Hostname: page-component-cd9895bd7-p9bg8 Total loading time: 0 Render date: 2024-12-18T21:59:34.394Z Has data issue: false hasContentIssue false

A vortex theory of animal flight. Part 1. The vortex wake of a hovering animal

Published online by Cambridge University Press:  19 April 2006

J. M. V. Rayner
Affiliation:
Department of Applied Mathematics and Theoretical Physics, University of Cambridge Present address: Department of Zoology, University of Bristol.

Abstract

The distribution of vorticity in the wake of a hovering bird or insect is considered. The wake is modelled by a chain of coaxial small-cored circular vortex rings stacked one upon another; each member of the chain is generated by a single wing-stroke. Circulation is determined by the animal's weight and the time for which a single ring must provide lift; ring size is calculated from the circulation distribution on the animal's wing. The theory is equally applicable to birds and insects, although the mechanism of ring formation differs. This approach avoids the use of lift and drag coefficients and is not bound by the constraints of steady-state aerodynamics; it gives a wake configuration in agreement with experimental observations. The classical momentum jet approach has steady momentum flux in the wake, and is difficult to relate to the wing motions of a hovering bird or insect; the vortex wake can be related to the momentum jet, but adjacent vortex elements are disjoint and momentum flux is periodic.

The evolution of the wake starting from rest is considered by releasing vortex rings at appropriate time intervals and allowing them to interact in their own velocity fields. The resulting configuration depends on the feathering parameter f (which depends on the animal's morphology); f increases with body size. At the lower end of the wake rings coalesce to form a single large vortex, which breaks away from the rest of the wake at intervals. Wake contraction depends on f; the minimum areal contraction of one-half (as in momentum-jet theory) occurs only in the limit f → 0, but values calculated for smaller insects of just over one-half suggest that the momentum jet may be a good approximation to the wake when f is small.

Induced power in hovering is calculated as the limit of the mean rate of increase of wake kinetic energy as time progresses. It can be related to the classical momentum-jet induced power by a simple conversion factor. For an insect or hummingbird the usual momentum-jet estimate may be between 10 and 15% too low, but for a bird it may be as much as 50% too low. This suggests that few, if any, birds are able to sustain aerobic hovering, and that as small a value of f as possible would be necessary if the bird were to hover.

Tip losses (energy cost of the vortex-ring wake compared with the equivalent momentum jet) are negligible for insects, but can be in the range 15–20% for birds.

Type
Research Article
Copyright
© 1979 Cambridge University Press

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

Batchelor, G. K. 1967 An Introduction to Fluid Dynamics. Cambridge University Press.
Cone, C. D. 1968 The aerodynamics of flapping birdflight. Spec. Sci. Rep. Va Inst. Mar. Sci. no. 52.Google Scholar
Ellington, C. P. 1978 The aerodynamics of normal hovering flight. In Comparative Physiology — Water, Ions and Fluid Mechanics (ed. K. Schmidt-Nielsen, L. Bolis & S. H. P. Maddrell), pp. 327345. Cambridge University Press.
Fraenkel, L. E. 1970 On steady vortex rings of small cross-section in an ideal fluid. Proc. Roy. Soc. A 316, 2962.Google Scholar
Glauert, H. 1935 Airplane propellors. In Aerodynamic Theory (ed. W. H. Durand), vol. 4, pp. 169360. Springer.
Gradshteyn, I. S. & Rhyzhik, I. M. 1965 Tables of Integrals, Series and Products. Academic Press.
Greenewalt, C. H. 1962 Dimensional relationships for flying animals. Smithson. Misc. Collns 144, 2.Google Scholar
Hainsworth, F. R. & Wolf, J. L. 1972 Power for hovering flight in relation to body size in hummingbirds. Am. Nat. 106, 589596.Google Scholar
Hoff, W. 1919 Der Flug der Insekten. Naturwissenschaften 7, 159164.Google Scholar
Kármán, T. von & Burgers, J. M. 1935 General aerodynamic theory — perfect fluids. In Aerodynamic theory (ed. W. H. Durand), vol. 2, pp. 315352. Springer.
Lamb, H. 1932 Hydrodynamics. Cambridge University Press.
Levy, H. & Forsdyke, A. G. 1927 The stability of an infinite system of circular vortices. I. Proc. Roy. Soc. A 114, 594604.Google Scholar
Lighthill, M. J. 1973 On the Weis—Fogh mechanism of lift generation. J. Fluid Mech. 60, 117.Google Scholar
Lighthill, M. J. 1977 Introduction to the scaling of aerial locomotion. In Scale Effects in Animal Locomotion (ed. T. J. Pedley), pp. 365404. Academic Press.
Magnan, A., Perrilliat-Botonet, C. & Girard, H. 1938 Essais d'enregistrements cinématographiques simultanées dans trois directions perpendiculaires deux à deux de l’écoulement de l'air autour d'un oiseau en vol. C. R. hebd. Séanc. Acad. Sci., Paris 206, 462464.Google Scholar
Maxworthy, T. 1972 The structure and stability of vortex rings. J. Fluid Mech. 51, 1532.Google Scholar
Maxworthy, T. 1977 Some experimental studies of vortex rings. J. Fluid Mech. 81, 465495.Google Scholar
Nachtigall, W. 1974 Insects in Flight. London: George Allen & Unwin.
Nachtigall, W. & Kempf, B. 1971 Vergleichende Untersuchungen zur flugbiologischen Funktion des Daumenfittichs (Alula spuria) bei Vögeln. Z. vergl. Physiol. 71, 326341.Google Scholar
Norberg, U. M. 1975 Hovering flight in the pied flycatcher (Ficedula hypoleuca). In Swimming and Flying in Nature (ed. T. Y. Wu, C. J. Brokaw & C. Brennen), vol. 2, pp. 869881. Plenum.
Norberg, U. M. 1976 Aerodynamics of hovering flight in the long-eared bat Plecotus auritus. J. Exp. Biol. 65, 459470.Google Scholar
Norbury, J. 1973 A family of steady vortex rings. J. Fluid Mech. 57, 417431.Google Scholar
Oehme, H. & Kitzler, U. 1974 Untersuchungen zur Flugbiophysik und Flugphysiologie der Vögel. I. Über die Kinematik des Flügelschlages beim unbeschleunigten Horizontalflug. Zool. Jb. Physiol. 78, 401512.Google Scholar
Oehme, H. & Kitzler, U. 1975a Untersuchungen zur Flugbiophysik und Flugphysiologie der Vögel. II. Zur Geometric des Vögelflügels. Zool. Jb. Physiol. 79, 402424.Google Scholar
Oehme, H. & Kitzler, U. 1975b Untersuchungen zur Flugbiophysik und Flugphysiologie der Vögel. III. Die Bestimmung der Muskelleistung beim Kraftflug der Vögel aus kinematischen und morphologischen Daten. Zool. Jb. Physiol. 79, 425458.Google Scholar
Osborne, M. F. M. 1951 Aerodynamics of flapping flight with applications to insects. J. Exp. Biol. 28, 221245.Google Scholar
Pennycuick, C. J. 1968 Power requirements for horizontal flight in the pigeon Columba livia. J. Exp. Biol. 49, 527555.Google Scholar
Rayner, J. M. V. 1979a A vortex theory of animal flight. Part 2. The forward flight of birds. J. Fluid Mech. 91, 739771.Google Scholar
Rayner, J. M. V. 1979b A new theory of animal flight mechanics. J. Exp. Biol. (In the Press.)Google Scholar
Rüppell, G. 1977 Bird Flight. Van Nostrand—Rheinhold.
Saffman, P. G. 1970 The velocity of viscous vortex rings. SIAM J. 49, 371380.Google Scholar
Vogel, S. 1967 Flight in Drosophila. III. Aerodynamic characteristics of fly wings and wing models. J. Exp. Biol. 46, 431443.Google Scholar
Weis-Fogh, T. 1972 Energetics of hovering flight in hummingbirds and Drosophila. J. Exp. Biol. 56, 79104.Google Scholar
Weis-Fook, T. 1973 Quick estimates of flight fitness in hovering animals, including novel mechanisms for lift production. J. Exp. Biol. 59, 169230.Google Scholar
Weis-Fogh, T. 1977 Dimensional analysis of hovering flight. In Scale Effects in Animal Locomotion (ed. T. J. Pedley), pp. 405420. Academic Press.
Weis-Fogh, T. & Alexander, R. McN. 1977 The sustained power output from striated muscle. In Scale Effects in Animal Locomotion (ed. T. J. Pedley), pp. 511525. Academic Press.
Widnall, S. E. & Sullivan, J. P. 1973 On the stability of vortex rings. Proc. Roy. Soc. A 332, 335353.Google Scholar
Widnall, S. E. & Tsai, C.-Y. 1977 The instability of the thin vortex ring of constant vorticity. Phil. Trans. Roy. Soc. A 287, 273305.Google Scholar