Entrainment to extinction of physiological tremor by spindle afferent input.

In this study the systematic modulation of wrist flexor muscle activity by imposed joint movement was examined. Ten subjects maintained a constant contraction level (25% of the maximum; trial duration: 20 s) in flexor carpi radialis while their wrists were perturbed with 50 different quasi-sinusoidal signals (frequency range: 0.5-9.5 Hz; amplitude: 0.3-4.2 degrees ). The frequency spectra of wrist position and the rectified and filtered electromyogram (EMG) were determined. The muscle activity was only weakly entrained to imposed movements of small amplitude and low frequency, as shown by a small peak in the EMG spectrum at the frequency of movement, while the most prominent peak in the spectrum was between 9 and 15 Hz, corresponding to the frequency range of physiological tremor. The entrainment of muscle activity increased markedly as the amplitude and frequency of the imposed movement increased, to the point of saturation of modulation and harmonic peaks in the spectrum. In parallel with this increase in entrainment, the 9-15 Hz tremor peak was progressively extinguished. The results are consistent with a coupled oscillator model in which the central oscillatory source(s) of tremor became fully entrained to the imposed movement at the highest amplitudes and frequencies. Such coupling depends on communication between the external forcing oscillator and the central oscillator(s), the I (a) afferent signal from the imposed movement being the most likely candidate to provide the entraining signal for the central oscillator(s).