SUMMARY Flies are among the most agile of flying insects, a capacity that ultimately results from their nervous system's control over steering muscles and aerodynamic forces during flight. In order to investigate the relationships among neuromuscular control, musculo-skeletal mechanics and flight forces, we captured high-speed, three-dimensional wing kinematics of the blowfly, Calliphora vicina, while simultaneously recording electromyogram signals from prominent steering muscles during visually induced turns. We used the quantified kinematics to calculate the translational and rotational components of aerodynamic forces and moments using a theoretical quasi-steady model of force generation, confirmed using a dynamically scaled mechanical model of a Calliphora wing. We identified three independently controlled features of the wingbeat trajectory –downstroke deviation, dorsal amplitude and mode. Modulation of each of these kinematic features corresponded to both activity in a distinct steering muscle group and a distinct manipulation of the aerodynamic force vector. This functional specificity resulted from the independent control of downstroke and upstroke forces rather than the independent control of separate aerodynamic mechanisms. The predicted contributions of each kinematic feature to body lift, thrust, roll, yaw and pitch are discussed.
SUMMARY Determining how the motor patterns of the nervous system are converted into the mechanical and behavioral output of the body is a central goal in the study of locomotion. In the case of dipteran flight, a population of small steering muscles controls many of the subtle changes in wing kinematics that allow flies to maneuver rapidly. We filmed the wing motion of tethered Calliphora vicina at high speed and simultaneously recorded multi-channel electromyographic signals from some of the prominent steering muscles in order to correlate kinematics with muscle activity. Using this analysis, we found that the timing of each spike in the basalare muscles was strongly correlated with changes in the deviation of the stroke plane during the downstroke. The relationship was non-linear such that the magnitude of the kinematic response to each muscle spike decreased with increasing levels of stroke deviation. This result suggests that downstroke deviation is controlled in part via the mechanical summation of basalare activity. We also found that interactions among the basalares and muscles III2–III4 determine the maximum forward amplitude of the wingstroke. In addition, activity in muscle I1 appears to participate in a wingbeat gearing mechanism, as previously proposed. Using these results, we have been able to correlate changes in wing kinematics with alteration in the spike rate, firing phase and combinatorial activity of identified steering muscles.
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