Intersegmental coordination of limb movements during locomotion: mathematical models predict circuits that drive...

Normal locomotion in arthropods and vertebrates is a complex behavior, and the neural mechanisms that coordinate their limbs during locomotion at different speeds are unknown. The neural modules that drive cyclic movements of swimmerets respond to changes in excitation by changing the period of the motor pattern. As period changes, however, both intersegmental phase differences and the relative durations of bursts of impulses in different sets of motor neurons are preserved. To investigate these phenomena, we constructed a cellular model of the local pattern-generating circuit that drives each swimmeret. We then constructed alternative intersegmental circuits that might coordinate these local circuits. The structures of both the model of the local circuit and the alternative models of the coordinating circuit were based on and constrained by previous experimental results on pattern-generating neurons and coordinating interneurons. To evaluate the relative merits of these alternatives, we compared their dynamics with the performance of the real circuit when the level of excitation was changed. Many of the alternative coordinating circuits failed. One coordinating circuit, however, did effectively match the performance of the real system as period changed from 1 to 3.2 Hz. With this coordinating circuit, both the intersegmental phase differences and the relative durations of activity within each of the local modules fell within the ranges characteristic of the normal motor pattern and did not change significantly as period changed. These results predict a mechanism of coordination and a pattern of intersegmental connections in the CNS that is amenable to experimental test.