Active control of free flight manoeuvres in a hawkmoth, Agrius convolvuli

By combining optical triangulation with the comb-fringe technique and dual-channel telemetry, wing kinematics and body attitudes accompanying muscle activities of free-flying male hawkmoths were recorded synchronously when they performed flight manoeuvres elicited by a female sex pheromone. The results indicate that the wing leading edge angular position at the ventral stroke reversal, which can be decomposed by two orthogonal angular parameters (a flapping angle and a deviation angle), is well controllable. Two specific flight muscles, the dorsal-ventral muscle (DVM, indirect muscle, a wing elevator) and the third axillary muscle (3AXM, direct muscle, a wing retractor), can modulate the flapping angle and the deviation angle, respectively, by means of regulating the firing timing of muscle activities. The firing timing can be expressed by the firing latency absolutely, which is just before the timing of ventral stroke reversal. The results illustrate that lengthening the firing latency of the DVM and of the 3AXM can increase the flapping angle and the deviation angle, respectively, which both strengthen the downstroke at the ventral stroke reversal. The relationship of bilateral asymmetry shows that the bilateral differences in the firing latency of the DVM and of the 3AXM will cause bilateral differences in the wing position, which accompany the variations of yaw and roll angles in time course. This implies the contribution of the two muscles to active steering controls during turning or banking, though the DVM being an indirect muscle was generally treated as a power generator. Finally, the relationship between the pitch angle and the 3AXM latency, deduced from the relationships between the pitch angle and the deviation angle and between the deviation angle and the 3AXM latency, shows that lengthening the 3AXM latency can increase the pitch angle at the ventral stroke reversal by moving the wing tip far away from the centre of gravity of the body, which indicates a functional role of the 3AXM in active pitching control.