The Aerodynamics of Hovering Insect Flight. III. Kinematics
24 Feb 1984-Philosophical Transactions of the Royal Society B (The Royal Society)-Vol. 305, Iss: 1122, pp 41-78
TL;DR: In this paper, a projection analysis technique is described that solves for the orientation of the animal with respect to a cam era-based coordinate system, giving full kinematic details for the longitudinal wing and body axes from single-view films.
Abstract: Insects in free flight were filmed at 5000 frames per second to determine the motion of their wings and bodies. General comments are offered on flight behaviour and manoeuvrability. Changes in the tilt of the stroke plane with respect to the horizontal provides kinematic control of manoeuvres, analogous to the type of control used for helicopters. A projection analysis technique is described that solves for the orientation of the animal with respect to a cam era-based coordinate system, giving full kinematic details for the longitudinal wing and body axes from single-view films. The technique can be applied to all types of flight where the wing motions are bilaterally symmetrical: forward, backward and hovering flight, as well as properly banked turns. An analysis of the errors of the technique is presented, and shows that the reconstructed angles for wing position should be accurate to within 1-2° in general. Although measurement of the angles of attack was not possible, visual estimations are given. Only 11 film sequences show flight velocities and accelerations that are small enough for the flight to be considered as ‘hovering’. Two sequences are presented for a hover-fly using an inclined stroke plane, and nine sequences of hovering with a horizontal stroke plane by another hover-fly, two crane-flies, a drone-fly, a ladybird beetle, a honey bee, and two bumble bees. In general, oscillations in the body position from its mean motion are within measurement error, about 1-2 % of the wing length. The amplitudes of oscillation for the body angle are only a few degrees, but the phase relation of this oscillation to the wingbeat cycle could be determined for a few sequences. The phase indicates that the pitching moments governing the oscillations result from the wing lift at the ends of the wingbeat, and not from the wing drag or inertial forces. The mean pitching moment of the wings, which determines the mean body angle, is controlled by shifting the centre of lift over the cycle by changing the mean positional angle of the flapping wings. Deviations of the wing tip path from the stroke plane are never large, and no consistent pattern could be found for the wing paths of different insects; indeed, variations in the path were even observed for individual insects. The wing motion is not greatly different from simple harmonic motion, but does show a general trend towards higher accelerations and decelerations at either end of the wingbeat, with constant velocities during the middle of half-strokes. Root mean square and cube root mean cube angular velocities are on average about 4 and 9% lower than simple harmonic motion. Angles of attack are nearly constant during the middle of half-strokes, typically 35° at a position 70 % along the wing length. The wing is twisted along its length, with angles of attack at the wing base some 10-20° greater than at the tip. The wings rotate through about 110° at either end of the wingbeat during 10-20 % of the cycle period. The mean velocity of the wing edges during rotation is similar to the mean flapping velocity of the wing tip and greater than the flapping velocity for more proximal wing regions, which indicates that vortex shedding during rotation is com parable with that during flapping. The wings tend to rotate as a flat plate during the first half of rotation, which ends just before, or at, the end of the half-stroke. The hover-fly using an inclined stroke plane provides a notable exception to this general pattern : pronation is delayed and overlaps the beginning of the downstroke. The wing profile flexes along a more or less localized longitudinal axis during the second half of rotation, generating the ‘flip’ profile postulated by Weis-Fogh for the hover-flies. This profile occurs to some extent for all of the insects, and is not exceptionally pronounced for the hover-fly. By the end of rotation the wings are nearly flat again, although a slight camber can sometimes be seen. Weis-Fogh showed that beneficial aerodynamic interference can result when the left and right wings come into contact during rotation at the end of the wingbeat. His ‘fling’ mechanism creates the circulation required for wing lift on the subsequent half-stroke, and can be seen on my films of the Large Cabbage White butterfly, a plum e moth, and the Mediterranean flour moth. However, their wings ‘peel’ apart like two pieces of paper being separated, rather than fling open rigidly about the trailing edges. A ‘partial fling’ was found for some insects, with the wings touching only along posterior wing areas. A ‘ near fling ’ with the wings separated by a fraction of the chord was also observed for m any insects. There is a continuous spectrum for the separation distance between the wings, in fact, and the separation can vary for a given insect during different manoeuvres. It is suggested that these variants on Weis-Fogh’s fling mechanism also generate circulation for wing lift, although less effectively than a complete fling, and that changes in the separation distance may provide a fine control over the amount of lift produced.
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TL;DR: 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 are reviewed.
Abstract: 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.
1,182 citations
TL;DR: In this article, a review of the recent progress in flapping wing aerodynamics and aeroelasticity is presented, where it is realized that a variation of the Reynolds number (wing sizing, flapping frequency, etc.) leads to a change in the leading edge vortex (LEV) and spanwise flow structures, which impacts the aerodynamic force generation.
877 citations
Cites background from "The Aerodynamics of Hovering Insect..."
...Moreover, the body angle and the stroke plane angle vary in accordance with the flight speed and flapping wing kinematics of biological flyers [1,28,29]....
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TL;DR: Design characteristics of insect-based flying machines are presented, along with estimates of the mass supported, the mechanical power requirement and maximum flight speeds over a wide range of sizes and frequencies.
Abstract: The wing motion in free flight has been described for insects ranging from 1 to 100 mm in wingspan. To support the body weight, the wings typically produce 2–3 times more lift than can be accounted for by conventional aerodynamics. Some insects use the fling mechanism: the wings are clapped together and then flung open before the start of the downstroke, creating a lift-enhancing vortex around each wing. Most insects, however, rely on a leading-edge vortex (LEV) created by dynamic stall during flapping; a strong spanwise flow is also generated by the pressure gradients on the flapping wing, causing the LEV to spiral out to the wingtip. Technical applications of the fling are limited by the mechanical damage that accompanies repeated clapping of the wings, but the spiral LEV can be used to augment the lift production of propellers, rotors and micro-air vehicles (MAVs). Design characteristics of insect-based flying machines are presented, along with estimates of the mass supported, the mechanical power requirement and maximum flight speeds over a wide range of sizes and frequencies. To support a given mass, larger machines need less power, but smaller ones operating at higher frequencies will reach faster speeds.
764 citations
TL;DR: A standard quasi-steady model of insect flight is modified to include rotational forces, translational forces and the added mass inertia, and the revised model predicts the time course of force generation for several different patterns of flapping kinematics more accurately than a model based solely on translational force coefficients.
Abstract: We used a dynamically scaled model insect to measure the rotational forces produced by a flapping insect wing. A steadily translating wing was rotated at a range of constant angular velocities, and the resulting aerodynamic forces were measured using a sensor attached to the base of the wing. These instantaneous forces were compared with quasi-steady estimates based on translational force coefficients. Because translational and rotational velocities were constant, the wing inertia was negligible, and any difference between measured forces and estimates based on translational force coefficients could be attributed to the aerodynamic effects of wing rotation. By factoring out the geometry and kinematics of the wings from the rotational forces, we determined rotational force coefficients for a range of angular velocities and different axes of rotation. The measured coefficients were compared with a mathematical model developed for two-dimensional motions in inviscid fluids, which we adapted to the three-dimensional case using blade element theory. As predicted by theory, the rotational coefficient varied linearly with the position of the rotational axis for all angular velocities measured. The coefficient also, however, varied with angular velocity, in contrast to theoretical predictions. Using the measured rotational coefficients, we modified a standard quasi-steady model of insect flight to include rotational forces, translational forces and the added mass inertia. The revised model predicts the time course of force generation for several different patterns of flapping kinematics more accurately than a model based solely on translational force coefficients. By subtracting the improved quasi-steady estimates from the measured forces, we isolated the aerodynamic forces due to wake capture.
746 citations
Cites background or methods or result from "The Aerodynamics of Hovering Insect..."
...Bennett (1970) conducted experiments with a dynamically scaled flapping model wing and showed that rotations alter aerodynamic forces at Reynolds numbers in the range 10 2 to approximately 10 3 . In a detailed overview of insect flight aerodynamics, Ellington (1984c) proposed a scheme to include wing rotation with translation in quasi-steady models....
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...Among insects, values of x̂0 are thought to lie between 0.25 and 0.5 (Ellington, 1984d), although very few studies have attempted to measure this parameter precisely....
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...To determine the validity of the quasi-steady 1094 assumption, Ellington (1984a) constructed the following logical argument....
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...In a detailed overview of insect flight aerodynamics, Ellington (1984c) proposed a scheme to include wing rotation with translation in quasi-steady models....
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...A quasi-steady treatment of the aerodynamic force due to wing rotation was derived by Fung (1969) (see also Theodorsen, 1935; Sedov, 1965; Ellington, 1984c ) for small-amplitude flutter on thin, rigid wings....
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TL;DR: A dynamically scaled mechanical model of the fruit fly Drosophila melanogaster is used to study how changes in wing kinematics influence the production of unsteady aerodynamic forces in insect flight, finding no evidence that stroke deviation can augment lift, but it nevertheless may be used to modulate forces on the two wings.
Abstract: We used a dynamically scaled mechanical model of the fruit fly Drosophila melanogaster to study how changes in wing kinematics influence the production of unsteady aerodynamic forces in insect flight. We examined 191 separate sets of kinematic patterns that differed with respect to stroke amplitude, angle of attack, flip timing, flip duration and the shape and magnitude of stroke deviation. Instantaneous aerodynamic forces were measured using a two-dimensional force sensor mounted at the base of the wing. The influence of unsteady rotational effects was assessed by comparing the time course of measured forces with that of corresponding translational quasi-steady estimates. For each pattern, we also calculated mean stroke-averaged values of the force coefficients and an estimate of profile power. The results of this analysis may be divided into four main points.
(i) For a short, symmetrical wing flip, mean lift was optimized by a stroke amplitude of 180° and an angle of attack of 50°. At all stroke amplitudes, mean drag increased monotonically with increasing angle of attack. Translational quasi-steady predictions better matched the measured values at high stroke amplitude than at low stroke amplitude. This discrepancy was due to the increasing importance of rotational mechanisms in kinematic patterns with low stroke amplitude.
(ii) For a 180° stroke amplitude and a 45° angle of attack, lift was maximized by short-duration flips occurring just slightly in advance of stroke reversal. Symmetrical rotations produced similarly high performance. Wing rotation that occurred after stroke reversal, however, produced very low mean lift.
(iii) The production of aerodynamic forces was sensitive to changes in the magnitude of the wing’s deviation from the mean stroke plane (stroke deviation) as well as to the actual shape of the wing tip trajectory. However, in all examples, stroke deviation lowered aerodynamic performance relative to the no deviation case. This attenuation was due, in part, to a trade-off between lift and a radially directed component of total aerodynamic force. Thus, while we found no evidence that stroke deviation can augment lift, it nevertheless may be used to modulate forces on the two wings. Thus, insects might use such changes in wing kinematics during steering maneuvers to generate appropriate force moments.
(iv) While quasi-steady estimates failed to capture the time course of measured lift for nearly all kinematic patterns, they did predict with reasonable accuracy stroke-averaged values for the mean lift coefficient. However, quasi-steady estimates grossly underestimated the magnitude of the mean drag coefficient under all conditions. This discrepancy was due to the contribution of rotational effects that steady-state estimates do not capture. This result suggests that many prior estimates of mechanical power based on wing kinematics may have been grossly underestimated.
726 citations
Cites background or methods or result from "The Aerodynamics of Hovering Insect..."
...…2 ν −1 AR −1 , where Φ is stroke amplitude, n is wingbeat frequency, R is wing length, ν is kinematic viscosity, aspect ratio AR is 4R 2 S −1 and S is the surface area of a wing pair; Ellington, 1984c) and the reduced frequency parameter (body velocity/wing velocity) constant (Spedding, 1993)....
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...It is worth noting that the range of CD – values is much higher than has been previously reported for Drosophila virilis wings under steady-state conditions (Vogel, 1967) or estimated on the basis of Reynolds number (CD≈0.7; Ellington, 1984c)....
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...From the forces on each wing, we calculated the corresponding mean force coefficients using an equation derived from blade element theory (Ellington, 1984c; Dickinson et al., 1999): where F – is the magnitude of a specific force component (lift, drag, radial, total) averaged over the stroke, Φ is…...
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...These patterns were chosen because they roughly approximate patterns described for a variety of insects (Ellington, 1984b; Zanker, 1990a)....
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...The waveform was smoothed to minimize inertial accelerations during stroke reversal and to match more closely published stroke kinematics from a variety of insects (Ellington, 1984b; Zanker, 1990a)....
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References
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TL;DR: In this article, the average lift coefficient, Reynolds number, the aerodynamic power, the moment of inertia of the wing mass and the dynamic efficiency in animals which perform normal hovering with horizontally beating wings are derived.
Abstract: 1. On the assumption that steady-state aerodynamics applies, simple analytical expressions are derived for the average lift coefficient, Reynolds number, the aerodynamic power, the moment of inertia of the wing mass and the dynamic efficiency in animals which perform normal hovering with horizontally beating wings. 2. The majority of hovering animals, including large lamellicorn beetles and sphingid moths, depend mainly on normal aerofoil action. However, in some groups with wing loading less than 10 N m -2 (1 kgf m -2 ), non-steady aerodynamics must play a major role, namely in very small insects at low Reynolds number, in true hover-flies (Syrphinae), in large dragonflies (Odonata) and in many butterflies (Lepidoptera Rhopalocera). 3. The specific aerodynamic power ranges between 1.3 and 4.7 WN -1 (11-40 cal h -1 gf -1 ) but power output does not vary systematically with size, inter alia because the lift/drag ratio deteriorates at low Reynolds number. 4. Comparisons between metabolic rate, aerodynamic power and dynamic efficiency show that the majority of insects require and depend upon an effective elastic system in the thorax which counteracts the bending moments caused by wing inertia. 5. The free flight of a very small chalcid wasp Encarsia formosa has been analysed by means of slow-motion films. At this low Reynolds number (10-20), the high lift co-efficient of 2 or 3 is not possible with steady-state aerodynamics and the wasp must depend almost entirely on non-steady flow patterns. 6. The wings of Encarsia are moved almost horizontally during hovering, the body being vertical, and there are three unusual phases in the wing stroke: the clap , the fling and the flip . In the clap the wings are brought together at the top of the morphological upstroke. In the fling, which is a pronation at the beginning of the morphological downstroke, the opposed wings are flung open like a book, hinging about their posterior margins. In the flip, which is a supination at the beginning of the morphological upstroke, the wings are rapidly twisted through about 180°. 7. The fling is a hitherto undescribed mechanism for creating lift and for setting up the appropriate circulation over the wing in anticipation of the downstroke. In the case of Encarsia the calculated and observed wing velocities at which lift equals body weight are in agreement, and lift is produced almost instantaneously from the beginning of the downstroke and without any Wagner effect. The fling mechanism seems to be involved in the normal flight of butterflies and possibly of Drosophila and other small insects. Dimensional and other considerations show that it could be a useful mechanism in birds and bats during take-off and in emergencies. 8. The flip is also believed to be a means of setting up an appropriate circulation around the wing, which has hitherto escaped attention; but its operation is less well understood. It is not confined to Encarsia but operates in other insects, not only at the beginning of the upstroke (supination) but also at the beginning of the downstroke where a flip (pronation) replaces the clap and fling of Encarsia . A study of freely flying hover-flies strongly indicates that the Syrphinae (and Odonata) depend almost entirely upon the flip mechanism when hovering. In the case of these insects a transient circulation is presumed to be set up before the translation of the wing through the air, by the rapid pronation (or supination) which affects the stiff anterior margin before the soft posterior portions of the wing. In the flip mechanism vortices of opposite sense must be shed, and a Wagner effect must be present. 9. In some hovering insects the wing twistings occur so rapidly that the speed of propagation of the elastic torsional wave from base to tip plays a significant role and appears to introduce beneficial effects. 10. Non-steady periods, particularly flip effects, are present in all flapping animals and they will modify and become superimposed upon the steady-state pattern as described by the mathematical model presented here. However, the accumulated evidence indicates that the majority of hovering animals conform reasonably well with that model. 11. Many new types of analysis are indicated in the text and are now open for future theoretical and experimental research.
1,279 citations
Journal Article•
375 citations
TL;DR: It is argued that the tilt of the stroke plane relative to the horizontal is an adaptation to the geometrically unfavourable induced wind and to the relatively large lift/drag ratio seen in many insects.
Abstract: 1. Expressions have been derived for an estimate of the average coefficient of lift, for the variation in bending moment or torque caused by wind forces and by inertia forces, and for the power output during hovering flight on one spot when the wings move according to a horizontal figure-of-eight. 2. In both hummingbirds and Drosophila the flight is consistent with steady-state aerodynamics, the average lift coefficient being 1.8 in the hummingbird and 0.8 in Drosophila. 3. The aerodynamic or hydraulic efficiency is 0.5 in the hummingbird and 0.3 in Drosophila, and in both types the aerodynamic power output is 22-24 cal/g body weight/h. 4. The total mechanical power output is 39 cal g-1 h-1 in the hummingbird because of the extra energy needed to accelerate the wing-mass. It is 24 cal g-1 h-1 in Drosophila in which the inertia term is negligible because the wing-stroke frequency is reduced to the lowest possible value for sustained flight. 5. In both animals the mechanical efficiency of the flight muscles is 0.2. 6. It is argued that the tilt of the stroke plane relative to the horizontal is an adaptation to the geometrically unfavourable induced wind and to the relatively large lift/drag ratio seen in many insects. The vertical movements at the extreme ends may serve to reduce the interaction between the shed ‘stopping’ vortex and the new bound vortex of opposite sense which has to be built up during the early part of the return stroke. 7. Two additional non-steady flow situations may exist at either end of the stroke, delayed stall and delayed build-up of circulation (Wagner effect), but since they have opposite effects it is probable that the resultant force is of about the same magnitude as that estimated for a steady-state situation. 8. Most insects have an effective elastic system to counteract the adverse effect of wing-inertia, but small fast-moving vertebrates have not. It is argued that the only material available for this purpose in this group is elastin and that it is unsuited at the rates of deformation required because recent measurements have shown that the damping is relatively high, probably due to molecular factors.
349 citations
TL;DR: Weis-Fogh as mentioned in this paper proposed a new mechanism of lift generation, which could work even in inviscid two-dimensional motions starting from rest, when Kelvin's theorem states that the total circulation round a body must vanish, but does not exclude the possibility that if the body breaks into two pieces then there may be equal and opposite circulations round them, each suitable for generating the lift required in the pieces' subsequent motions.
Abstract: Weis-Fogh (1973) proposed a new mechanism of lift generation of fundamental interest. Surprisingly, it could work even in inviscid two-dimensional motions starting from rest, when Kelvin's theorem states that the total circulation round a body must vanish, but does not exclude the possibility that if the body breaks into two pieces then there may be equal and opposite circulations round them, each suitable for generating the lift required in the pieces’ subsequent motions! The ‘fling’ of two insect wings of chord c (figure 1) turning with angular velocity Ω generates irrotational motions associated with the sucking of air into the opening gap which are calculated in § 2 as involving circulations −0·69Ωc2 and + 0.69Ωc2 around the wings when their trailing edges, which are stagnation points of those irrotational motions, break apart (position (f)). Viscous modifications to this irrotational flow pattern by shedding of vorticity at the boundary generate (§ 3) a leading-edge separation bubble, and tend to increase slightly the total bound vorticity. Its role in a three-dimensional picture of the Weis-Fogh mechanism of lift generation, involving formation of trailing vortices at the wing tips, and including the case of a hovering insect like Encarsia formosa moving those tips in circular paths, is investigated in § 4. The paper ends with the comment that the far flow field of such very small hovering insects should take the form of the exact solution (Landau 1944; Squire 1951) of the Navier-Stokes equations for the effect of a concentrated force (the weight mg of the animal) acting on a fluid of kinematic viscosity v and density p, whenever the ratio mg/pv2 is small enough for that jet-type induced motion to be stable.
341 citations
TL;DR: Comparative experiments with the housefly Musca domestica indicate that the principle of independent torque and thrust control by vision is adopted in at least two different species.
Abstract: Apparent motion was simulated in the visual system of the tethered fruitfly Drosophila melanogaster by projecting moving stripe patterns onto stationary screens positioned in front of the lateral eye regions. The reactions of the animal were recorded under conditions of stationary flight in still air. It was found that visual stimulation modifies, independently, torque and thrust of the flight system. The responses appear suitable to counteract involuntary changes of direction and altitude in free flight.
322 citations