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      Efficiency of Lift Production in Flapping and Gliding Flight of Swifts

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          Abstract

          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.

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          Most cited references13

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          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.
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            Leading-edge vortex lifts swifts.

            The current understanding of how birds fly must be revised, because birds use their hand-wings in an unconventional way to generate lift and drag. Physical models of a common swift wing in gliding posture with a 60 degrees sweep of the sharp hand-wing leading edge were tested in a water tunnel. Interactions with the flow were measured quantitatively with digital particle image velocimetry at Reynolds numbers realistic for the gliding flight of a swift between 3750 and 37,500. The results show that gliding swifts can generate stable leading-edge vortices at small (5 degrees to 10 degrees) angles of attack. We suggest that the flow around the arm-wings of most birds can remain conventionally attached, whereas the swept-back hand-wings generate lift with leading-edge vortices.
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              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.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                28 February 2014
                : 9
                : 2
                : e90170
                Affiliations
                [1 ]Department of Biology, Lund University, Lund, Sweden
                [2 ]Structure & Motion Lab, The Royal Veterinary College, University of London, Hatfield, Hertfordshire, United Kingdom
                Scientific Institute Foundation Santa Lucia, Italy
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: PH AH. Performed the experiments: PH. Analyzed the data: PH AH RJB. Wrote the paper: PH AH RJB.

                Article
                PONE-D-13-36043
                10.1371/journal.pone.0090170
                3938594
                24587260
                f9321c32-3337-4c0a-b27b-66f51d459f1f
                Copyright @ 2014

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 2 September 2013
                : 31 January 2014
                Page count
                Pages: 7
                Funding
                This study was supported financially by grants from the Swedish Research Council and the Knut and Alice Wallenberg Foundation. It also received support from the Centre for Animal Movement Research (CAnMove) financed by a Linnaeus grant (349-2007-8690) from the Swedish Research Council and Lund University, the EPSRC (EP/H004025/1) and BBSRC (BB/J001244/1). RJB holds an EPSRC Career Acceleration Fellowship. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology
                Anatomy and physiology
                Musculoskeletal system
                Biomechanics
                Flight mechanics
                Biophysics
                Biomechanics
                Flight mechanics
                Zoology
                Ornithology
                Physics
                Biophysics
                Biomechanics
                Flight mechanics

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                Uncategorized

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