Neuromuscular control of aerodynamic forces and moments in the blowfly, Calliphora vicina

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.