Coordination of fore and hind leg stepping in cats on a transversely-split treadmill.

To gain insight into the mechanism of coordination of stepping in the fore and hind legs of quadrupeds, we examined the kinematics of leg movements and the motor patterns in fore and hind leg flexor muscles in decerebrate walking cats when the two pairs of legs stepped on separate treadmills running at different speeds. When the front treadmill was slowed progressively from 0.6 to 0.3 m/s with the rear treadmill running at 0.6 m/s, the rate of stepping in both the fore and hind legs decreased and a 1:1 stepping ratio was maintained. The decrease in the rate of stepping in the hind legs was due primarily to an increase in the duration of the swing phase. Slowing the speed of the rear treadmill while keeping the front treadmill speed at 0.6 m/s decreased the rate of stepping of the hind legs, but had relatively little influence on the average rate of stepping in the forelegs. In this situation stepping in the fore and hind legs was uncoupled and the time of stepping in one hind leg relative to the ipsilateral foreleg progressively shifted during a walking sequence. Analysis of the timing of electromyographic (EMG) recordings from flexor muscles of the hip and elbow joints yielded insight into the neuronal mechanisms underlying the asymmetry in slowing either the front or rear treadmill. We propose that ipsilateral pattern generating networks are asymmetrically coupled via descending inhibitory pathways and an ascending excitatory pathway. We discuss how the characteristics of these linkages are functionally appropriate for establishing the normal timing of stepping in the hind and forelegs during slow walking.