AbstractCentimeter-scale fliers that combine wings with springy elements must contend with the high power requirements and mechanical constraints of flapping wing flight. Insects utilize elastic energy exchange to reduce the inertial costs of flapping wing flight and potentially match wingbeat frequencies to a mechanical resonance. Flying at resonance may be energetically favorable under steady conditions, but it is difficult to modulate the frequency of a resonant system. Evidence suggests that insects utilize frequency modulation over long time scales to adjust aerodynamic forces, but it remains an open question the extent to which insects can modulate frequency on the wingstroke-to-wingstroke timescale. If wingbeat frequencies deviate from resonance, the musculature must work against the elastic flight system, thereby potentially increasing energetic costs. To assess how insects address the simultaneous needs for power and control, we tested the capacity for wingstroke-to-wingstroke wingbeat frequency modulation by perturbing free hovering Manduca sexta with vortex rings while recording high-speed video at 2000 fps. Because hawkmoth flight muscles are synchronous, there is at least the potential for the nervous system to modulate frequency on each wingstroke. We observed ± 16% wingbeat frequency modulation in just a few wing strokes. Via instantaneous phase analysis of wing kinematics, we found that over 85% of perturbation responses required active changes in motor input frequency. Unlike their robotic counterparts that explicitly abdicate frequency modulation in favor of energy efficiency, we find that wingstroke-to-wingstroke frequency modulation is an underappreciated control strategies that complements other strategies for maneuverability and stability in insect flight.
Hawkmoths use wingstroke-to-wingstroke frequency modulation for aerial recovery to vortex ring perturbations
