Bd. Anderson et al., AXIAL KINEMATICS AND MUSCLE-ACTIVITY DURING TERRESTRIAL LOCOMOTION OFTHE CENTIPEDE SCOLOPENDRA-HEROS, Journal of Experimental Biology, 198(5), 1995, pp. 1185-1195
For centipedes moving steadily on a treadmill at speeds of 0.5, 1.0 an
d 1.5 L s(-1), where L is body length, we obtained video tapes of move
ment that were synchronized with electromyograms (EMGs) from lateral f
lexor muscles at six standardized longitudinal positions, Field-by-hel
d analysis of video tapes revealed posteriorly propagated waves of ben
ding 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 co
nsistent with electrical activity during the shortening of muscle fibe
rs; therefore, activity of the axial musculature appears to cause late
ral bending, Analysis of variance revealed widespread effects of speed
on both kinematic and electromyographic variables, whereas longitudin
al position within the centipede (between body segments 8 and 18) gene
rally did not have significant effects on the same variables, For exam
ple, as speed increased from 0.5 to 1.5 L s(-1), the amplitude of late
ral displacement approximately doubled and the amplitude of lateral be
nding increased approximately threefold, Lag times (in seconds) indica
ting the propagation of kinematic and EMG events along the length of t
he centipede decreased significantly with speed, Phase lags among long
itudinal sites decreased significantly with increased speed, indicatin
g that the kinematic and EMG wavelengths increased with increased spee
d, EMG duration approximated 50% of cycle duration and was unaffected
by speed, and the phase of the EMG activity relative to lateral bendin
g 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 steppi
ng pattern of the legs and that bending is resisted by axial muscles.