William B. Dickson

Force production and flow structure of the leading edge vortex on flapping wings at high and low Reynolds numbers

SUMMARY The elevated aerodynamic performance of insects has been attributed in part to the generation and maintenance of a stable region of vorticity known as the leading edge vortex (LEV). One explanation for the stability of the LEV is that spiraling axial flow within the vortex core drains energy into the tip vortex, forming a leading-edge spiral vortex analogous to the flow structure generated by delta wing aircraft. However, whereas spiral flow is a conspicuous feature of flapping wings at Reynolds numbers (Re) of 5000, similar experiments at Re=100 failed to identify a comparable structure. We used a dynamically scaled robot to investigate both the forces and the flows created by a wing undergoing identical motion at Re of∼ 120 and ∼1400. In both cases, motion at constant angular velocity and fixed angle of attack generated a stable LEV with no evidence of shedding. At Re=1400, flow visualization indicated an intense narrow region of spanwise flow within the core of the LEV, a feature conspicuously absent at Re=120. The results suggest that the transport of vorticity from the leading edge to the wake that permits prolonged vortex attachment takes different forms at different Re.

The effect of advance ratio on the aerodynamics of revolving wings

SUMMARY Recent studies have demonstrated that a quasi-steady model closely matches the instantaneous force produced by an insect wing during hovering flight. It is not clear, however, if such methods extend to forward flight. In this study we use a dynamically scaled robotic model of the fruit fly Drosophila melanogaster to investigate the forces produced by a wing revolving at constant angular velocity while simultaneously translating at velocities appropriate for forward flight. Because the forward and angular velocities were constant wing inertia was negligible, and the measured forces can be attributed to fluid dynamic phenomena. The combined forward and revolving motions of the wing produce a time-dependent free-stream velocity profile, which suggests that added mass forces make a contribution to the measured forces. We find that the forces due added mass make a small, but measurable, component of the total force and are in excellent agreement with theoretical values. Lift and drag coefficients are calculated from the force traces after subtracting the contributions due to added mass. The lift and drag coefficients, for fixed angle of attack, are not constant for non-zero advance ratios, but rather vary in magnitude throughout the stroke. This observation implies that modifications of the quasi-steady model are required in order to predict accurately the instantaneous forces produced during forward flight. We show that the dependence of the lift and drag coefficients upon advance ratio and stroke position can be characterized effectively in terms of the tip velocity ratio – the ratio of the chordwise components of flow velocity at the wing tip due to translation and revolution. On this basis we develop a modified quasi-steady model that can account for the varying magnitudes of the lift and drag coefficients. Our model may also resolve discrepancies in past measurements of wing performance based on translational and revolving motion.

Leading-Edge Vortices Elevate Lift of Autorotating Plant Seeds

Helicopter Seed Lift The “helicopter” seeds of maple trees and other similar autorotating seeds detach from their parent tree under windy conditions and gyrate as they are dispersed by the wind. The reproductive success of the tree depends on the flight performance of its seeds. Autorotating seeds are known to generate high lift as they slowly descend through the air, but the means by which they do so is unclear. Lentink et al. (p. 1438, see the cover) have elucidated the aerodynamic mechanism for high lift in autorotating seeds using a robot model seed and a three-dimensional flow measurement technique. Maple seeds and a hornbeam seed create a prominent leading-edge vortex that is similar to the flow structures that are responsible for the high lift generated by the wings of hovering insects and bats. Thus, both animals and plants have converged on an identical aerodynamic solution for generating lift. Winged plant seeds use leading edge vortices to create lift, in the same way that flying animals do. As they descend, the autorotating seeds of maples and some other trees generate unexpectedly high lift, but how they attain this elevated performance is unknown. To elucidate the mechanisms responsible, we measured the three-dimensional flow around dynamically scaled models of maple and hornbeam seeds. Our results indicate that these seeds attain high lift by generating a stable leading-edge vortex (LEV) as they descend. The compact LEV, which we verified on real specimens, allows maple seeds to remain in the air more effectively than do a variety of nonautorotating seeds. LEVs also explain the high lift generated by hovering insects, bats, and possibly birds, suggesting that the use of LEVs represents a convergent aerodynamic solution in the evolution of flight performance in both animals and plants.

Integrative Model of Drosophila Flight

This paper presents a framework for simulating the flight dynamics and control strategies of the fruit fly Drosophila melanogaster. The framework consists of five main components: an articulated rigid-body simulation, a model of the aerodynamic forces and moments, a sensory systems model, a control model, and an environment model. In the rigid-body simulation the fly is represented by a system of three rigid bodies connected by a pair of actuated ball joints. At each instant of the simulation, the aerodynamic forces and moments acting on the wings and body of the fly are calculated using an empirically derived quasi-steady model. The pattern of wing kinematics is based on data captured from high-speed video sequences. The forces and moments produced by the wings are modulated by deforming the base wing kinematics along certain characteristic actuation modes. Models of the fly’s visual and mechanosensory systems are used to generate inputs to a controller that sets the magnitude of each actuation mode, thus modulating the forces produced by the wings. This simulation framework provides a quantitative test bed for examining the possible control strategies employed by flying insects. Examples demonstrating pitch rate, velocity, altitude, and flight speed control, as well as visually guided centering in a corridor are presented.

A linear systems analysis of the yaw dynamics of a dynamically scaled insect model

SUMMARY Recent studies suggest that fruit flies use subtle changes to their wing motion to actively generate forces during aerial maneuvers. In addition, it has been estimated that the passive rotational damping caused by the flapping wings of an insect is around two orders of magnitude greater than that for the body alone. At present, however, the relationships between the active regulation of wing kinematics, passive damping produced by the flapping wings and the overall trajectory of the animal are still poorly understood. In this study, we use a dynamically scaled robotic model equipped with a torque feedback mechanism to study the dynamics of yaw turns in the fruit fly Drosophila melanogaster. Four plausible mechanisms for the active generation of yaw torque are examined. The mechanisms deform the wing kinematics of hovering in order to introduce asymmetry that results in the active production of yaw torque by the flapping wings. The results demonstrate that the stroke-averaged yaw torque is well approximated by a model that is linear with respect to both the yaw velocity and the magnitude of the kinematic deformations. Dynamic measurements, in which the yaw torque produced by the flapping wings was used in real-time to determine the rotation of the robot, suggest that a first-order linear model with stroke-average coefficients accurately captures the yaw dynamics of the system. Finally, an analysis of the stroke-average dynamics suggests that both damping and inertia will be important factors during rapid body saccades of a fruit fly.

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.

Neuromuscular control of wingbeat kinematics in Anna's hummingbirds (Calypte anna)

SUMMARY Hummingbirds can maintain the highest wingbeat frequencies of any flying vertebrate – a feat accomplished by the large pectoral muscles that power the wing strokes. An unusual feature of these muscles is that they are activated by one or a few spikes per cycle as revealed by electromyogram recordings (EMGs). The relatively simple nature of this activation pattern provides an opportunity to understand how motor units are recruited to modulate limb kinematics. Hummingbirds made to fly in low-density air responded by moderately increasing wingbeat frequency and substantially increasing the wing stroke amplitude as compared with flight in normal air. There was little change in the number of spikes per EMG burst in the pectoralis major muscle between flight in normal and low-density heliox (mean=1.4 spikes cycle−1). However the spike amplitude, which we take to be an indication of the number of active motor units, increased in concert with the wing stroke amplitude, 1.7 times the value in air. We also challenged the hummingbirds using transient load lifting to elicit maximum burst performance. During maximum load lifting, both wing stroke amplitude and wingbeat frequency increased substantially above those values during hovering flight. The number of spikes per EMG burst increased to a mean of 3.3 per cycle, and the maximum spike amplitude increased to approximately 1.6 times those values during flight in heliox. These results suggest that hummingbirds recruit additional motor units (spatial recruitment) to regulate wing stroke amplitude but that temporal recruitment is also required to maintain maximum stroke amplitude at the highest wingbeat frequencies.

Unsteadiness in Flow over a Flat Plate at Angle-of-Attack at Low Reynolds Numbers

Flow over an impulsively started low-aspect-ratio flat plate at angle-of-attack is investigated for a Reynolds number of 300. Numerical simulations, validated by a companion experiment, are performed to study the influence of aspect ratio, angle of attack, and planform geometry on the interaction of the leading-edge and tip vortices and resulting lift and drag coecients. Aspect ratio is found to significantly influence the wake pattern and the force experienced by the plate. For large aspect ratio plates, leading-edge vortices evolved into hairpin vortices that eventually detached from the plate, interacting with the tip vortices in a complex manner. Separation of the leading-edge vortex is delayed to some extent by having convective transport of the spanwise vorticity as observed in flow over elliptic, semicircular, and delta-shaped planforms. The time at which lift achieves its maximum is observed to be fairly constant over dierent aspect ratios, angles of attack, and planform geometries during the initial transient. Preliminary results are also presented for flow over plates with steady actuation near the leading edge.

Turning dynamics and passive damping in flapping flight

We investigated whether flapping flight has an inherent stability by analyzing the inertial and aerodynamic effects of flapping wings on body dynamics. Based on wing and body kinematics of free flying fruit flies during rapid maneuvers, we found a passive counter torque due to body rotation. It is indentified both in simulation through quasi-steady state aerodynamic model and through experiments on a dynamically scaled robotic wing. An analytical form is derived correspondingly. In the turning yaw axis, the estimated damping coefficient of flapping wings is significantly higher than body frictional damping; this indicates a passive deceleration during turning. By simulating insect to rotate about each principal axis of inertial and body frames, we calculated the corresponding damping coefficients, and further analyzed the attitude stability. The result reveals that, passive damping of flapping flight, while does not necessarily lead to a stable full body dynamics, provides a considerable passive restoring torque that could be critical for flight stabilization and control in the design of micro aerial vehicles. Preliminary analysis on the scaling parameters of passive damping was also performed.

Prior Mating Experience Modulates the Dispersal of Drosophila in Males More Than in Females

Cues from both an animal’s internal physiological state and its local environment may influence its decision to disperse. However, identifying and quantifying the causative factors underlying the initiation of dispersal is difficult in uncontrolled natural settings. In this study, we automatically monitored the movement of fruit flies and examined the influence of food availability, sex, and reproductive status on their dispersal between laboratory environments. In general, flies with mating experience behave as if they are hungrier than virgin flies, leaving at a greater rate when food is unavailable and staying longer when it is available. Males dispersed at a higher rate and were more active than females when food was unavailable, but tended to stay longer in environments containing food than did females. We found no significant relationship between weight and activity, suggesting the behavioral differences between males and females are caused by an intrinsic factor relating to the sex of a fly and not simply its body size. Finally, we observed a significant difference between the dispersal of the natural isolate used throughout this study and the widely-used laboratory strain, Canton-S, and show that the difference cannot be explained by allelic differences in the foraging gene.