Interlimb coordination during stepping in the cat: in-phase stepping and gait transitions

Abstract

The coordination of step cycles between all 4 limbs during in-phase stepping and during transitions to and from alternate stepping was studied in 12 adult cats during repeated overground stepping trials. The temporal spacing of step cycles of the different limbs was determined from analysis of electromyographic (EMG) activity in a single extensor muscle of each limb. Patterns of coordination of the different limbs were established on the basis of the frequency with which phase values separating step cycles were encountered. Steps in which the phasing of step cycles of the two hindlimbs were closer to true in-phase coordination than true alternation (phase between 270 degrees and 90 degrees) and where similar coupling was found in both the preceding and following steps were defined as steady state conditions. Distinct patterns of coordination of forelimb-forelimb and forelimb-hindlimb step cycles were noted under steady state conditions. During stepping sequences which include transitions either to or from alternate stepping, both gradual and abrupt phase changes were found. The changes in both forelimb-forelimb and forelimb-hindlimb phase relationships were more often gradual than abrupt. Where abrupt changes were encountered in the change in phase relationships between one such limb pair the phase change in the other pair was gradual. Changes in hindlimb-hindlimb phase relationships during transitions were nearly always abrupt. It is concluded that the 4 limbs are coordinated during in-phase stepping according to a few patterns, but that the variability about these patterns makes their association with simple neural circuitry rather speculative. The finding that transitions were most often gradual is interpreted in terms of a state-dependent model of interlimb control, in which the type of transition utilized depends on the strength of neural coupling of step cycles of all 4 limbs at the time that the transition is initiated.