Dynamics of hinged wings in strong upward gusts.

A bird's wings are articulated to its body via highly mobile shoulder joints. The joints confer an impressive range of motion, enabling the wings to make broad, sweeping movements that can modulate quite dramatically the production of aerodynamic load. This is enormously useful in challenging flight environments, especially the gusty, turbulent layers of the lower atmosphere. In this study, we develop a dynamics model to examine how a bird-scale gliding aircraft can use wing-root hinges (analogous to avian shoulder joints) to reject the initial impact of a strong upward gust. The idea requires that the spanwise centre of pressure and the centre of percussion of the hinged wing start, and stay, in good initial alignment (the centre of percussion here is related to the idea of a 'sweet spot' on a bat, as in cricket or baseball). We propose a method for achieving this rejection passively, for which the essential ingredients are (i) appropriate lift and mass distributions; (ii) hinges under constant initial torque; and (iii) a wing whose sections stall softly. When configured correctly, the gusted wings will first pivot on their hinges without disturbing the fuselage of the aircraft, affording time for other corrective actions to engage. We expect this system to enhance the control of aircraft that fly in gusty conditions.

A method for continuous study of soaring and windhovering birds.

Avian flight continues to inspire aircraft designers. Reducing the scale of autonomous aircraft to that of birds and large insects has resulted in new control challenges when attempting to hold steady flight in turbulent atmospheric wind. Some birds, however, are capable of remarkably stable hovering flight in the same conditions. This work describes the development of a wind tunnel configuration that facilitates the study of flapless windhovering (hanging) and soaring bird flight in wind conditions replicating those in nature. Updrafts were generated by flow over replica "hills" and turbulence was introduced through upstream grids, which had already been developed to replicate atmospheric turbulence in prior studies. Successful flight tests with windhovering nankeen kestrels (Falco cenchroides) were conducted, verifying that the facility can support soaring and wind hovering bird flight. The wind tunnel allows the flow characteristics to be carefully controlled and measured, providing great advantages over outdoor flight tests. Also, existing wind tunnels may be readily configured using this method, providing a simpler alternative to the development of dedicated bird flight wind tunnels such as tilting wind tunnels, and the large test section allows for the replication of orographic soaring. This methodology holds promise for future testing investigating the flight behaviour and control responses employed by soaring and windhovering birds.

Quantifying avian inertial properties using calibrated computed tomography.

Estimating centre of mass and mass moments of inertia is an important aspect of many studies in biomechanics. Characterising these parameters accurately in three dimensions is challenging with traditional methods requiring dissection or suspension of cadavers. Here, we present a method to quantify the three-dimensional centre of mass and inertia tensor of birds of prey using calibrated computed tomography (CT) scans. The technique was validated using several independent methods, providing body segment mass estimates within approximately 1% of physical dissection measurements and moment of inertia measurements with a 0.993 R2 correlation with conventional trifilar pendulum measurements. Calibrated CT offers a relatively straightforward, non-destructive approach that yields highly detailed mass distribution data that can be used for three-dimensional dynamics modelling in biomechanics. Although demonstrated here with birds, this approach should work equally well with any animal or appendage capable of being CT scanned.

Raptor wing morphing with flight speed.

In gliding flight, birds morph their wings and tails to control their flight trajectory and speed. Using high-resolution videogrammetry, we reconstructed accurate and detailed three-dimensional geometries of gliding flights for three raptors (barn owl, Tyto alba; tawny owl, Strix aluco, and goshawk, Accipiter gentilis). Wing shapes were highly repeatable and shoulder actuation was a key component of reconfiguring the overall planform and controlling angle of attack. The three birds shared common spanwise patterns of wing twist, an inverse relationship between twist and peak camber, and held their wings depressed below their shoulder in an anhedral configuration. With increased speed, all three birds tended to reduce camber throughout the wing, and their wings bent in a saddle-shape pattern. A number of morphing features suggest that the coordinated movements of the wing and tail support efficient flight, and that the tail may act to modulate wing camber through indirect aeroelastic control.

Bird wings act as a suspension system that rejects gusts.

Musculoskeletal systems cope with many environmental perturbations without neurological control. These passive preflex responses aid animals to move swiftly through complex terrain. Whether preflexes play a substantial role in animal flight is uncertain. We investigated how birds cope with gusty environments and found that their wings can act as a suspension system, reducing the effects of vertical gusts by elevating rapidly about the shoulder. This preflex mechanism rejected the gust impulse through inertial effects, diminishing the predicted impulse to the torso and head by 32% over the first 80 ms, before aerodynamic mechanisms took effect. For each wing, the centre of aerodynamic loading aligns with the centre of percussion, consistent with enhancing passive inertial gust rejection. The reduced motion of the torso in demanding conditions simplifies crucial tasks, such as landing, prey capture and visual tracking. Implementing a similar preflex mechanism in future small-scale aircraft will help to mitigate the effects of gusts and turbulence without added computational burden.

High aerodynamic lift from the tail reduces drag in gliding raptors.

Many functions have been postulated for the aerodynamic role of the avian tail during steady-state flight. By analogy with conventional aircraft, the tail might provide passive pitch stability if it produced very low or negative lift. Alternatively, aeronautical principles might suggest strategies that allow the tail to reduce inviscid, induced drag: if the wings and tail act in different horizontal planes, they might benefit from biplane-like aerodynamics; if they act in the same plane, lift from the tail might compensate for lift lost over the fuselage (body), reducing induced drag with a more even downwash profile. However, textbook aeronautical principles should be applied with caution because birds have highly capable sensing and active control, presumably reducing the demand for passive aerodynamic stability, and, because of their small size and low flight speeds, operate at Reynolds numbers two orders of magnitude below those of light aircraft. Here, by tracking up to 20,000, 0.3 mm neutrally buoyant soap bubbles behind a gliding barn owl, tawny owl and goshawk, we found that downwash velocity due to the body/tail consistently exceeds that due to the wings. The downwash measured behind the centreline is quantitatively consistent with an alternative hypothesis: that of constant lift production per planform area, a requirement for minimizing viscous, profile drag. Gliding raptors use lift distributions that compromise both inviscid induced drag minimization and static pitch stability, instead adopting a strategy that reduces the viscous drag, which is of proportionately greater importance to lower Reynolds number fliers.

Avian surface reconstruction in free flight with application to flight stability analysis of a barn owl and peregrine falcon.

Birds primarily create and control the forces necessary for flight through changing the shape and orientation of their wings and tail. Their wing geometry is characterised by complex variation in parameters such as camber, twist, sweep and dihedral. To characterise this complexity, a multi-view stereo-photogrammetry setup was developed for accurately measuring surface geometry in high resolution during free flight. The natural patterning of the birds was used as the basis for phase correlation-based image matching, allowing indoor or outdoor use while being non-intrusive for the birds. The accuracy of the method was quantified and shown to be sufficient for characterising the geometric parameters of interest, but with a reduction in accuracy close to the wing edge and in some localised regions. To demonstrate the method's utility, surface reconstructions are presented for a barn owl (Tyto alba) and peregrine falcon (Falco peregrinus) during three instants of gliding flight per bird. The barn owl flew with a consistent geometry, with positive wing camber and longitudinal anhedral. Based on flight dynamics theory, this suggests it was longitudinally statically unstable during these flights. The peregrine falcon flew with a consistent glide angle, but at a range of air speeds with varying geometry. Unlike the barn owl, its glide configuration did not provide a clear indication of longitudinal static stability/instability. Aspects of the geometries adopted by both birds appeared to be related to control corrections and this method would be well suited for future investigations in this area, as well as for other quantitative studies into avian flight dynamics.

Head movements quadruple the range of speeds encoded by the insect motion vision system in hawkmoths.

Flying insects use compensatory head movements to stabilize gaze. Like other optokinetic responses, these movements can reduce image displacement, motion and misalignment, and simplify the optic flow field. Because gaze is imperfectly stabilized in insects, we hypothesized that compensatory head movements serve to extend the range of velocities of self-motion that the visual system encodes. We tested this by measuring head movements in hawkmoths Hyles lineata responding to full-field visual stimuli of differing oscillation amplitudes, oscillation frequencies and spatial frequencies. We used frequency-domain system identification techniques to characterize the head's roll response, and simulated how this would have affected the output of the motion vision system, modelled as a computational array of Reichardt detectors. The moths' head movements were modulated to allow encoding of both fast and slow self-motion, effectively quadrupling the working range of the visual system for flight control. By using its own output to drive compensatory head movements, the motion vision system thereby works as an adaptive sensor, which will be especially beneficial in nocturnal species with inherently slow vision. Studies of the ecology of motion vision must therefore consider the tuning of motion-sensitive interneurons in the context of the closed-loop systems in which they function.

Fine-scale flight strategies of gulls in urban airflows indicate risk and reward in city living.

Birds modulate their flight paths in relation to regional and global airflows in order to reduce their travel costs. Birds should also respond to fine-scale airflows, although the incidence and value of this remains largely unknown. We resolved the three-dimensional trajectories of gulls flying along a built-up coastline, and used computational fluid dynamic models to examine how gulls reacted to airflows around buildings. Birds systematically altered their flight trajectories with wind conditions to exploit updraughts over features as small as a row of low-rise buildings. This provides the first evidence that human activities can change patterns of space-use in flying birds by altering the profitability of the airscape. At finer scales still, gulls varied their position to select a narrow range of updraught values, rather than exploiting the strongest updraughts available, and their precise positions were consistent with a strategy to increase their velocity control in gusty conditions. Ultimately, strategies such as these could help unmanned aerial vehicles negotiate complex airflows. Overall, airflows around fine-scale features have profound implications for flight control and energy use, and consideration of this could lead to a paradigm-shift in the way ecologists view the urban environment.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

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.

The flow fields involved in hydrodynamic imaging by blind Mexican cave fish (Astyanax fasciatus). Part I: open water and heading towards a wall.

Blind Mexican cave fish (Astyanax fasciatus) sense the presence of nearby objects by sensing changes in the water flow around their body. The information available to the fish using this hydrodynamic imaging ability depends on the properties of the flow field it generates while gliding and how this flow field is altered by the presence of objects. Here, we used particle image velocimetry to measure the flow fields around gliding blind cave fish as they moved through open water and when heading towards a wall. These measurements, combined with computational fluid dynamics models, were used to estimate the stimulus to the lateral line system of the fish. Our results showed that there was a high-pressure region around the nose of the fish, low-pressure regions corresponding to accelerated flow around the widest part of the body and a thick laminar boundary layer down the body. When approaching a wall head-on, the changes in the stimulus to the lateral line were confined to approximately the first 20% of the body. Assuming that the fish are sensitive to a certain relative change in lateral line stimuli, it was found that swimming at higher Reynolds numbers slightly decreased the distance at which the fish could detect a wall when approaching head-on, which is the opposite to what has previously been expected. However, when the effects of environmental noise are considered, swimming at higher speed may improve the signal to noise ratio of the stimulus to the lateral line.

The flow fields involved in hydrodynamic imaging by blind Mexican cave fish (Astyanax fasciatus). Part II: gliding parallel to a wall.

Blind Mexican cave fish (Astyanax fasciatus) are able to sense detailed information about objects by gliding alongside them and sensing changes in the flow field around their body using their lateral line sensory system. Hence the fish are able to build hydrodynamic images of their surroundings. This study measured the flow fields around blind cave fish using particle image velocimetry (PIV) as they swam parallel to a wall. Computational fluid dynamics models were also used to calculate the flow fields and the stimuli to the lateral line sensory system. Our results showed that characteristic changes in the form of the flow field occurred when the fish were within approximately 0.20 body lengths (BL) of a wall. The magnitude of these changes increased steadily as the distance between the fish and the wall was reduced. When the fish were within 0.02 BL of the wall there was a change in the form of the flow field owing to the merging of the boundary layers on the body of the fish and the wall. The stimuli to the lateral line appears to be sufficient for fish to detect walls when they are 0.10 BL away (the mean distance at which they normally swim from a wall), but insufficient for the fish to detect a wall when 0.25 BL away. This suggests that the nature of the flow fields surrounding the fish are such that hydrodynamic imaging can only be used by fish to detect surfaces at short range.

The influence of viscous hydrodynamics on the fish lateral-line system.

Fish exhibit many behaviors that involve sensing water flows with their lateral-line system. In many situations, viscosity affects how the flow interacts with the body of the fish and the neuromasts of the lateral line. Here we discuss how viscosity influences the stimulus to the fish lateral-line system. The movement of a fish's body creates flows that can interfere with the detection of external signals, but these flows can also serve as a source of information about nearby obstacles and the fish's own hydrodynamic performance. The viscous boundary layer on the surface of the skin alters external signals by attenuating the low-frequency components of stimuli. The stimulus to each neuromast depends on the interaction of the fluid surrounding the neuromast and the structural properties of that neuromast, including the number of mechanosensory hair cells it contains. A consideration of the influences of viscosity on flow, at both the whole-body and receptor levels, offers the promise of a more comprehensive understanding of the signals involved in behaviors mediated by the lateral-line system.

Swimming kinematics and hydrodynamic imaging in the blind Mexican cave fish (Astyanax fasciatus).

Blind Mexican cave fish (Astyanax fasciatus) lack a functioning visual system, and are known to use self-generated water motion to sense their surroundings; an ability termed hydrodynamic imaging. Nearby objects distort the flow field created by the motion of the fish. These flow distortions are sensed by the mechanosensory lateral line. Here we used image processing to measure detailed kinematics, along with a new behavioural technique, to investigate the effectiveness of hydrodynamic imaging. In a head-on approach to a wall, fish reacted to avoid collision with the wall at an average distance of only 4.0+/-0.2 mm. Contrary to previous expectation, there was no significant correlation between the swimming velocity of the fish and the distance at which they reacted to the wall. Hydrodynamic imaging appeared to be most effective when the fish were gliding with their bodies held straight, with the proportion of approaches to the wall that resulted in collision increasing from 11% to 73% if the fish were beating their tails rather than gliding as they neared the wall. The swimming kinematics of the fish were significantly different when swimming beside a wall compared with when swimming away from any walls. Blind cave fish frequently touched walls when swimming alongside them, indicating that they use both tactile and hydrodynamic information in this situation. We conclude that although hydrodynamic imaging can provide effective collision avoidance, it is a short-range sense that may often be used synergistically with direct touch.