Flies compensate for unilateral wing damage through modular adjustments of wing and body kinematics.

Using high-speed videography, we investigated how fruit flies compensate for unilateral wing damage, in which loss of area on one wing compromises both weight support and roll torque equilibrium. Our results show that flies control for unilateral damage by rolling their body towards the damaged wing and by adjusting the kinematics of both the intact and damaged wings. To compensate for the reduction in vertical lift force due to damage, flies elevate wingbeat frequency. Because this rise in frequency increases the flapping velocity of both wings, it has the undesired consequence of further increasing roll torque. To compensate for this effect, flies increase the stroke amplitude and advance the timing of pronation and supination of the damaged wing, while making the opposite adjustments on the intact wing. The resulting increase in force on the damaged wing and decrease in force on the intact wing function to maintain zero net roll torque. However, the bilaterally asymmetrical pattern of wing motion generates a finite lateral force, which flies balance by maintaining a constant body roll angle. Based on these results and additional experiments using a dynamically scaled robotic fly, we propose a simple bioinspired control algorithm for asymmetric wing damage.

Body saccades of Drosophila consist of stereotyped banked turns.

The flight pattern of many fly species consists of straight flight segments interspersed with rapid turns called body saccades, a strategy that is thought to minimize motion blur. We analyzed the body saccades of fruit flies (Drosophila hydei), using high-speed 3D videography to track body and wing kinematics and a dynamically scaled robot to study the production of aerodynamic forces and moments. Although the size, degree and speed of the saccades vary, the dynamics of the maneuver are remarkably stereotypic. In executing a body saccade, flies perform a quick roll and counter-roll, combined with a slower unidirectional rotation around their yaw axis. Flies regulate the size of the turn by adjusting the magnitude of torque that they produce about these control axes, while maintaining the orientation of the rotational axes in the body frame constant. In this way, body saccades are different from escape responses in the same species, in which the roll and pitch component of banking is varied to adjust turn angle. Our analysis of the wing kinematics and aerodynamics showed that flies control aerodynamic torques during the saccade primarily by adjusting the timing and amount of span-wise wing rotation.

Strategies for the stabilization of longitudinal forward flapping flight revealed using a dynamically-scaled robotic fly.

The ability to regulate forward speed is an essential requirement for flying animals. Here, we use a dynamically-scaled robot to study how flapping insects adjust their wing kinematics to regulate and stabilize forward flight. The results suggest that the steady-state lift and thrust requirements at different speeds may be accomplished with quite subtle changes in hovering kinematics, and that these adjustments act primarily by altering the pitch moment. This finding is consistent with prior hypotheses regarding the relationship between body pitch and flight speed in fruit flies. Adjusting the mean stroke position of the wings is a likely mechanism for trimming the pitch moment at all speeds, whereas changes in the mean angle of attack may be required at higher speeds. To ensure stability, the flapping system requires additional pitch damping that increases in magnitude with flight speed. A compensatory reflex driven by fast feedback of pitch rate from the halteres could provide such damping, and would automatically exhibit gain scheduling with flight speed if pitch torque was regulated via changes in stroke deviation. Such a control scheme would provide an elegant solution for stabilization across a wide range of forward flight speeds.

Flies evade looming targets by executing rapid visually directed banked turns.

Avoiding predators is an essential behavior in which animals must quickly transform sensory cues into evasive actions. Sensory reflexes are particularly fast in flying insects such as flies, but the means by which they evade aerial predators is not known. Using high-speed videography and automated tracking of flies in combination with aerodynamic measurements on flapping robots, we show that flying flies react to looming stimuli with directed banked turns. The maneuver consists of a rapid body rotation followed immediately by an active counter-rotation and is enacted by remarkably subtle changes in wing motion. These evasive maneuvers of flies are substantially faster than steering maneuvers measured previously and indicate the existence of sensory-motor circuitry that can reorient the fly's flight path within a few wingbeats.

The influence of sensory delay on the yaw dynamics of a flapping insect.

In closed-loop systems, sensor feedback delays may have disastrous implications for performance and stability. Flies have evolved multiple specializations to reduce this latency, but the fastest feedback during flight involves a delay that is still significant on the timescale of body dynamics. We explored the effect of sensor delay on flight stability and performance for yaw turns using a dynamically scaled robotic model of the fruitfly, Drosophila. The robot was equipped with a real-time feedback system that performed active turns in response to measured torque about the functional yaw axis. We performed system response experiments for a proportional controller in yaw velocity for a range of feedback delays, similar in dimensionless timescale to those experienced by a fly. The results show a fundamental trade-off between sensor delay and permissible feedback gain, and suggest that fast mechanosensory feedback in flies, and most probably in other insects, provide a source of active damping which compliments that contributed by passive effects. Presented in the context of these findings, a control architecture whereby a haltere-mediated inner-loop proportional controller provides damping for slower visually mediated feedback is consistent with tethered-flight measurements, free-flight observations and engineering design principles.