Cockroaches are remarkably stable runners, exhibiting rapid recovery from external perturbations. To uncover the mechanisms behind this important behavioral trait, we recorded leg kinematics of freely running animals in both undisturbed and perturbed trials. Functional coupling underlying inter-leg coordination was monitored before and during localized perturbations, which were applied to single legs via magnetic impulses. The resulting transient effects on all legs and the recovery times to normal pre-perturbation kinematics were studied. We estimated coupling architecture and strength by fitting experimental data to a six-leg-unit phase oscillator model. Using maximum-likelihood techniques, we found that a network with nearest-neighbor inter-leg coupling best fitted the data and that, although coupling strengths vary among preparations, the overall inputs entering each leg are approximately balanced and consistent. Simulations of models with different coupling strengths encountering perturbations suggest that the coupling schemes estimated from our experiments allow animals relatively fast and uniform recoveries from perturbations.
The American cockroach, Periplaneta americana, provides a successful model for the study of legged locomotion. Sensory regulation and the relative importance of sensory feedback vs. central control in animal locomotion are key aspects in our understanding of locomotive behavior. Here we introduce the cockroach model and describe the basic characteristics of the neural generation and control of walking and running in this insect. We further provide a brief overview of some recent studies, including mathematical modeling, which have contributed to our knowledge of sensory control in cockroach locomotion. We focus on two sensory mechanisms and sense organs, those providing information related to loading and unloading of the body and the legs, and leg-movement-related sensory receptors, and present evidence for the instrumental role of these sensory signals in inter-leg locomotion control. We conclude by identifying important open questions and indicate future perspectives.
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