Simon M. Walker

Details of Insect Wing Design and Deformation Enhance Aerodynamic Function and Flight Efficiency

Locust Wing Aerodynamics Insect wings function as deformable aerofoils, but the precise aerodynamic benefits of the observed deformations have remained obscure. Previous models have treated the wing as a flat plate, lacking any deformation, even though it is clear that the locust wing can twist and rotate along its length. Young et al. (p. 1549) validate a computational fluid dynamic model, using particle imaging velocimetry and smoke visualization of the flow around actual locusts, and use the model to investigate the effect of measured changes in wing shape during a stroke cycle. The complexity of insect wing venation directly affects the aerodynamics of flight via the intermediary of wing deformation. Measurements of locust wing kinematics validate a fluid dynamics model of the aerodynamic effects of wing deformation. Insect wings are complex structures that deform dramatically in flight. We analyzed the aerodynamic consequences of wing deformation in locusts using a three-dimensional computational fluid dynamics simulation based on detailed wing kinematics. We validated the simulation against smoke visualizations and digital particle image velocimetry on real locusts. We then used the validated model to explore the effects of wing topography and deformation, first by removing camber while keeping the same time-varying twist distribution, and second by removing camber and spanwise twist. The full-fidelity model achieved greater power economy than the uncambered model, which performed better than the untwisted model, showing that the details of insect wing topography and deformation are important aerodynamically. Such details are likely to be important in engineering applications of flapping flight.

Deformable wing kinematics in free-flying hoverflies

Here, we present a detailed analysis of the deforming wing kinematics of free-flying hoverflies (Eristalis tenax, Linnaeus) during hovering flight. We used four high-speed digital video cameras to reconstruct the motion of approximately 22 points on each wing using photogrammetric techniques. While the root-flapping motion of the wing is similar in both the downstroke and upstroke, and is well modelled as a simple harmonic motion, other wing kinematic parameters show substantial variation between the downstroke and upstroke. Whereas the magnitude of the angle of incidence varies considerably within and between different hoverflies, the twist distribution along the wing is highly stereotyped. The angle of incidence and camber both show a recoil effect as they change abruptly at stroke reversal. Pronation occurs consistently after stroke reversal, which is perhaps surprising, because this has been found to reduce lift production in modelling studies. We find that the alula, a hinged flap near the base of the wing, operates in two discrete states: either in plane with the wing, or flipped approximately normal to it. We hypothesize that the alula may be acting as a flow-control device.

Photogrammetric reconstruction of high-resolution surface topographies and deformable wing kinematics of tethered locusts and free-flying hoverflies

Here, we present a suite of photogrammetric methods for reconstructing insect wing kinematics, to provide instantaneous topographic maps of the wing surface. We filmed tethered locusts (Schistocerca gregaria) and free-flying hoverflies (Eristalis tenax) using four high-speed digital video cameras. We digitized multiple natural features and marked points on the wings using manual and automated tracking. Epipolar geometry was used to identify additional points on the hoverfly wing outline which were anatomically indistinguishable. The cameras were calibrated using a bundle adjustment technique that provides an estimate of the error associated with each individual data point. The mean absolute three-dimensional measurement error was 0.11 mm for the locust and 0.03 mm for the hoverfly. The error in the angle of incidence was at worst 0.51° (s.d.) for the locust and 0.88° (s.d.) for the hoverfly. The results we present are of unprecedented spatio-temporal resolution, and represent the most detailed measurements of insect wing kinematics to date. Variable spanwise twist and camber are prominent in the wingbeats of both the species, and are of such complexity that they would not be adequately captured by lower resolution techniques. The role of spanwise twist and camber in insect flight has yet to be fully understood, and accurate insect wing kinematics such as we present here are required to be sure of making valid predictions about their aerodynamic effects.

Deformable wing kinematics in the desert locust: how and why do camber, twist and topography vary through the stroke?

Here, we present a detailed analysis of the wing kinematics and wing deformations of desert locusts (Schistocerca gregaria, Forskål) flying tethered in a wind tunnel. We filmed them using four high-speed digital video cameras, and used photogrammetry to reconstruct the motion of more than 100 identified points. Whereas the hindwing motions were highly stereotyped, the forewing motions showed considerable variation, consistent with a role in flight control. Both wings were positively cambered on the downstroke. The hindwing was cambered through an ‘umbrella effect’ whereby the trailing edge tension compressed the radial veins during the downstroke. Hindwing camber was reversed on the upstroke as the wing fan corrugated, reducing the projected area by 30 per cent, and releasing the tension in the trailing edge. Both the wings were strongly twisted from the root to the tip. The linear decrease in incidence along the hindwing on the downstroke precisely counteracts the linear increase in the angle of attack that would otherwise occur in root flapping for an untwisted wing. The consequent near-constant angle of attack is reminiscent of the optimum for a propeller of constant aerofoil section, wherein a linear twist distribution allows each section to operate at the unique angle of attack maximizing the lift to drag ratio. This implies tuning of the structural, morphological and kinematic parameters of the hindwing for efficient aerodynamic force production.

In Vivo Time- Resolved Microtomography Reveals the Mechanics of the Blowfly Flight Motor

Time-resolved X-ray microtomography permits a real-time view of the blowfly in flight at a previously unprecedented level of detail, revealing how the tiny steering muscles work.

Morphomechanical Innovation Drives Explosive Seed Dispersal

Summary How mechanical and biological processes are coordinated across cells, tissues, and organs to produce complex traits is a key question in biology. Cardamine hirsuta, a relative of Arabidopsis thaliana, uses an explosive mechanism to disperse its seeds. We show that this trait evolved through morphomechanical innovations at different spatial scales. At the organ scale, tension within the fruit wall generates the elastic energy required for explosion. This tension is produced by differential contraction of fruit wall tissues through an active mechanism involving turgor pressure, cell geometry, and wall properties of the epidermis. Explosive release of this tension is controlled at the cellular scale by asymmetric lignin deposition within endocarp b cells—a striking pattern that is strictly associated with explosive pod shatter across the Brassicaceae plant family. By bridging these different scales, we present an integrated mechanism for explosive seed dispersal that links evolutionary novelty with complex trait innovation. Video Abstract

Four-dimensional in vivo X-ray microscopy with projection-guided gating

Visualizing fast micrometer scale internal movements of small animals is a key challenge for functional anatomy, physiology and biomechanics. We combine phase contrast tomographic microscopy (down to 3.3 μm voxel size) with retrospective, projection-based gating (in the order of hundreds of microseconds) to improve the spatiotemporal resolution by an order of magnitude over previous studies. We demonstrate our method by visualizing 20 three-dimensional snapshots through the 150 Hz oscillations of the blowfly flight motor.

Smart wing rotation and trailing-edge vortices enable high frequency mosquito flight

Mosquitoes exhibit unusual wing kinematics; their long, slender wings flap at remarkably high frequencies for their size (>800 Hz)and with lower stroke amplitudes than any other insect group. This shifts weight support away from the translation-dominated, aerodynamic mechanisms used by most insects, as well as by helicopters and aeroplanes, towards poorly understood rotational mechanisms that occur when pitching at the end of each half-stroke. Here we report free-flight mosquito wing kinematics, solve the full Navier–Stokes equations using computational fluid dynamics with overset grids, and validate our results with in vivo flow measurements. We show that, although mosquitoes use familiar separated flow patterns, much of the aerodynamic force that supports their weight is generated in a manner unlike any previously described for a flying animal. There are three key features: leading-edge vortices (a well-known mechanism that appears to be almost ubiquitous in insect flight), trailing-edge vortices caused by a form of wake capture at stroke reversal, and rotational drag. The two new elements are largely independent of the wing velocity, instead relying on rapid changes in the pitch angle (wing rotation) at the end of each half-stroke, and they are therefore relatively immune to the shallow flapping amplitude. Moreover, these mechanisms are particularly well suited to high aspect ratio mosquito wings.

Aerodynamics of aerofoil sections measured on a free-flying bird:

Abstract Birds are adapted to a wide range of flight conditions, from steady fixed-wing glides to high angle of attack manoeuvres involving unsteady separated flows. They naturally control and exploit the transitional Reynolds number regime of Re≈ 105 that is currently of interest in unmanned air vehicle technologies. This article presents a reconstruction of the inner portion of a wing of an eagle in free flight, during a rapid pitch-up manoeuvre at the end of a shallow glide to an elevated perch. Photogrammetric techniques were used to map the identified points on the wing and these were used to fit a mathematical model of the upper and lower surface topography using polynomial regression techniques. The surface model accounts for spanwise twist, spanwise bending, and varying chord distribution, as well as for the shape of the aerofoil. The aerodynamics of the two-dimensional aerofoil sections were analysed using XFOIL and were compared against two technical aerofoils, namely the Selig S1223 and Clark Y aerofoils, at 1×105≤Re≤2×105. The bird aerofoil maintains a robust, near-constant drag coefficient over a wide lift coefficient range.

Mechanics and aerodynamics of perching manoeuvres in a large bird of prey

This paper reviews recent results on the mechanics and aerodynamics of perching in a large bird of prey, the Steppe Eagle Aquila nipalensis . Data collected using onboard and high-speed video cameras are used to examine gross morphing of the wing planform by the flight muscles, and smaller-scale morphing of the wing profile by aeroelastic deflection of the feathers, Carruthers et al . High-resolution still images are used to reconstruct the shape of the wing using multi-station photogrammetry, and the performance of the measured wing profile is analysed using a panel code, Carruthers et al . In bringing these lines of research together, we examine the role of aeroelastic feather deflection, and show that the key to perching in birds lies not in high-lift aerodynamics, but in the way in which the wings and tail morph to allow the bird to transition quickly from a steady glide into a deep stall.