The demonstrated importance of unsteady effects in insect flight has prompted recent development of better experimental and computational tools to investigate unsteady forces and vortical flows around a flapping wing. In particular, dynamically scaled robots of both the hawkmoth (Ellington et al., 1996) and fruitfly (Dickinson et al., 1999) have been

We investigate aspects of hovering insect flight by finding the optimal wing kinematics which minimize power consumption while still providing enough lift to maintain a time-averaged constant altitude over one flapping period. In particular, we study the flight of three insects whose masses vary by approximately three orders of magnitude: fruitfly (Drosophila melanogaster), bumblebee (Bombus terrestris), and hawkmoth (Manduca sexta). Here, we model an insect wing as a rigid body with three rotational degrees of freedom. The aerodynamic forces are modelled via a quasisteady model of a thin plate interacting with the surrounding fluid. The advantage of this model, as opposed to the more computationally costly method of direct numerical simulation via computational fluid dynamics, is that it allows us to perform optimization procedures and detailed sensitivity analyses which require many cost function evaluations. The optimal solutions are found via a hybrid optimization algorithm combining aspects of a genetic algorithm and a gradient-based optimizer. We find that the results of this optimization yield kinematics which are qualitatively and quantitatively similar to previously observed data. We also perform sensitivity analyses on parameters of the optimal kinematics to gain insight into the values of the observed optima. Additionally, we find that all of the optimal kinematics found here maintain the same leading edge throughout the stroke, as is the case for nearly all insect wing motions. We show that this type of stroke takes advantage of a passive wing rotation in which aerodynamic forces help to reverse the wing pitch, similar to the turning of a free-falling leaf.

We investigate the aerodynamics of freely falling plates in a quasi-two-dimensional flow at Reynolds number of 10 3 , which is typical for a leaf or business card falling in air. We quantify the trajectories experimentally using high-speed digital video at sufficient resolution to determine the instantaneous plate accelerations and thus to deduce the instantaneous fluid forces. We compare the measurements with direct numerical solutions of the two-dimensional Navier-Stokes equation. Using inviscid theory as a guide, we decompose the fluid forces into contributions due to acceleration, translation, and rotation of the plate. For both fluttering and tumbling we find that the fluid circulation is dominated by a rotational term proportional to the angular velocity of the plate, as opposed to the translational velocity for a glider with fixed angle of attack. We find that the torque on a freely falling plate is small, i.e. the torque is one to two orders of magnitude smaller than the torque on a glider with fixed angle of attack. Based on these results we revise the existing ODE models of freely falling plates. We get access to different kinds of dynamics by exploring the phase diagram spanned by the Reynolds number, the dimensionless moment of inertia, and the thickness-to-width ratio. In agreement with previous experiments, we find fluttering, tumbling, and apparently chaotic motion. We further investigate the dependence on initial conditions and find brief transients followed by periodic fluttering described by simple harmonics and tumbling with a pronounced period-two structure. Near the cusp-like turning points, the plates elevate, a feature which would be absent if the lift depended on the translational velocity alone.

What force does an insect wing generate?" Finding answers to this enduring question is an essential step toward our understanding of interactions of moving objects with fluids that enable most living species such as insects, birds, and fish to travel efficiently and us to follow similar suit with sails, oars, and airfoils. We give a brief history of research in insect flight and discuss recent findings in unsteady aerodynamics of flapping flight at intermediate range Reynolds numbers (10-10 4). In particular, we examine the unsteady mechanisms in uniform and accelerated motions, forward and hovering flight, as well as passive flight of free-falling objects. The results obtained by "taking the insects apart" helped us to resolve previous puzzles about the force estimates in hovering insects, to ellucidate basic mechanisms essential to flapping flight, and to gain insights about the efficieny of flight.

Just as the Wright brothers implemented controls to achieve stable airplane flight, flying insects have evolved behavioral strategies that ensure recovery from flight disturbances. Pioneering studies performed on tethered and dissected insects demonstrate that the sensory, neurological, and musculoskeletal systems play important roles in flight control. Such studies, however, cannot produce an integrative model of insect flight stability because they do not incorporate the interaction of these systems with free-flight aerodynamics. We directly investigate control and stability through the application of torque impulses to freely flying fruit flies (Drosophila melanogaster) and measurement of their behavioral response. High-speed video and a new motion tracking method capture the aerial "stumble," and we discover that flies respond to gentle disturbances by accurately returning to their original orientation. These insects take advantage of a stabilizing aerodynamic influence and active torque generation to recover their heading to within 2°in <60 ms. To explain this recovery behavior, we form a feedback control model that includes the fly's ability to sense body rotations, process this information, and actuate the wing motions that generate corrective aerodynamic torque. Thus, like early man-made aircraft and modern fighter jets, the fruit fly employs an automatic stabilization scheme that reacts to short time-scale disturbances.flight control | insect flight | stability | perturbation | fruit fly L ocomotion through natural environments demands mechanisms that maintain stability in the face of unpredictable disturbances. Behavioral strategies play a particularly important role in controlling the flight of insects (1-7), because even gentle air currents can cause large disruptions to the intended flight path. Insects must also contend with the intrinsic instability of flapping flight (8, 9) and the large fluctuations in aerodynamic forces caused by slight variations in wing motions (10, 11). Corrective behavior often takes advantage of vision (1, 2). For fruit flies, however, reaction time to visual stimuli is at least 10 wingbeats (12), so these insects must employ faster sensory circuits to recover from short time-scale disturbances and instabilities. To probe this fast control strategy, we devised an experimental method that imposes impulsive mechanical disturbances (6, 13) to flying insects while allowing us to measure relevant aspects of flight behavior. We first glue tiny ferromagnetic pins to fruit flies and image their free flight using three orthogonally oriented highspeed video cameras (Methods and SI Text). When a fly enters the filming volume, an optical trigger detects the insect, initiates recording, and activates a pair of Helmholtz coils that produce a magnetic field. The field and pin are both oriented horizontally, so the resulting torque on the pin reorients the yaw, or heading angle, of the insect (Fig. 1). We then use a new motion tracking technique to extract the three-dimensional body and w...