Although many studies have investigated reflexes after stimulation of either cutaneous or proprioceptive afferents, much less is known about responses after more natural perturbations, such as stumbling over an obstacle. In particular, the phase dependency of these responses and their relation to the stumbling behavior has received little attention. Hence response strategies during stumbling reactions after perturbations at different times in the swing phase of gait were studied. While subjects walked on a treadmill, a rigid obstacle unexpectedly obstructed the forward sway of the foot. All subjects showed an "elevating strategy" after early swing perturbations and a "lowering strategy" after late swing perturbations. During the elevating strategy, the foot was directly lifted over the obstacle through extra knee flexion assisted by ipsilateral biceps femoris (iBF) responses and ankle dorsiflexion assisted by tibialis anterior (iTA) responses. Later, large rectus femoris (iRF) activations induced knee extension to place the foot on the treadmill. During the lowering strategy, the foot was quickly placed on the treadmill and was lifted over the obstacle in the subsequent swing. Foot placement was actively controlled by iRF and iBF responses related to knee extension and deceleration of the forward sway. Activations of iTA mostly preceded the main ipsilateral soleus (iSO) responses. For both strategies, four response peaks could be distinguished with latencies of approximately 40 ms (RP1), approximately 75 ms (RP2), approximately 110 ms (RP3), and approximately 160 ms (RP4). The amplitudes of these response peaks depended on the phase in the step cycle. The phase-dependent modulation of the responses could not be accounted for by differences in stimulation or in background activity and therefore is assumed to be premotoneuronal in origin. In mid swing, both the elevating and lowering strategy could occur. For this phase, the responses of the two strategies could be compared in the absence of phase-dependent response modulation. Both strategies had the same initial electromyographic responses till approximately 100 ms (RP1-RP2) after perturbation. The earliest response (RP1) is assumed to be a short-latency stretch reflex evoked by the considerable impact of the collision, whereas the second (RP2) has features reminiscent of cutaneous and proprioceptive responses. Both these responses did not determine the behavioral response strategy. The functionally important response strategies depended on later responses (RP3-RP4). These data suggest that during stumbling reactions, as a first line of defense, the CNS releases a relatively aspecific response, which is followed by an appropriate behavioral response to avoid the obstacle.

The purpose of the present study was to determine whether tactile cutaneous reflexes from the skin of the foot contain location-specific information during human walking. Muscular responses to non-nociceptive electrical stimulation of the sural, posterior tibial, and superficial peroneal nerves, each supplying a different skin area of the foot, were studied in both legs during walking on a treadmill. For all three nerves the major responses in all muscles were observed at a similar latency of ϳ80-85 msec. In the ipsilateral leg these reflex responses and their phase-dependent modulation were highly nerve-specific. During most of the stance phase, for example, the peroneal and tibial nerves generally evoked small responses in the biceps femoris muscle. In contrast, during late swing large facilitations generally occurred for the peroneal nerve, whereas suppressions were observed for the tibial nerve. In the contralateral leg

A new method to study the reactions to unexpected mechanical perturbations during human walking on a treadmill is presented.Perturbations consisted of an obstruction of the forward swinging foot during the early swing phase. These were caused by obstacles which were dropped on the treadmill in front of the subject The timing o f the perturbation was controlled by an electromagnet which released the obstacle at a preprogrammed delay after left or right heel strike. This kind of perturbation evoked stumbling reactions. The electromyographic (EM G ) responses during these stumbling reactions had mean latencies of 76 ms in both the ipsilateral biceps femoris and rectus femoris when perturbations were applied in early swing. During the perturbed swing, increased flexion in the knee occurred to lift the foot over the obstacle. Both the E M G and kinesiologic responses were reproducible when perturbations were presented in the same part of the swing phase of different step cycles.

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1. Cutaneous reflex responses were elicited during human running (8 km/h) on a treadmill by electrical stimulation of the sural nerve at the ankle. Stimulus trains (5 pulses of 1 ms at 200 Hz) at three nonnociceptive intensities, which were 1.5, 2.0, and 2.5 times perception threshold (PT), were delivered at 16 phases of the step cycle. For 11 subjects the surface electromyographic (EMG) activity of both the ipsilateral and contralateral long head of the biceps femoris (iBF and cBF, respectively), the semitendinosus (iST and cST), the rectus femoris (iRF and cRF), and the tibialis anterior (iTA and cTA) were recorded. 2. During human running nonnociceptive sural nerve stimulation appears to be sufficient to elicit large, widespread and statistically significant reflex responses, with a latency of approximately 80 ms and a duration of approximately 30 ms. These reflex responses seem to be an elementary property of human locomotion. This is indicated by the occurrence of the responses in all subjects, the consistency of most of the reflex patterns across the subjects and, apart from a small amount of habituation, the reproducibility of the responses during the course of the experiment. 3. The responses are modulated continuously throughout the step cycle such that their magnitude does not in general covary with the background locomotor activities. This is observed most clearly in iST, iTA, and cTA for which statistically significant reflex reversals are demonstrated, and in cRF and cTA for which the responses are gated during most of the step cycle. 4. The response magnitude generally increases as a function of increasing intensity, whereas the phase-dependent reflex modulation is intensity independent. 5. A functional dissociation within the ipsilateral hamstring muscles is demonstrated: the iBF and iST show an antagonistic reflex pattern (facilitatory and suppressive, respectively) during the periods of synergistic background locomotor activity in the step cycle. Contralaterally, however, the cBF and cST are reflexively activated as close synergists during these periods. 6. The reflex responses and their phase-dependent modulation are different for the homologous muscles in the two legs. Yet, some similarities are observed. These are present rather with respect to the phase of the corresponding leg than with respect to the phase of the stimulated leg. Both observations suggest that the phase-dependent reflex modulation is controlled separately in the ipsilateral and contralateral legs. 7. The response simultaneity in all investigated muscles supports the notion of a coordinated cutaneous interlimb reflex during human running.(ABSTRACT TRUNCATED AT 400 WORDS)