Per Henningsson

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

SUMMARY 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.

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.

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.

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.

Compensation for wind drift by migrating swifts

We investigated the orientation in relation to wind of common swifts, Apus apus, during nocturnal spring and autumn migration. Swifts are highly adapted to a life in the air, showing wind-dependent orientation during nocturnal roosting flights, and may be expected to be more efficient in their wind drift/compensation behaviour than nocturnal passerine migrants, which are usually subjected to full or partial wind drift. A tracking radar at Lund in southern Sweden was used to record the orientation of common swifts (identified by their characteristic radar echo signature) on nocturnal migration flights and to measure wind conditions at the altitudes at which the birds were flying. Comparing track and heading directions under easterly and westerly winds revealed that the swifts shifted their heading distinctly into the wind, with the result that track directions were similar under the different wind conditions. As this pattern of complete compensation for drift from cross-winds occurred during both spring and autumn migration, there were no indications of differences between age classes. In addition, we found an effect of side-winds on equivalent airspeeds, with swifts increasing their airspeed with increasing wind speed. Such a response has been theoretically predicted as part of an optimal behaviour for counteracting wind drift but has hereto not been empirically demonstrated. There was also a positive correlation between overall wind speed and equivalent airspeed, making it difficult to interpret whether the swifts respond to the total wind speed rather than specifically to the side-wind effect. Our results suggest that important differences may exist between species in their capacities to orient in relation to the wind and that the swift may be particularly efficient in adjusting heading direction and airspeed to obtain complete compensation for wind drift during high-altitude nocturnal migratory flights

Enhanced flight performance by genetic manipulation of wing shape in Drosophila

Insect wing shapes are remarkably diverse and the combination of shape and kinematics determines both aerial capabilities and power requirements. However, the contribution of any specific morphological feature to performance is not known. Using targeted RNA interference to modify wing shape far beyond the natural variation found within the population of a single species, we show a direct effect on flight performance that can be explained by physical modelling of the novel wing geometry. Our data show that altering the expression of a single gene can significantly enhance aerial agility and that the Drosophila wing shape is not, therefore, optimized for certain flight performance characteristics that are known to be important. Our technique points in a new direction for experiments on the evolution of performance specialities in animals.

Span efficiency in hawkmoths

Flight in animals is the result of aerodynamic forces generated as flight muscles drive the wings through air. Aerial performance is therefore limited by the efficiency with which momentum is imparted to the air, a property that can be measured using modern techniques. We measured the induced flow fields around six hawkmoth species flying tethered in a wind tunnel to assess span efficiency, ei, and from these measurements, determined the morphological and kinematic characters that predict efficient flight. The species were selected to represent a range in wingspan from 40 to 110 mm (2.75 times) and in mass from 0.2 to 1.5 g (7.5 times) but they were similar in their overall shape and their ecology. From high spatio-temporal resolution quantitative wake images, we extracted time-resolved downwash distributions behind the hawkmoths, calculating instantaneous values of ei throughout the wingbeat cycle as well as multi-wingbeat averages. Span efficiency correlated positively with normalized lift and negatively with advance ratio. Average span efficiencies for the moths ranged from 0.31 to 0.60 showing that the standard generic value of 0.83 used in previous studies of animal flight is not a suitable approximation of aerodynamic performance in insects.

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.

Aerodynamic flight performance in flap-gliding birds and bats.

Many birds use a flight mode called undulating or flap-gliding flight, where they alternate between flapping and gliding phases, while only a few bats make use of such a flight mode. Among birds, flap-gliding is commonly used by medium to large species, where it is regarded to have a lower energetic cost than continuously flapping flight. Here, we introduce a novel model for estimating the energetic flight economy of flap-gliding animals, by determining the lift-to-drag ratio for flap-gliding based on empirical lift-to-drag ratio estimates for continuous flapping flight and for continuous gliding flight, respectively. We apply the model to flight performance data of the common swift (Apus apus) and of the lesser long-nosed bat (Leptonycteris yerbabuenae). The common swift is a typical flap-glider while-to the best of our knowledge-the lesser long-nosed bat does not use flap-gliding. The results show that, according to the model, the flap-gliding common swift saves up to 15% energy compared to a continuous flapping swift, and that this is primarily due to the exceptionally high lift-to-drag ratio in gliding flight relative to that in flapping flight for common swifts. The lesser long-nosed bat, on the other hand, seems not to be able to reduce energetic costs by flap-gliding. The difference in relative costs of flap-gliding flight between the common swift and the lesser long-nosed bat can be explained by differences in morphology, flight style and wake dynamics. The model presented here proves to be a valuable tool for estimating energetic flight economy in flap-gliding animals. The results show that flap-gliding flight that is naturally used by common swifts is indeed the most economic one of the two flight modes, while this is not the case for the non-flap-gliding lesser long-nosed bat.

Efficiency of lift production in flapping and gliding flight of swifts.

Many flying animals use both flapping and gliding flight as part of their routine behaviour. These two kinematic patterns impose conflicting requirements on wing design for aerodynamic efficiency and, in the absence of extreme morphing, wings cannot be optimised for both flight modes. In gliding flight, the wing experiences uniform incident flow and the optimal shape is a high aspect ratio wing with an elliptical planform. In flapping flight, on the other hand, the wing tip travels faster than the root, creating a spanwise velocity gradient. To compensate, the optimal wing shape should taper towards the tip (reducing the local chord) and/or twist from root to tip (reducing local angle of attack). We hypothesised that, if a bird is limited in its ability to morph its wings and adapt its wing shape to suit both flight modes, then a preference towards flapping flight optimization will be expected since this is the most energetically demanding flight mode. We tested this by studying a well-known flap-gliding species, the common swift, by measuring the wakes generated by two birds, one in gliding and one in flapping flight in a wind tunnel. We calculated span efficiency, the efficiency of lift production, and found that the flapping swift had consistently higher span efficiency than the gliding swift. This supports our hypothesis and suggests that even though swifts have been shown previously to increase their lift-to-drag ratio substantially when gliding, the wing morphology is tuned to be more aerodynamically efficient in generating lift during flapping. Since body drag can be assumed to be similar for both flapping and gliding, it follows that the higher total drag in flapping flight compared with gliding flight is primarily a consequence of an increase in wing profile drag due to the flapping motion, exceeding the reduction in induced drag.