Intersegmental Coordination: Influence of a Single Walking Leg on the Neighboring Segments in the Stick Insect Walking System

Abstract

A key element of walking is the coordinated interplay of multiple limbs to achieve a stable locomotor pattern that is adapted to the environment. We investigated intersegmental coordination of walking in the stick insect, Carausius morosus by examining the influence a single stepping leg has on the motoneural activity of the other hemiganglia, and whether this influence changes with the walking direction. We used a reduced single leg walking preparation with only one intact front, middle, or hind leg. The intact leg performed stepping movements on a treadmill, thus providing intersegmental signals about its stepping to the other hemiganglia. The activity of coxal motoneurons was simultaneously recorded extracellularly in all other segments. Stepping sequences of any given single leg in either walking direction were accompanied by an increase in coxal motoneuron (MN) activity of all other segments, which was mostly modulated and slightly in phase with stance of the walking leg. In addition, forward stepping of the front leg and, to a lesser extent, backward stepping of the hind leg elicited alternating activity in mesothoracic coxal MNs. Forward and backward stepping of the middle leg did not elicit alternating activity in coxal MNs in any other hemiganglia, indicating that the influence of middle leg stepping is qualitatively different from that of forward front and backward hind leg stepping. Our results indicate that in an insect walking system individual segments differ with respect to their intersegmental influences and thus cannot be treated as similar within the chain of segmental walking pattern generators. Consequences for the current concepts on intersegmental coordination are discussed.