflex' or 'action TSR' has been used previously to describe a reflex response to continuous stretch, separated from voluntary activity by means of a cross-correlational and spectrographic analysis. In this paper it is proposed that the action TSR play a functional role during voluntary movement by damping the transient oscillations associated with the natural resonant frequencies of the limbs. It is suggested that oscillations excite an action TSR force response with phase lead ahead of muscle stretch. This force response can be resolved into two force components, one of which has a 90 degree phase lead ahead of muscle stretch and behaves like a viscous friction reaction force causing damping of oscillations. Three experiments which support this proposition are described. (1) Analog computer model stimulation studies of a muscle supporting a mass-spring load reveal that damping only occurs when the force response has a phase lead ahead of muscle stretch. When the force response has a phage lag behind muscle stretch, the system is unstable and the amplitude of oscillation increases with time. (2) It is demonstrated that when a mass-spring load is supported as rigidly as possible by the human arm, reaction forces from the arm damp and mass-spring oscillations more rapidly than when the mass-spring is rigidly supported. Electromyogram (EMG) recordings reveal that mass-spring oscillations excite action TSR responses with phase lead ahead of muscle stretch. (3) Recordings of elbow angle and biceps EMG during rapid forearm flexion or extension movements followed by sudden stops reveal critically damped oscillations in the elbow angle signal (i.e. no more than one or two small overshoots), which are accompanied by EMG action TSR responses with phase lead ahead of muscle stretch.