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J Neurophysiol 95: 53-66, 2006. First published September 14, 2005; doi:10.1152/jn.00051.2005
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Parallel Neuronal Mechanisms Underlying Physiological Force Tremor in Steady Muscle Contractions of Humans

Constantinos N. Christakos, Nikos A. Papadimitriou and Sophia Erimaki

Laboratory of Systems Physiology, Division of Basic Sciences, Medical School, University of Crete; and Computational Neuroscience Group, Institute of Applied and Computational Mathematics, Foundation for Research and Technology Hellas, Heraklion, Greece

Submitted 18 January 2005; accepted in final form 6 September 2005

We present results from a study of the 6-to 12-Hz force tremor in relation to motor unit (MU) firing synchrony. Our experimental observations from 32 subjects, 321 contractions, and 427 recorded MUs reveal that tremor is accompanied by corresponding, in-phase MU rhythms that are additional to the ones at the MU intrinsic firing rates. This rhythmical synchrony is widespread and has a uniform strength that ranges from near zero to very large (MU/MU coherence > 0.50) in different contractions. Both the synchrony and the tremor are suppressed during ischemia, and this strongly suggests an involvement of spindle feedback in their generation. Furthermore, in the presence of substantial synchrony, the tremor enhancement, relative to the minimal tremor of ischemia, reflects the strength of the synchrony. Theoretical considerations based on these observations indicate that the muscle force signal is expected to show 1) frequency components in the band of the firing rates of the last-recruited, large MUs, and 2) because of the synchronized MU rhythms, an additional, distinct component with a size reflecting the strength of synchrony. Furthermore, synchronized MU rhythms, with frequencies in the 6- to 12-Hz range, are expected to arise from self-oscillations in the monosynaptic stretch reflex loop, due primarily to the associated muscle delay (several tens of milliseconds). Our results therefore reveal the parallel action of two tremor mechanisms, one of which involves MU synchrony probably caused by loop action. Clearly, the results on the synchrony and its impact also apply to other possible generators of tremor synchrony, including supraspinal ones.


Address for reprint requests and other correspondence: C. N. Christakos, Div. of Basic Sciences, Medical School, Univ. of Crete, 71003 Heraklion, Crete, Greece (E-mail: ccnncc{at}med.uoc.gr)




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