Kinematic and behavioral evidence for a distinction between trotting and ambling gaits in the cockroachBlaberus discoidalis

Abstract

Earlier observations had suggested that cockroaches might show multiple patterns of leg coordination, or gaits, but these were not followed by detailed behavioral or kinematic measurements that would allow a definite conclusion. We measured the walking speeds of cockroaches exploring a large arena and found that the body movements tended to cluster at one of two preferred speeds, either very slow (<10 cm s(-1)) or fairly fast (∼30 cm s(-1)). To highlight the neural control of walking leg movements, we experimentally reduced the mechanical coupling among the various legs by tethering the animals and allowing them to walk in place on a lightly oiled glass plate. Under these conditions, the rate of stepping was bimodal, clustering at fast and slow speeds. We next used high-speed videos to extract three-dimensional limb and joint kinematics for each segment of all six legs. The angular excursions and three-dimensional motions of the leg joints over the course of a stride were variable, but had different distributions in each gait. The change in gait occurs at a Froude number of ∼0.4, a speed scale at which a wide variety of animals show a transition between walking and trotting. We conclude that cockroaches do have multiple gaits, with corresponding implications for the collection and interpretation of data on the neural control of locomotion.