Beyond robins: aerodynamic analyses of animal flight

Recent progress in studies of animal flight mechanics is reviewed. A range of birds, and now bats, has been studied in wind tunnel facilities, revealing an array of wake patterns caused by the beating wings and also by the drag on the body. Nevertheless, the quantitative analysis of these complex wake structures shows a degree of similarity among all the different wake patterns and a close agreement with standard quasi-steady aerodynamic models and predictions. At the same time, new data on the flow over a bat wing in mid-downstroke show that, at least in this case, such simplifications cannot be useful in describing in detail either the wing properties or control prospects. The reasons for these apparently divergent results are discussed and prospects for future advances are considered.     

Computational investigation of cicada aerodynamics in forward flight

Free forward flight of cicadas is investigated through high-speed photogrammetry, three-dimensional surface reconstruction and computational fluid dynamics simulations. We report two new vortices generated by the cicada''s wide body. One is the thorax-generated vortex, which helps the downwash flow, indicating a new phenomenon of lift enhancement. Another is the cicada posterior body vortex, which entangles with the vortex ring composed of wing tip, trailing edge and wing root vortices. Some other vortex features include: independently developed left- and right-hand side leading edge vortex (LEV), dual-core LEV structure at the mid-wing region and near-wake two-vortex-ring structure. In the cicada forward flight, approximately 79% of the total lift is generated during the downstroke. Cicada wings experience drag in the downstroke, and generate thrust during the upstroke. Energetics study shows that the cicada in free forward flight consumes much more power in the downstroke than in the upstroke, to provide enough lift to support the weight and to overcome drag to move forward.     

Bird migration flight altitudes studied by a network of operational weather radars

A fully automated method for the detection and quantification of bird migration was developed for operational C-band weather radar, measuring bird density, speed and direction as a function of altitude. These weather radar bird observations have been validated with data from a high-accuracy dedicated bird radar, which was stationed in the measurement volume of weather radar sites in The Netherlands, Belgium and France for a full migration season during autumn 2007 and spring 2008. We show that weather radar can extract near real-time bird density altitude profiles that closely correspond to the density profiles measured by dedicated bird radar. Doppler weather radar can thus be used as a reliable sensor for quantifying bird densities aloft in an operational setting, which—when extended to multiple radars—enables the mapping and continuous monitoring of bird migration flyways. By applying the automated method to a network of weather radars, we observed how mesoscale variability in weather conditions structured the timing and altitude profile of bird migration within single nights. Bird density altitude profiles were observed that consisted of multiple layers, which could be explained from the distinct wind conditions at different take-off sites. Consistently lower bird densities are recorded in The Netherlands compared with sites in France and eastern Belgium, which reveals some of the spatial extent of the dominant Scandinavian flyway over continental Europe.     

Tomographic particle image velocimetry of desert locust wakes: instantaneous volumes combine to reveal hidden vortex elements and rapid wake deformation

Aerodynamic structures generated by animals in flight are unstable and complex. Recent progress in quantitative flow visualization has advanced our understanding of animal aerodynamics, but measurements have hitherto been limited to flow velocities at a plane through the wake. We applied an emergent, high-speed, volumetric fluid imaging technique (tomographic particle image velocimetry) to examine segments of the wake of desert locusts, capturing fully three-dimensional instantaneous flow fields. We used those flow fields to characterize the aerodynamic footprint in unprecedented detail and revealed previously unseen wake elements that would have gone undetected by two-dimensional or stereo-imaging technology. Vortex iso-surface topographies show the spatio-temporal signature of aerodynamic force generation manifest in the wake of locusts, and expose the extent to which animal wakes can deform, potentially leading to unreliable calculations of lift and thrust when using conventional diagnostic methods. We discuss implications for experimental design and analysis as volumetric flow imaging becomes more widespread.     

Time-varying span efficiency through the wingbeat of desert locusts

The flight performance of animals depends greatly on the efficacy with which they generate aerodynamic forces. Accordingly, maximum range, load-lifting capacity and peak accelerations during manoeuvres are all constrained by the efficiency of momentum transfer to the wake. Here, we use high-speed particle image velocimetry (1 kHz) to record flow velocities in the near wake of desert locusts (Schistocerca gregaria, Forskål). We use the measured flow fields to calculate time-varying span efficiency throughout the wing stroke cycle. The locusts are found to operate at a maximum span efficiency of 79 per cent, typically at a plateau of about 60 per cent for the majority of the downstroke, but at lower values during the upstroke. Moreover, the calculated span efficiencies are highest when the largest lift forces are being generated (90% of the total lift is generated during the plateau of span efficiency) suggesting that the combination of wing kinematics and morphology in locust flight perform most efficiently when doing the most work.     

The aerodynamic cost of flight in the short-tailed fruit bat (Carollia perspicillata): comparing theory with measurement

Aerodynamic theory has long been used to predict the power required for animal flight, but widely used models contain many simplifications. It has been difficult to ascertain how closely biological reality matches model predictions, largely because of the technical challenges of accurately measuring the power expended when an animal flies. We designed a study to measure flight speed-dependent aerodynamic power directly from the kinetic energy contained in the wake of bats flying in a wind tunnel. We compared these measurements with two theoretical predictions that have been used for several decades in diverse fields of vertebrate biology and to metabolic measurements from a previous study using the same individuals. A high-accuracy displaced laser sheet stereo particle image velocimetry experimental design measured the wake velocities in the Trefftz plane behind four bats flying over a range of speeds (3–7 m s⁻¹). We computed the aerodynamic power contained in the wake using a novel interpolation method and compared these results with the power predicted by Pennycuick''s and Rayner''s models. The measured aerodynamic power falls between the two theoretical predictions, demonstrating that the models effectively predict the appropriate range of flight power, but the models do not accurately predict minimum power or maximum range speeds. Mechanical efficiency—the ratio of aerodynamic power output to metabolic power input—varied from 5.9% to 9.8% for the same individuals, changing with flight speed.     

Comparative aerodynamic performance of flapping flight in two bat species using time-resolved wake visualization

Bats are unique among extant actively flying animals in having very flexible wings, controlled by multi-jointed fingers. This gives the potential for fine-tuned active control to optimize aerodynamic performance throughout the wingbeat and thus a more efficient flight. But how bat wing performance scales with size, morphology and ecology is not yet known. Here, we present time-resolved fluid wake data of two species of bats flying freely across a range of flight speeds using stereoscopic digital particle image velocimetry in a wind tunnel. From these data, we construct an average wake for each bat species and speed combination, which is used to estimate the flight forces throughout the wingbeat and resulting flight performance properties such as lift-to-drag ratio (L/D). The results show that the wake dynamics and flight performance of both bat species are similar, as was expected since both species operate at similar Reynolds numbers (Re) and Strouhal numbers (St). However, maximum L/D is achieved at a significant higher flight speed for the larger, highly mobile and migratory bat species than for the smaller non-migratory species. Although the flight performance of these bats may depend on a range of morphological and ecological factors, the differences in optimal flight speeds between the species could at least partly be explained by differences in their movement ecology.     

Predicting power-optimal kinematics of avian wings

A theoretical model of avian flight is developed which simulates wing motion through a class of methods known as predictive simulation. This approach uses numerical optimization to predict power-optimal kinematics of avian wings in hover, cruise, climb and descent. The wing dynamics capture both aerodynamic and inertial loads. The model is used to simulate the flight of the pigeon, Columba livia, and the results are compared with previous experimental measurements. In cruise, the model unearths a vast range of kinematic modes that are capable of generating the required forces for flight. The most efficient mode uses a near-vertical stroke–plane and a flexed-wing upstroke, similar to kinematics recorded experimentally. In hover, the model predicts that the power-optimal mode uses an extended-wing upstroke, similar to hummingbirds. In flexing their wings, pigeons are predicted to consume 20% more power than if they kept their wings full extended, implying that the typical kinematics used by pigeons in hover are suboptimal. Predictions of climbing flight suggest that the most energy-efficient way to reach a given altitude is to climb as steeply as possible, subjected to the availability of power.     

Elytra boost lift,but reduce aerodynamic efficiency in flying beetles

Flying insects typically possess two pairs of wings. In beetles, the front pair has evolved into short, hardened structures, the elytra, which protect the second pair of wings and the abdomen. This allows beetles to exploit habitats that would otherwise cause damage to the wings and body. Many beetles fly with the elytra extended, suggesting that they influence aerodynamic performance, but little is known about their role in flight. Using quantitative measurements of the beetle''s wake, we show that the presence of the elytra increases vertical force production by approximately 40 per cent, indicating that they contribute to weight support. The wing-elytra combination creates a complex wake compared with previously studied animal wakes. At mid-downstroke, multiple vortices are visible behind each wing. These include a wingtip and an elytron vortex with the same sense of rotation, a body vortex and an additional vortex of the opposite sense of rotation. This latter vortex reflects a negative interaction between the wing and the elytron, resulting in a single wing span efficiency of approximately 0.77 at mid downstroke. This is lower than that found in birds and bats, suggesting that the extra weight support of the elytra comes at the price of reduced efficiency.     

A quantitative comparison of bird and bat wakes

Qualitative comparison of bird and bat wakes has demonstrated significant differences in the structure of the far wake. Birds have been found to have a unified vortex wake of the two wings, while bats have a more complex wake with gradients in the circulation along the wingspan, and with each wing generating its own vortex structure. Here, we compare quantitative measures of the circulation in the far wake of three bird and one bat species. We find that bats have a significantly stronger normalized circulation of the start vortex than birds. We also find differences in how the circulation develops during the wingbeat as demonstrated by the ratio of the circulation of the dominant start vortex and the total circulation of the same sense. Birds show a more prominent change with changing flight speed and a relatively weaker start vortex at minimum power speed than bats. We also find that bats have a higher normalized wake loading based on the start vortex, indicating higher relative induced drag and therefore less efficient lift generation than birds. Our results thus indicate fundamental differences in the aerodynamics of bird and bat flight that will further our understanding of the evolution of vertebrate flight.     

Aerodynamic effects of flexibility in flapping wings

Recent work on the aerodynamics of flapping flight reveals fundamental differences in the mechanisms of aerodynamic force generation between fixed and flapping wings. When fixed wings translate at high angles of attack, they periodically generate and shed leading and trailing edge vortices as reflected in their fluctuating aerodynamic force traces and associated flow visualization. In contrast, wings flapping at high angles of attack generate stable leading edge vorticity, which persists throughout the duration of the stroke and enhances mean aerodynamic forces. Here, we show that aerodynamic forces can be controlled by altering the trailing edge flexibility of a flapping wing. We used a dynamically scaled mechanical model of flapping flight (Re ≈ 2000) to measure the aerodynamic forces on flapping wings of variable flexural stiffness (EI). For low to medium angles of attack, as flexibility of the wing increases, its ability to generate aerodynamic forces decreases monotonically but its lift-to-drag ratios remain approximately constant. The instantaneous force traces reveal no major differences in the underlying modes of force generation for flexible and rigid wings, but the magnitude of force, the angle of net force vector and centre of pressure all vary systematically with wing flexibility. Even a rudimentary framework of wing veins is sufficient to restore the ability of flexible wings to generate forces at near-rigid values. Thus, the magnitude of force generation can be controlled by modulating the trailing edge flexibility and thereby controlling the magnitude of the leading edge vorticity. To characterize this, we have generated a detailed database of aerodynamic forces as a function of several variables including material properties, kinematics, aerodynamic forces and centre of pressure, which can also be used to help validate computational models of aeroelastic flapping wings. These experiments will also be useful for wing design for small robotic insects and, to a limited extent, in understanding the aerodynamics of flapping insect wings.     

Development and Validation of a Novel Bird Strike Resistant Composite Leading Edge Structure

A novel design of a fibre-reinforced composite Leading Edge (LE) of a Horizontal Tail Plain (HTP) is proposed. The development and validation approach of the innovative composite LE structure are described. The main design goal is the satisfactory impact resistance of the novel composite LE in the case of bird strike. The design concept is based on the absorption of the major portion of the bird kinetic energy by the composite skins, in order to protect the ribs and the inner LE structure from damaging, thus preserving the tail plane functionality for safe landing. To this purpose, the LE skin is fabricated from specially designed composite panels, so called ‘tensor skin’ panels, comprising folded layers, which unfold under the impact load and increase the energy absorption capability of the LE. A numerical model simulating the bird strike process is developed and bird strike experimental testing is performed, in order to validate the proposed layout and prove the capability of the structure to successfully withstand the impact loading. The numerical modelling issues and the critical parameters of the simulation are discussed. The present work is part of the European Aeronautics Research Project, ‘Crashworthiness of aircraft for high velocity impact – CRAHVI’ [1].     

Active and passive stabilization of body pitch in insect flight

Flying insects have evolved sophisticated sensory–motor systems, and here we argue that such systems are used to keep upright against intrinsic flight instabilities. We describe a theory that predicts the instability growth rate in body pitch from flapping-wing aerodynamics and reveals two ways of achieving balanced flight: active control with sufficiently rapid reactions and passive stabilization with high body drag. By glueing magnets to fruit flies and perturbing their flight using magnetic impulses, we show that these insects employ active control that is indeed fast relative to the instability. Moreover, we find that fruit flies with their control sensors disabled can keep upright if high-drag fibres are also attached to their bodies, an observation consistent with our prediction for the passive stability condition. Finally, we extend this framework to unify the control strategies used by hovering animals and also furnish criteria for achieving pitch stability in flapping-wing robots.     

In search of the pitching momentum that enables some lizards to sustain bipedal running at constant speeds

The forelimbs of lizards are often lifted from the ground when they start sprinting. Previous research pointed out that this is a consequence of the propulsive forces from the hindlimbs. However, despite forward acceleration being hypothesized as necessary to lift the head, trunk and forelimbs, some species of agamids, teiids and basilisks sustain running in a bipedal posture at a constant speed for a relatively long time. Biomechanical modelling of steady bipedal running in the agamid Ctenophorus cristatus now shows that a combination of three mechanisms must be present to generate the angular impulse needed to cancel or oppose the effect of gravity. First, the trunk must be lifted significantly to displace the centre of mass more towards the hip joint. Second, the nose-up pitching moment resulting from aerodynamic forces exerted at the lizard''s surface must be taken into account. Third, the vertical ground-reaction forces at the hindlimb must show a certain degree of temporal asymmetry with higher forces closer to the instant of initial foot contact. Such asymmetrical vertical ground-reaction force profiles, which differ from the classical spring-mass model of bipedal running, seem inherent to the windmilling, splayed-legged running style of lizards.     

Vision-based flight control in the hawkmoth Hyles lineata

Vision is a key sensory modality for flying insects, playing an important role in guidance, navigation and control. Here, we use a virtual-reality flight simulator to measure the optomotor responses of the hawkmoth Hyles lineata, and use a published linear-time invariant model of the flight dynamics to interpret the function of the measured responses in flight stabilization and control. We recorded the forces and moments produced during oscillation of the visual field in roll, pitch and yaw, varying the temporal frequency, amplitude or spatial frequency of the stimulus. The moths’ responses were strongly dependent upon contrast frequency, as expected if the optomotor system uses correlation-type motion detectors to sense self-motion. The flight dynamics model predicts that roll angle feedback is needed to stabilize the lateral dynamics, and that a combination of pitch angle and pitch rate feedback is most effective in stabilizing the longitudinal dynamics. The moths’ responses to roll and pitch stimuli coincided qualitatively with these functional predictions. The moths produced coupled roll and yaw moments in response to yaw stimuli, which could help to reduce the energetic cost of correcting heading. Our results emphasize the close relationship between physics and physiology in the stabilization of insect flight.     

Lift enhancement by bats' dynamically changing wingspan

This paper elucidates the aerodynamic role of the dynamically changing wingspan in bat flight. Based on direct numerical simulations of the flow over a slow-flying bat, it is found that the dynamically changing wingspan can significantly enhance the lift. Further, an analysis of flow structures and lift decomposition reveal that the elevated vortex lift associated with the leading-edge vortices intensified by the dynamically changing wingspan considerably contributed to enhancement of the time-averaged lift. The nonlinear interaction between the dynamically changing wing and the vortical structures plays an important role in the lift enhancement of a flying bat in addition to the geometrical effect of changing the lifting-surface area in a flapping cycle. In addition, the dynamically changing wingspan leads to the higher efficiency in terms of generating lift for a given amount of the mechanical energy consumed in flight.