Bees with attitude: the effects of directed gusts on flight trajectories

Flight is a complicated task at small scales in part due to the ubiquitous unsteady air which contains it. Flying organisms deal with these difficulties using active and passive control mechanisms to steer their body motion. Body attitudes of flapping organisms are linked with their resultant flight trajectories and performance, yet little is understood about how discrete unsteady aerodynamic phenomena affect the interlaced dynamics of such systems. In this study, we examined freely flying bumblebees subject to a single discrete gust to emulate aerodynamic disturbances encountered in nature. Bumblebees are expert commanders of the aerial domain as they persistently forage within complex terrain elements. By tracking the three-dimensional dynamics of bees flying through gusts, we determined the sequences of motion that permit flight in three disturbance conditions: sideward, upward and downward gusts. Bees executed a series of passive impulsive maneuvers followed by active recovery maneuvers. Impulsive motion was unique in each gust direction, maintaining control by passive manipulation of the body. Bees pitched up and slowed-down at the beginning of recovery in every disturbance, followed by corrective maneuvers which brought attitudes back to their original state. Bees were displaced the most by the sideward gust, displaying large lateral translations and roll deviations. Upward gusts were easier for bees to fly through, causing only minor flight changes and minimal recovery times. Downward gusts severely impaired the control response of bees, inflicting strong adverse forces which sharply upset trajectories. Bees used a variety of control strategies when flying in each disturbance, offering new insights into insect-scale flapping flight and bio-inspired robotic systems.