Neuromechanical models for insect locomotion: Stability, maneuverability, and proprioceptive feedback.

We describe a hierarchy of models for legged locomotion, emphasizing relationships among feedforward (preflexive) stability, maneuverability, and reflexive feedback. We focus on a hexapedal geometry representative of insect locomotion in the ground plane that includes a neural central pattern generator circuit, nonlinear muscles, and a representative proprioceptive sensory pathway. Although these components of the model are rather complex, neglect of leg mass yields a neuromechanical system with only three degrees of freedom, and numerical simulations coupled with a Poincaré map analysis shows that the feedforward dynamics is strongly stable, apart from one relatively slow mode and a neutral mode in body yaw angle. These modes moderate high frequency perturbations, producing slow heading changes that can be corrected by a stride-to-stride steering strategy. We show that the model's response to a lateral impulsive perturbation closely matches that of a cockroach subject to a similar impulse. We also describe preliminary studies of proprioceptive leg force feedback, showing how a reflexive pathway can reinforce the preflexive stability inherent in the system.