Occurrence of widespread motor-unit firing correlations in muscle contractions: Their role in the generation of tremor and time-varying voluntary force

The firing behavior of motor units (MUs) of the first dorsal interrosseus muscle of the hand was examined during both constant-force and varying-force (sinusoidal or broadband random variations) isometric contractions in healthy adults. The emphasis was on the analysis of MU synchrony with an efficient and sensitive method. In static contractions, widespread and strong MU firing correlations, with the MUs in phase, were present at the frequency of muscle tremor, when the tremor was regular (narrowband) and large. MU correlations could also exist in contractions where the tremor of a subject was irregular (broadband) overall, but they were generally weak. These correlations were at the frequency of the subject's regular tremor, and the corresponding distinct tremor component was sometimes discernible within the broad tremor-band. In contrast, the MUs did not show any such correlations in the case of purely irregular and small tremor. On the basis of these observations, it is concluded that the rhythms in the force contributions of the last-recruited, large MUs, which fire near their threshold rate, compose the broadband frequency content of physiological muscle tremor in every contraction. Within this band, there is an additional distinct tremor component when MU correlations are present. For widespread and strong MU correlations, this component dominates and constitutes the observed regular tremor. In dynamic contractions, the firing of all MUs was modulated in the frequency band of both the sinusoidal and the complex variations of the force. The MU modulations showed a time-lead over the force variations and were strongly correlated both to these variations and among themselves. Thus widespread and strong correlations of MU firing modulations seem to provide a mechanism for generation of time-varying voluntary force, under general dynamic conditions. Finally, when regular tremor was present in dynamic contractions, widespread and fairly strong MU correlations also existed at the tremor frequency. It is concluded that at least two mechanisms can cause widespread MU synchrony, and they can act in parallel. They involve two types of correlated inputs to the cu-motoneurons (presumably from the muscle spindles and the cortex), whose effects combine at the level of the membrane potential of the cells.