The aerodynamics of insect flight

SUMMARY The flight of insects has fascinated physicists and biologists for more than a century. Yet, until recently, researchers were unable to rigorously quantify the complex wing motions of flapping insects or measure the forces and flows around their wings. However, recent developments in high-speed videography and tools for computational and mechanical modeling have allowed researchers to make rapid progress in advancing our understanding of insect flight. These mechanical and computational fluid dynamic models, combined with modern flow visualization techniques, have revealed that the fluid dynamic phenomena underlying flapping flight are different from those of non-flapping, 2-D wings on which most previous models were based. In particular, even at high angles of attack, a prominent leading edge vortex remains stably attached on the insect wing and does not shed into an unsteady wake, as would be expected from non-flapping 2-D wings. Its presence greatly enhances the forces generated by the wing, thus enabling insects to hover or maneuver. In addition, flight forces are further enhanced by other mechanisms acting during changes in angle of attack, especially at stroke reversal, the mutual interaction of the two wings at dorsal stroke reversal or wing–wake interactions following stroke reversal. This progress has enabled the development of simple analytical and empirical models that allow us to calculate the instantaneous forces on flapping insect wings more accurately than was previously possible. It also promises to foster new and exciting multi-disciplinary collaborations between physicists who seek to explain the phenomenology, biologists who seek to understand its relevance to insect physiology and evolution, and engineers who are inspired to build micro-robotic insects using these principles. This review covers the basic physical principles underlying flapping flight in insects, results of recent experiments concerning the aerodynamics of insect flight, as well as the different approaches used to model these phenomena.

[1]  H A Hazen,et al.  THE MECHANICS OF FLIGHT. , 1893, Science.

[2]  H. J.,et al.  Hydrodynamics , 1924, Nature.

[3]  Herbert Wagner Über die Entstehung des dynamischen Auftriebes von Tragflügeln , 1925 .

[4]  Max M Munk Note on the Air Forces on a Wing Caused by Pitching. , 1925 .

[5]  Max M. Munk Elements of the Wing Section Theory and of the Wing Theory , 1925 .

[6]  H. Glauert The elements of aerofoil and airscrew theory , 1926 .

[7]  Elliott G Reid Airfoil lift with changing angle of attack , 1927 .

[8]  H. Glauert The force and moment on an oscillating aerofoil , 1930 .

[9]  O. Tietjens,et al.  Applied hydro- and aeromechanics , 1934 .

[10]  T. Theodorsen General Theory of Aerodynamic Instability and the Mechanism of Flutter , 1934 .

[11]  O. Tietjens,et al.  Fundamentals of hydro- and aeromechanics , 1934 .

[12]  W. Farren,et al.  THE REACTION ON A WING WHOSE ANGLE OF INCIDENCE IS CHANGING RAPIDLY WIND TUNNEL EXPERIMENTS WITH A SHORT PERIOD RECORDING BALANCE , 1935 .

[13]  I. E. Garrick Propulsion of a flapping and oscillating airfoil , 1936 .

[14]  Abe Silverstein,et al.  Experimental Verification of the Theory of Oscillating Airfoils , 1939 .

[15]  W. J. Duncan Theoretical Aerodynamics , 1948, Nature.

[16]  M. Osborne Aerodynamics of flapping flight with application to insects. , 1951, The Journal of experimental biology.

[17]  R. L. Halfman Experimental aerodynamic derivatives of a sinusoidally oscillating airfoil in two-dimensional flow , 1952 .

[18]  C. Truesdell The Kinematics Of Vorticity , 1954 .

[19]  Yuan-Cheng Fung,et al.  An introduction to the theory of aeroelasticity , 1955 .

[20]  M. Jensen Biology and physics of locust flight. III. The aerodynamics of locust flight , 1956, Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences.

[21]  P. Libby,et al.  Two-dimensional Problems in Hydrodynamics and Aerodynamics , 1965 .

[22]  S. Vogel Flight in Drosophila : III. Aerodynamic Characteristics of Fly Wing Sand Wing Models , 1967 .

[23]  S. Vogel Flight in Drosophila , 1967 .

[24]  G. Batchelor,et al.  An Introduction to Fluid Dynamics , 1968 .

[25]  L. Bennett Insect Flight: Lift and Rate of Change of Incidence , 1970, Science.

[26]  E. Polhamus Predictions of vortex-lift characteristics based on a leading-edge suction analogy. , 1971 .

[27]  M. Lighthill On the Weis-Fogh mechanism of lift generation , 1973, Journal of Fluid Mechanics.

[28]  T. Weis-Fogh Quick estimates of flight fitness in hovering animals , 1973 .

[29]  C. Brennen,et al.  Swimming and Flying in Nature , 1975, Springer US.

[30]  R. Norberg Hovering Flight of the Dragonfly Aeschna Juncea L., Kinematics and Aerodynamics , 1975 .

[31]  C. Rees Aerodynamic properties of an insect wing section and a smooth aerofoil compared , 1975, Nature.

[32]  L. Bennett Clap and Fling Aerodynamics-An Experimental Evaluation , 1977 .

[33]  B. G. Newman,et al.  The Role of Vortices and Unsteady Effects During the Hovering Flight of Dragonflies , 1979 .

[34]  J. Rayner A vortex theory of animal flight. Part 2. The forward flight of birds , 1979, Journal of Fluid Mechanics.

[35]  M. Cloupeau,et al.  Direct Measurements of Instantaneous Lift in Desert Locust; Comparison with Jensen'S Experiments on Detached Wings , 1979 .

[36]  J. Rayner A vortex theory of animal flight. Part 1. The vortex wake of a hovering animal , 1979, Journal of Fluid Mechanics.

[37]  T. N. Stevenson,et al.  Fluid Mechanics , 2021, Nature.

[38]  R. Buckholz Measurements of Unsteady Periodic Forces generated by the Blowfly Flying in a Wind Tunnel , 1981 .

[39]  J. Wu Theory for Aerodynamic Force and Moment in Viscous Flows , 1981 .

[40]  C. Ellington The Aerodynamics of Hovering Insect Flight. III. Kinematics , 1984 .

[41]  C. Ellington The Aerodynamics of Hovering Insect Flight. I. The Quasi-Steady Analysis , 1984 .

[42]  T. Daniel Unsteady Aspects of Aquatic Locomotion , 1984 .

[43]  C. Ellington THE AERODYNAMICS OF HOVERING INSECT FLIGHT. V. A VORTEX THEORY , 1984 .

[44]  C. Ellington The Aerodynamics of Hovering Insect Flight. II. Morphological Parameters , 1984 .

[45]  C. Ellington The Aerodynamics of Hovering Insect Flight. VI. Lift and Power Requirements , 1984 .

[46]  C. Ellington The Aerodynamics of Hovering Insect Flight. IV. Aeorodynamic Mechanisms , 1984 .

[47]  C. Somps,et al.  Dragonfly Flight: Novel Uses of Unsteady Separated Flows , 1985, Science.

[48]  I. Watanabe,et al.  Flight Mechanics of a Dragonfly , 1985 .

[49]  A. M. Kuethe,et al.  Foundations of aerodynamics: bases of aerodynamic design , 1986 .

[50]  G. Spedding,et al.  The generation of circulation and lift in a rigid two-dimensional fling , 1986, Journal of Fluid Mechanics.

[51]  J. Marden Maximum Lift Production During Takeoff in Flying Animals , 1987 .

[52]  A. Azuma,et al.  Flight Performance of a Dragonfly , 1988 .

[53]  A. R. Ennos INERTIAL AND AERODYNAMIC TORQUES ON THE WINGS OF DIPTERA IN FLIGHT , 1989 .

[54]  G. Rüppell Kinematic Analysis of Symmetrical Flight Manoeuvres of Odonata , 1989 .

[55]  A. R. Ennos The kinematics and aerodynamics of the free flight of some diptera , 1989 .

[56]  K. Götz,et al.  The Wing Beat of Drosophila Melanogaster. II. Dynamics , 1990 .

[57]  D. Acheson Elementary Fluid Dynamics , 1990 .

[58]  R. Dudley,et al.  Mechanics of Forward Flight in Bumblebees: I. Kinematics and Morphology , 1990 .

[59]  R. Dudley,et al.  Mechanics of Forward Flight in Bumblebees: II. QUASI-STEADY LIFT AND POWER REQUIREMENTS , 1990 .

[60]  R. Dudley Biomechanics of Flight in Neotropical Butterflies: Aerodynamics and Mechanical Power Requirements , 1991 .

[61]  S. Sunada,et al.  FUNDAMENTAL ANALYSIS OF THREE-DIMENSIONAL ‘NEAR FLING’ , 1993 .

[62]  M. Dickinson,et al.  The active control of wing rotation by Drosophila. , 1993, The Journal of experimental biology.

[63]  M. Dickinson,et al.  UNSTEADY AERODYNAMIC PERFORMANCE OF MODEL WINGS AT LOW REYNOLDS NUMBERS , 1993 .

[64]  M. Denny,et al.  Air and water : the biology and physics of life's media , 1993 .

[65]  A. K. Brodskiĭ,et al.  The evolution of insect flight , 1994 .

[66]  Dickinson,et al.  THE EFFECTS OF WING ROTATION ON UNSTEADY AERODYNAMIC PERFORMANCE AT LOW REYNOLDS NUMBERS , 1994, The Journal of experimental biology.

[67]  Smith,et al.  The advantages of an unsteady panel method in modelling the aerodynamic forces on rigid flapping wings , 1996, The Journal of experimental biology.

[68]  Adrian L. R. Thomas,et al.  Leading-edge vortices in insect flight , 1996, Nature.

[69]  Sato,et al.  The flight performance of a damselfly Ceriagrion melanurum Selys , 1997, The Journal of experimental biology.

[70]  M. Dickinson,et al.  The changes in power requirements and muscle efficiency during elevated force production in the fruit fly Drosophila melanogaster. , 1997, The Journal of experimental biology.

[71]  J. Spurk Boundary Layer Theory , 2019, Fluid Mechanics.

[72]  Willmott,et al.  Measuring the angle of attack of beating insect wings: robust three-dimensional reconstruction from two-dimensional images , 1997, The Journal of experimental biology.

[73]  C. Ellington,et al.  The three–dimensional leading–edge vortex of a ‘hovering’ model hawkmoth , 1997 .

[74]  C. Ellington,et al.  The mechanics of flight in the hawkmoth Manduca sexta. II. Aerodynamic consequences of kinematic and morphological variation. , 1997, The Journal of experimental biology.

[75]  S. Vogel,et al.  Life in Moving Fluids , 2020 .

[76]  K. Kawachi,et al.  A Numerical Study of Insect Flight , 1998 .

[77]  Ellington,et al.  A computational fluid dynamic study of hawkmoth hovering , 1998, The Journal of experimental biology.

[78]  C. Ellington The novel aerodynamics of insect flight: applications to micro-air vehicles. , 1999, The Journal of experimental biology.

[79]  M. Dickinson,et al.  Wing rotation and the aerodynamic basis of insect flight. , 1999, Science.

[80]  Mao Sun,et al.  Aerodynamic forces and flow structures of an airfoil in some unsteady motions at small Reynolds number , 2000 .

[81]  J A Walker,et al.  Mechanical performance of aquatic rowing and flying , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[82]  Z. J. Wang Two dimensional mechanism for insect hovering , 2000 .

[83]  H. Hamdani,et al.  A study on the mechanism of high-lift generation by an airfoil in unsteady motion at low reynolds number , 2001 .

[84]  M. Dickinson,et al.  Spanwise flow and the attachment of the leading-edge vortex on insect wings , 2001, Nature.

[85]  M. Dickinson,et al.  The control of flight force by a flapping wing: lift and drag production. , 2001, The Journal of experimental biology.

[86]  S. Sunada,et al.  Unsteady Forces on a Two-Dimensional Wing in Plunging and Pitching Motions , 2001 .

[87]  Jeffrey A. Walker,et al.  Rotational lift: something different or more of the same? , 2002, The Journal of experimental biology.

[88]  F. Minotti Unsteady two-dimensional theory of a flapping wing. , 2002, Physical review. E, Statistical, nonlinear, and soft matter physics.

[89]  M. Dickinson,et al.  The aerodynamic effects of wing rotation and a revised quasi-steady model of flapping flight. , 2002, The Journal of experimental biology.

[90]  J. Usherwood,et al.  The aerodynamics of revolving wings II. Propeller force coefficients from mayfly to quail. , 2002, The Journal of experimental biology.

[91]  A. Abbott Alliance for cellular signaling: Into unknown territory , 2002, Nature.

[92]  R. B. Srygley,et al.  Unconventional lift-generating mechanisms in free-flying butterflies , 2002, Nature.

[93]  Mao Sun,et al.  Unsteady aerodynamic force generation by a model fruit fly wing in flapping motion. , 2002, The Journal of experimental biology.

[94]  R. Zbikowski On aerodynamic modelling of an insect–like flapping wing in hover for micro air vehicles , 2002, Philosophical Transactions of the Royal Society of London. Series A: Mathematical, Physical and Engineering Sciences.

[95]  J. Usherwood,et al.  The aerodynamics of revolving wings I. Model hawkmoth wings. , 2002, The Journal of experimental biology.

[96]  R. Ramamurti,et al.  A three-dimensional computational study of the aerodynamic mechanisms of insect flight. , 2002, The Journal of experimental biology.

[97]  J. Dyszlewicz Two-dimensional problems , 2004 .

[98]  C. Ellington MECHANICS OF FORWARD FLIGHT IN BUMBLEBEES , 2005 .