Axial kinematics and muscle activity during terrestrial locomotion of the centipede Scolopendra heros

For centipedes moving steadily on a treadmill at speeds of 0.5, 1.0 and 1.5 L s-1, where L is body length, we obtained video tapes of movement that were synchronized with electromyograms (EMGs) from lateral flexor muscles at six standardized longitudinal positions. Field-by-field analysis of video tapes revealed posteriorly propagated waves of bending at all speeds. Muscle activity was also propagated posteriorly at the same speed as the kinematic wave, and EMGs of the lateral flexors were generally unilateral and alternating (between the left and right sides). The timing of EMG activity relative to lateral bending was consistent with electrical activity during the shortening of muscle fibers; therefore, activity of the axial musculature appears to cause lateral bending. Analysis of variance revealed widespread effects of speed on both kinematic and electromyographic variables, whereas longitudinal position within the centipede (between body segments 8 and 18) generally did not have significant effects on the same variables. For example, as speed increased from 0.5 to 1.5 L s-1, the amplitude of lateral displacement approximately doubled and the amplitude of lateral bending increased approximately threefold. Lag times (in seconds) indicating the propagation of kinematic and EMG events along the length of the centipede decreased significantly with speed. Phase lags among longitudinal sites decreased significantly with increased speed, indicating that the kinematic and EMG wavelengths increased with increased speed. EMG duration approximated 50 % of cycle duration and was unaffected by speed, and the phase of the EMG activity relative to lateral bending was also unaffected by locomotor speed. Hence, all results from all speeds are consistent with active bending of the axial segments during centipede locomotion, conflicting with the widely accepted hypothesis that lateral bending is imposed on the body by the metachronal stepping pattern of the legs and that bending is resisted by axial muscles.