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Unconventional lift-generating mechanisms in free-flying butterflies

Abstract

Flying insects generate forces that are too large to be accounted for by conventional steady-state aerodynamics1,2. To investigate these mechanisms of force generation, we trained red admiral butterflies, Vanessa atalanta, to fly freely to and from artificial flowers in a wind tunnel, and used high-resolution, smoke-wire flow visualizations to obtain qualitative, high-speed digital images of the air flow around their wings. The images show that free-flying butterflies use a variety of unconventional aerodynamic mechanisms to generate force: wake capture3, two different types of leading-edge vortex3,4,5,6,7, active and inactive upstrokes8, in addition to the use of rotational mechanisms3 and the Weis–Fogh ‘clap-and-fling’ mechanism9,10,11,12. Free-flying butterflies often used different aerodynamic mechanisms in successive strokes. There seems to be no one ‘key’ to insect flight, instead insects rely on a wide array of aerodynamic mechanisms to take off, manoeuvre, maintain steady flight, and for landing.

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Figure 1: Butterfly leading edge vortex structure.
Figure 2: Flow over the wings during the downstroke.
Figure 3: Wake capture.
Figure 4: Upstrokes with positive, neutral and negative (downwards) loadings produced during a single flight by a 434-mg male flying into a 1.5 m s-1 wind.

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References

  1. Sane, S. P. & Dickinson, M. H. The aerodynamic effects of wing rotation and a revised quasi-steady model of flapping flight. J. Exp. Biol. 205, 1087–1096 (2002)

    PubMed  Google Scholar 

  2. Zbikowski, R. On aerodynamic modelling of an insect-like flapping wing in hover for micro air vehicles. Phil. Trans. R. Soc. Lond. A 360, 273–290 (2002)

    Article  ADS  Google Scholar 

  3. Dickinson, M. H., Lehmann, F.-O. & Sane, S. P. Wing rotation and the aerodynamic basis of insect flight. Science 284, 1954–1960 (1999)

    Article  CAS  Google Scholar 

  4. Maxworthy, T. The fluid-dynamics of insect flight. Ann. Rev. Fluid Mech. 13, 329–350 (1981)

    Article  ADS  Google Scholar 

  5. Ellington, C. P., van den Berg, C., Willmott, A. P. & Thomas, A. L. R. Leading-edge vortices in insect flight. Nature 384, 626–630 (1996)

    Article  ADS  CAS  Google Scholar 

  6. Van den Berg, C. & Ellington, C. P. The vortex wake of a hovering model hawkmoth. Phil. Trans. R. Soc. Lond B 352, 317–328 (1997)

    Article  ADS  Google Scholar 

  7. Birch, J. M. & Dickinson, M. H. Spanwise flow and the attachment of the leading-edge vortex on insect wings. Nature 412, 729–733 (2001)

    Article  ADS  CAS  Google Scholar 

  8. Saffman, P. G. & Sheffield, J. S. Flow over a wing with an attached free vortex. Studies Appl. Math. 57, 107–117 (1977)

    Article  MathSciNet  Google Scholar 

  9. Weis-Fogh, T. Quick estimates of flight fitness in hovering animals, including novel mechanisms for lift production. J. Exp. Biol. 59, 169–230 (1973)

    Google Scholar 

  10. Lighthill, J. On the Weis-Fogh mechanism of lift generation. J. Fluid Mech. 60, 1–17 (1973)

    Article  ADS  MathSciNet  Google Scholar 

  11. Maxworthy, T. Experiments on the Weis-Fogh mechanism of lift generation by insects in hovering flight. Part 1. Dynamics of the ‘fling’. J. Fluid Mech. 93, 47–63 (1979)

    Article  ADS  Google Scholar 

  12. Ellington, C. P. Biological Fluid Dynamics Symp. Soc. Exp. Biol. (eds Ellington, C. P. & Pedley, T. J.) Vol. 49, 109–129 (Company of Biologists, Cambridge, 1995)

    Google Scholar 

  13. Rossow, V. J. Lift enhancement by an externally trapped vortex. J. Aircraft 15, 618–625 (1978)

    Article  Google Scholar 

  14. Rossow, V. J. Two-fence concept for efficient trapping of vortices on airfoils. J. Aircraft 29, 847–855 (1992)

    Article  Google Scholar 

  15. Rossow, V. J. Aerodynamics of airfoils with vortex trapped by two spanwise fences. J. Aircraft 31, 146–153 (1994)

    Article  Google Scholar 

  16. Huang, M.-K. & Chow, C.-Y. Trapping of a free vortex by Joukowski airfoils. Am. Inst. Aeronaut. Astronaut. J. 20, 292–298 (1982)

    Article  Google Scholar 

  17. Mourtos, N. J. & Brooks, M. Flow past a flat plate with a vortex/sink combination. J. Appl. Mech. 63, 543–550 (1996)

    Article  ADS  Google Scholar 

  18. Riddle, T. W., Wadcock, A. J., Tso, J. & Cummings, R. M. An experimental analysis of vortex trapping techniques. J. Fluids Eng. 121, 555–559 (1999)

    Article  Google Scholar 

  19. Gursul, I. & Ho, C.-M. High aerodynamic loads on an airfoil submerged in an unsteady stream. Am. Inst. Aeronaut. Astronaut. J. 30, 1117–1119 (1992)

    Article  Google Scholar 

  20. Gad-el-Hak, M. & Ho, C.-M. Unsteady vortical lift around three-dimensional lifting surfaces. Am. Inst. Aeronaut. Astronaut. J. 24, 713–721 (1986)

    Article  Google Scholar 

  21. Délery, J. M., Legendre, R. & Werlé, H. Toward the elucidation of three-dimensional separation. Ann. Rev. Fluid Mech. 33, 129–154 (2001)

    Article  ADS  Google Scholar 

  22. Perry, A. E. & Chong, M. S. A description of eddying motions and flow patterns using critical-point concepts. Ann. Rev. Fluid Mech. 19, 125–155 (1987)

    Article  ADS  Google Scholar 

  23. Tobak, M. & Peake, D. J. Topology of three-dimensional separated flows. Ann. Rev. Fluid Mech. 14, 61–85 (1982)

    Article  ADS  MathSciNet  Google Scholar 

  24. Lighthill, M. J. Laminar Boundary Layer Theory Section II 2.6 (ed. Rosenhead, L.) 72–82 (Oxford Univ. Press, New York, 1963)

    Google Scholar 

  25. Legendre, R. Separation de L'ecoulement laminaire tridimensionnel. Rech. Aeronaut. 54, 3–8 (1956)

    Google Scholar 

  26. Poincaré, H. Les points singulièrs des équations différentielles. C.R. Acad. Sci. Paris 94, 416–418 (1882)

    MATH  Google Scholar 

  27. Hornung, H. & Perry, A. E. Some aspects of three-dimensional separation. I: Streamsurface bifurcations. Z. Flugwiss. Weltraumforsch. 8, 77–87 (1984)

    Google Scholar 

  28. Willmott, A. P., Ellington, C. P. & Thomas, A. L. R. Flow visualisation and unsteady aerodynamics in the flight of the hawkmoth Manduca sexta. Phil. Trans. R. Soc. Lond. B 352, 303–316 (1997)

    Article  ADS  Google Scholar 

  29. Rayner, J. M. V., Jones, G. & Thomas, A. L. R. Vortex flow visualizations reveal change in upstroke function with flight speed in bats. Nature 321, 162–164 (1986)

    Article  ADS  Google Scholar 

  30. Spedding, G. R. Advances in Comparative Environmental Physiology 11. Mechanics of Animal Locomotion (ed. Alexander, R. M.) (Springer, Berlin, 1993)

    Google Scholar 

Download references

Acknowledgements

We thank the Engineering and Physical Sciences Research Council instrument pool for use of their NAC500 high-speed video camera. R.B.S. was supported by a Biotechnology and Biological Sciences Research Council grant to A.L.R.T. A.L.R.T. was supported by a Royal Society University Research Fellowship.

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Correspondence to A. L. R. Thomas.

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Srygley, R., Thomas, A. Unconventional lift-generating mechanisms in free-flying butterflies. Nature 420, 660–664 (2002). https://doi.org/10.1038/nature01223

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