Early and late stretch responses of human foot muscles induced by perturbation of stance

In eight subjects standing on a movable platform, surface EMG activity was recorded from the foot muscles extensor digitorum brevis (EDB) and flexor digitorum brevis (FDB) and from the leg muscles soleus (Sol) and tibialis anterior (TA) during perturbations of upright stance. Perturbations inducing foot dorsiflexion (upward tilt and backward translation) evoked a short-latency response (SLR) and a medium-latency response (MLR) to stretch in the physiological extensors FDB and Sol, and a long-latency response (LLR) in the physiological flexors EDB and TA. Perturbations inducing plantar-flexion (downward tilt and forward translation) evoked the MLR in EDB and TA, and the LLR in FDB and Sol. The latency of the FDB and Sol SLR was compared to that of the H and T reflexes evoked in the same muscles by electrical or mechanical stimulation, respectively. In both muscles, the T reflex and the SLR followed the H reflex at delays accounted for by the different stimulation mode, indicating that the SLR induced in both muscles by upward tilt and backward translation was a true autogenetic stretch reflex from spindle primaries. The time interval between the onset of SLR and of MLR was significantly greater for the FDB than the Sol muscle, suggesting that MLR is a spinal reflex travelling through slower peripheral afferent pathways than SLR. From these latency differences and from the distance between the muscles, we calculated in four subjects the conduction velocity of the afferent fibres presumably responsible for the MLR in FDB. This was about 29 m/s. LLRs were evoked in TA and EDB during upward tilt and backward translation, and in Sol and FDB during downward tilt, but not forward translation. LLRs did not adhere to a proximal-to-distal pattern, since these could appear earlier in the foot than in the leg muscles. All responses were modulated by perturbation type (tilt vs translation) and body posture (normal stance vs forward leaning). Both the large amplitude of the foot muscle responses and their temporal pattern indicate that the muscles acting on the toes play a major role in stabilising posture. Their action increases in amplitude and extends in time the foot-ground reaction force, thereby improving the efficiency of the superimposed action of the leg muscle responses.