Acoustic startle/escape reactions in tethered flying locusts: motor patterns and wing kinematics underlying intentional steering.

We simultaneously recorded flight muscle activity and wing kinematics in tethered, flying locusts to determine the relationship between asymmetric depressor muscle activation and the kinematics of the stroke reversal at the onset of wing depression during attempted intentional steering manoeuvres. High-frequency, pulsed sounds produced bilateral asymmetries in forewing direct depressor muscles (M97, 98, 99) that were positively correlated with asymmetric forewing depression and asymmetries in stroke reversal timing. Bilateral asymmetries in hindwing depressor muscles (M127 and M128 but not M129) were positively correlated with asymmetric hindwing depression and asymmetries in the timing of the hindwing stroke reversal; M129 was negatively correlated with these shifts. Hindwing depressor asymmetries and wing kinematic changes were smaller and shifted in opposite direction than corresponding measurements of the forewings. These findings suggest that intentional steering manoeuvres employ bulk shifts in depressor muscle timing that affect the timing of the stroke reversals thereby establishing asymmetric wing depression. Finally, we found indications that locusts may actively control the timing of forewing rotation and speculate this may be a mechanism for generating steering torques. These effects would act in concert with forces generated by asymmetric wing depression and angle of attack to establish rapid changes in direction.