Stick insects walking on a wheel: Perturbations induced by obstruction of leg protraction

Summary

Stick insects (Carausius morosus) walking on a wheel were perturbed by restricting the forward protraction of individual legs. A barrier placed before a single middle or rear leg prevented that leg from reaching its normal protraction endpoint but allowed it unimpeded retraction. Upon striking the barrier, the protracting leg attempted to get past it and thereby prolonged protraction. This prolongation increased with the extent to which the obstruction infringed upon the leg's normal step range. Barriers placed near the midpoint of this range elicited large perturbations: the blocked leg often continued its protraction throughout many step cycles of the other legs (Fig. 1 E, F). For the most part walking was irregular and smooth forward progression was disrupted. Nevertheless, the infrequent steps by the affected leg usually were coordinated with those of the adjacent ipsilateral legs.

More rostral barrier positions elicited smaller perturbations: the blocked leg usually made one step in each step cycle of the other legs (Fig. 1 B, C, D, G). Measurements for these regular step sequences showed quantitatively that protraction duration increased in proportion to the severity of the infringement on normal leg movement (Figs. 3, 4). The fraction of the step period occupied by protraction increased from ca. 10% for normal walking to ca. 50% for caudal barrier positions. This proportionality is interpreted to show the importance of spatial components of the walking program.

When one leg was obstructed, its extended protraction influenced the stepping of the three adjacent legs as follows. First, the ipsilateral rostral leg showed the largest change: its protraction onset was regularly delayed for the duration of the extended protraction (Figs. 4, 7, 8), demonstrating a strong, centrally mediated inhibition. The presence of a further delay of up to 100 to 140 ms suggests that peripheral input from the protracting leg may be important for releasing this inhibition. Second, steps by an adjacent caudal leg were not measurably affected. However, the method may not have sufficed to reveal such effects because during regular walking middle leg protractions rarely lasted long enough to conflict with subsequent steps by the ipsilateral rear leg. Third, contralateral effects differed between middle and rear leg obstructions. If the obstructed leg was a middle leg, its extended protraction had little effect upon stepping by the contralateral middle leg: the latter leg frequently protracted while the blocked leg continued its protraction and there was no consistent change in the phase relation of these two legs (Table 1). In contrast, if the obstructed leg was a rear leg, protractions by the contralateral rear leg tended to be delayed (Table 1).