Thixotropic behaviour of human finger flexor muscles with accompanying changes in spindle and reflex responses to stretch.

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Abstract
1. Prompted by previous reports on muscle thixotropy, we have investigated changes in inherent and reflex stiffness of the finger flexor muscles of human subjects at rest, following transient conditioning manoeuvres involving contractions and/or length changes of the finger flexors. The stiffness measurements were combined with electromyographic recordings from forearm and hand muscles and with microneurographic recordings of afferent stretch responses in finger flexor nerve fascicles. 2. Finger flexor stiffness was evaluated by measuring (a) the flexion angle of the metacarpo-phalangeal joints at which the system during rest balanced the force of gravity and (b) the speed and amplitude of angular finger extensions induced by recurrent extension torque pulses of constant strength delivered by a torque motor. In the latter case, extension drifts in the resting position of the fingers were prevented by a weak flexion bias torque holding the fingers in a pre-determined, semiflexed position against a stop-bar. 3. Stiffness changes following passive large amplitude finger flexions and extensions were studied in subjects with nerve blocks or nerve lesions preventing neurally mediated contractions in the forearm and hand muscles. Inherent stiffness was enhanced following transient finger flexions and reduced following transient finger extensions. The after-effects gradually declined during observation periods of several minutes. 4. Similar results were obtained in subjects with intact innervation who succeeded during the pre- and post-conditioning periods in keeping the arm and hand muscles relaxed (i.e. showed no electromyographic activity). In these subjects it was also found that the after-effects were similar for active and passive finger movements and that isometric voluntary finger flexor contractions loosened the system in a way similar to finger extensions. 5. In some subjects electromyographic reflex discharges appeared in the finger flexors in response to the extension test pulses. When elicited by small ramp stretch stimuli of constant amplitude, the stretch reflex responses were found to vary in strength in parallel with the changes in inherent stiffness following the various conditioning manoeuvres. 6. The strength of the multi-unit afferent stretch discharges in the muscle nerve, used as index of muscle spindle stretch sensitivity, varied in parallel with the changes in inherent stiffness. Post-manoeuvre changes in muscle spindle stretch sensitivity were seen also when the spindles were de-efferented by a nerve block proximal to the recording site. 7. The results can be explained in terms of thixotropic behaviour of extra- and intrafusal muscle fibres. The in-parallel changes in reflex and inherent stiffness can be accounted for by assuming that the conditioning manoeuvres have similar mechanical after-effects on intra- and extrafusal muscle fibres, and that the primary spindle endings are more susceptible to the intra- than to the counteracting extrafusal stiffness changes. 8. Loosening of thixotropic bonds in extra- and intrafusal muscle fibres may contribute to the beneficial effects of limbering-up manoeuvres commonly employed by athletes, dancers and physiotherapists.