How swifts control their glide performance with morphing wings.

Gliding birds continually change the shape and size of their wings, presumably to exploit the profound effect of wing morphology on aerodynamic performance. That birds should adjust wing sweep to suit glide speed has been predicted qualitatively by analytical glide models, which extrapolated the wing's performance envelope from aerodynamic theory. Here we describe the aerodynamic and structural performance of actual swift wings, as measured in a wind tunnel, and on this basis build a semi-empirical glide model. By measuring inside and outside swifts' behavioural envelope, we show that choosing the most suitable sweep can halve sink speed or triple turning rate. Extended wings are superior for slow glides and turns; swept wings are superior for fast glides and turns. This superiority is due to better aerodynamic performance-with the exception of fast turns. Swept wings are less effective at generating lift while turning at high speeds, but can bear the extreme loads. Finally, our glide model predicts that cost-effective gliding occurs at speeds of 8-10 m s(-1), whereas agility-related figures of merit peak at 15-25 m s(-1). In fact, swifts spend the night ('roost') in flight at 8-10 m s(-1) (ref. 11), thus our model can explain this choice for a resting behaviour. Morphing not only adjusts birds' wing performance to the task at hand, but could also control the flight of future aircraft.

Aerodynamics of gliding flight in common swifts.

Gliding flight performance and wake topology of a common swift (Apus apus L.) were examined in a wind tunnel at speeds between 7 and 11 m s(-1). The tunnel was tilted to simulate descending flight at different sink speeds. The swift varied its wingspan, wing area and tail span over the speed range. Wingspan decreased linearly with speed, whereas tail span decreased in a nonlinear manner. For each airspeed, the minimum glide angle was found. The corresponding sink speeds showed a curvilinear relationship with airspeed, with a minimum sink speed at 8.1 m s(-1) and a speed of best glide at 9.4 m s(-1). Lift-to-drag ratio was calculated for each airspeed and tilt angle combinations and the maximum for each speed showed a curvilinear relationship with airspeed, with a maximum of 12.5 at an airspeed of 9.5 m s(-1). Wake was sampled in the transverse plane using stereo digital particle image velocimetry (DPIV). The main structures of the wake were a pair of trailing wingtip vortices and a pair of trailing tail vortices. Circulation of these was measured and a model was constructed that showed good weight support. Parasite drag was estimated from the wake defect measured in the wake behind the body. Parasite drag coefficient ranged from 0.30 to 0.22 over the range of airspeeds. Induced drag was calculated and used to estimate profile drag coefficient, which was found to be in the same range as that previously measured on a Harris' hawk.

Downstroke and upstroke conflict during banked turns in butterflies.

For all flyers, aeroplanes or animals, making banked turns involve a rolling motion which, due to higher induced drag on the outer than the inner wing, results in a yawing torque opposite to the turn. This adverse yaw torque can be counteracted using a tail, but how animals that lack tail, e.g. all insects, handle this problem is not fully understood. Here, we quantify the performance of turning take-off flights in butterflies and find that they use force vectoring during banked turns without fully compensating for adverse yaw. This lowers their turning performance, increasing turn radius, since thrust becomes misaligned with the flight path. The separation of function between downstroke (lift production) and upstroke (thrust production) in our butterflies, in combination with a more pronounced adverse yaw during the upstroke increases the misalignment of the thrust. This may be a cost the butterflies pay for the efficient thrust-generating upstroke clap, but also other insects fail to rectify adverse yaw during escape manoeuvres, suggesting a general feature in functionally two-winged insect flight. When lacking tail and left with costly approaches to counteract adverse yaw, costs of flying with adverse yaw may be outweighed by the benefits of maintaining thrust and flight speed.

Butterflies fly using efficient propulsive clap mechanism owing to flexible wings.

Butterflies look like no other flying animal, with unusually short, broad and large wings relative to their body size. Previous studies have suggested butterflies use several unsteady aerodynamic mechanisms to boost force production with upstroke wing clap being a prominent feature. When the wings clap together at the end of upstroke the air between the wings is pressed out, creating a jet, pushing the animal in the opposite direction. Although viewed, for the last 50 years, as a crucial mechanism in insect flight, quantitative aerodynamic measurements of the clap in freely flying animals are lacking. Using quantitative flow measurements behind freely flying butterflies during take-off and a mechanical clapper, we provide aerodynamic performance estimates for the wing clap. We show that flexible butterfly wings, forming a cupped shape during the upstroke and clap, thrust the butterfly forwards, while the downstroke is used for weight support. We further show that flexible wings dramatically increase the useful impulse (+22%) and efficiency (+28%) of the clap compared to rigid wings. Combined, our results suggest butterflies evolved a highly effective clap, which provides a mechanistic hypothesis for their unique wing morphology. Furthermore, our findings could aid the design of man-made flapping drones, boosting propulsive performance.

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected.

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.

See-saw rocking: an in vitro model for mechanotransduction research.

In vitro mechanotransduction studies, uncovering the basic science of the response of cells to mechanical forces, are essential for progress in tissue engineering and its clinical application. Many varying investigations have described a multitude of cell responses; however, as the precise nature and magnitude of the stresses applied are infrequently reported and rarely validated, the experiments are often not comparable, limiting research progress. This paper provides physical and biological validation of a widely available fluid stimulation device, a see-saw rocker, as an in vitro model for cyclic fluid shear stress mechanotransduction. This allows linkage between precisely characterized stimuli and cell monolayer response in a convenient six-well plate format. Models of one well were discretized and analysed extensively using computational fluid dynamics to generate convergent, stable and consistent predictions of the cyclic fluid velocity vectors at a rocking frequency of 0.5 Hz, accounting for the free surface. Validation was provided by comparison with flow velocities measured experimentally using particle image velocimetry. Qualitative flow behaviour was matched and quantitative analysis showed agreement at representative locations and time points. Maximum shear stress of 0.22 Pa was estimated near the well edge, and time-average shear stress ranged between 0.029 and 0.068 Pa. Human tenocytes stimulated using the system showed significant increases in collagen and GAG secretion at 2 and 7 day time points. This in vitro model for mechanotransduction provides a versatile, flexible and inexpensive method for the fluid shear stress impact on biological cells to be studied.

Time-resolved vortex wake of a common swift flying over a range of flight speeds.

The wake of a freely flying common swift (Apus apus L.) is examined in a wind tunnel at three different flight speeds, 5.7, 7.7 and 9.9 m s(-1). The wake of the bird is visualized using high-speed stereo digital particle image velocimetry (DPIV). Wake images are recorded in the transverse plane, perpendicular to the airflow. The wake of a swift has been studied previously using DPIV and recording wake images in the longitudinal plane, parallel to the airflow. The high-speed DPIV system allows for time-resolved wake sampling and the result shows features that were not discovered in the previous study, but there was approximately a 40 per cent vertical force deficit. As the earlier study also revealed, a pair of wingtip vortices are trailing behind the wingtips, but in addition, a pair of tail vortices and a pair of 'wing root vortices' are found that appear to originate from the wing/body junction. The existence of wing root vortices suggests that the two wings are not acting as a single wing, but are to some extent aerodynamically detached from each other. It is proposed that this is due to the body disrupting the lift distribution over the wing by generating less lift than the wings.

Vortex wake and flight kinematics of a swift in cruising flight in a wind tunnel.

In this paper we describe the flight characteristics of a swift (Apus apus) in cruising flight at three different flight speeds (8.0, 8.4 and 9.2 m s(-1)) in a low turbulence wind tunnel. The wingbeat kinematics were recorded by high-speed filming and the wake of the bird was visualized by digital particle image velocimetry (DPIV). Certain flight characteristics of the swift differ from those of previously studied species. As the flight speed increases, the angular velocity of the wingbeat remains constant, and so as the wingbeat amplitude increases, the frequency decreases accordingly, as though the flight muscles were contracting at a fixed rate. The wings are also comparatively inflexible and are flexed or retracted rather little during the upstroke. The upstroke is always aerodynamically active and this is reflected in the wake, where shedding of spanwise vorticity occurs throughout the wingbeat. Although the wake superficially resembles those of other birds in cruising flight, with a pair of trailing wingtip vortices connected by spanwise vortices, the continuous shedding of first positive vorticity during the downstroke and then negative vorticity during the upstroke suggests a wing whose circulation is gradually increasing and then decreasing during the wingbeat cycle. The wake (and implied wing aerodynamics) are not well described by discrete vortex loop models, but a new wake-based model, where incremental spanwise and streamwise variations of the wake impulse are integrated over the wingbeat, shows good agreement of the vertical momentum flux with the required weight support. The total drag was also estimated from the wake alone, and the calculated lift:drag ratio of approximately 13 for flapping flight is the highest measured yet for birds.