Static stability predicts the continuum of interleg coordination patterns in Drosophila

Abstract

During walking, insects must coordinate the movements of their six legs for efficient locomotion. This interleg coordination is speed dependent: fast walking in insects is associated with tripod coordination patterns, whereas slow walking is associated with more variable, tetrapod-like patterns. To date, however, there has been no comprehensive explanation as to why these speeddependent shifts in interleg coordination should occur in insects. Tripod coordination would be sufficient at low walking speeds. The fact that insects use a different interleg coordination pattern at lower speeds suggests that it is more optimal or advantageous at these speeds. Furthermore, previous studies focused on discrete tripod and tetrapod coordination patterns. Experimental data, however, suggest that changes observed in interleg coordination are part of a speed-dependent spectrum. Here, we explore these issues in relation to static stability as an important aspect for interleg coordination in Drosophila. We created a model that uses basic experimentally measured parameters in fruit flies to find the interleg phase relationships that maximize stability for a given walking speed. The model predicted a continuum of interleg coordination patterns spanning the complete range of walking speeds as well as an anteriorly directed swing phase progression. Furthermore, for low walking speeds, the model predicted tetrapod-like patterns to be most stable, whereas at high walking speeds, tripod coordination emerged as most optimal. Finally, we validated the basic assumption of a continuum of interleg coordination patterns in a large set of experimental data from walking fruit flies and compared these data with the model-based predictions.