Flies fly at a broad range of speeds and produce sophisticated aerial maneuvers with precisely controlled wing movements. Remarkably, only subtle changes in wing motion are used by flies to produce aerial maneuvers, resulting in little directional tilt of aerodynamic force vector relative to the body. Therefore, it is often considered that flies fly according to a helicopter model and control speed mainly via force-vectoring enabled primarily by body-pitch change. Here we examine the speed control of blue bottle flies using a magnetically-levitated (MAGLEV) flight mill, as they fly at different body pitch and with different augmented aerodynamic damping. We identify wing kinematic contributors to the changes of estimated aerodynamic force through testing two force-vectoring models. Results show that in addition to body pitch, flies also use a collection of wing kinematic variables to control both force magnitude and direction, the roles of which are analogous to those of throttle, collective and cyclic pitch of helicopters. Our results also suggest that the MAGLEV flight mill system can be potentially used to study the roles of visual and mechanosensory feedback in insect flight control.
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