Dynamic stabilization of rapid hexapedal locomotion.

To stabilize locomotion, animals must generate forces appropriate to overcome the effects of perturbations and to maintain a desired speed or direction of movement. We studied the stabilizing mechanism employed by rapidly running insects by using a novel apparatus to perturb running cockroaches (Blaberus discoidalis). The apparatus used chemical propellants to accelerate a small projectile, generating reaction force impulses of less than 10 ms duration. The apparatus was mounted onto the thorax of the insect, oriented to propel the projectile laterally and loaded with propellant sufficient to cause a nearly tenfold increase in lateral velocity relative to maxima observed during unperturbed locomotion. Cockroaches were able to recover from these perturbations in 27+/-12 ms (mean +/- S.D., N=9) when running on a high-friction substratum. Lateral velocity began to decrease 13+/-5 ms (mean +/- S.D., N=11) following the start of a perturbation, a time comparable with the fastest reflexes measured in cockroaches. Cockroaches did not require step transitions to recover from lateral perturbations. Instead, they exhibited viscoelastic behavior in the lateral direction, with spring constants similar to those observed during unperturbed locomotion. The rapid onset of recovery from lateral perturbations supports the possibility that, during fast locomotion, intrinsic properties of the musculoskeletal system augment neural stabilization by reflexes.