CORRECTIVE RESPONSES TO LOSS OF GROUND SUPPORT DURING WALKING .2. COMPARISON OF INTACT AND CHRONIC SPINAL CATS

1. The preceding study described a corrective response in cats when one hind leg steps into a hole. In this investigation we examine the extent to which this behavior is organized at the spinal level by comparing the responses elicited in intact and chronic spinal cats. 2. Adult cats were trained to step bipedally with their hind legs on a treadmill. After training, the responses to stepping into a hole cut in the treadmill belt were monitored with a video recorder and by recording electromyograms from muscles in both hind legs. The responses to stepping into the hole were also recorded in chronic spinal cats that had recovered the ability to step with their hind legs a few weeks after spinalization. 3. The behavioral responses in the two groups of animals differed in two respects. First, the latency of the onset of the flexion movement to remove the foot from the hole was shorter in intact animals (70-150 ms in intact vs. 130-350 ms in spinal animals). Second, the flexion movement in the intact animals was stronger. The exaggerated flexion movement in intact animals lifted the paw well clear of the hole and allowed support to be regained on the treadmill belt. The weaker flexion movement in spinal animals was usually insufficient to lift the paw completely from the hole. 4. Differences in the motor patterns recorded from flexor muscles during the corrective response in intact and spinal animals correspond with the differences in the kinematics. First, the onset of flexor activity after the foot entered the hole was delayed by similar to 100 ms in spinal animals relative to intact animals. Second, in intact animals the magnitudes of flexor bursts were increased relative to the flexor bursts associated with the swing phase during stepping, whereas in spinal animals flexor bursts during the corrective response resembled those occurring during swing. 5. Similarities in the duration and the timing of bursts in different flexor muscles in intact and spinal animals during the corrective response and during swing indicated that the corrective response involves activation of the spinal system that normally produces swing phase flexor activity. We conclude that activation of this system is facilitated by input from supraspinal structures during the corrective response in intact animals. 6. In all intact animals and three of five spinal animals, support of the hindquarters when the foot entered the hole was maintained by the contralateral leg. The stance phase of this leg was prolonged and extension significantly increased. The magnitude of extensor activity in the contralateral leg increased similar to 60 ms after the ipsilateral leg entered the hole, presumably because of the additional loading. These similarities indicate that the mechanism by which the contralateral leg maintains support during the corrective response is organized primarily at the spinal level.