Kinematic compensation for wing loss in flying damselflies

Flying insects can tolerate substantial wing wear before their ability to fly is entirely compromised. In order to keep flying with damaged wings, the entire flight apparatus needs to adjust its action to compensate for the reduced aerodynamic force and to balance the asymmetries in area and shape of the damaged wings. While several studies have shown that damaged wings change their flapping kinematics in response to partial loss of wing area, it is unclear how, in insects with four separate wings, the remaining three wings compensate for the loss of a fourth wing. We used high-speed video of flying blue-tailed damselflies (Ischnura elegans) to identify the wingbeat kinematics of the two wing pairs and compared it to the flapping kinematics after one of the hindwings was artificially removed. The insects remained capable of flying and precise maneuvering using only three wings. To compensate for the reduction in lift, they increased flapping frequency by 18 ± 15.4% on average. To achieve steady straight flight, the remaining intact hindwing reduced its flapping amplitude while the forewings changed their stroke plane angle so that the forewing of the manipulated side flapped at a shallower stroke plane angle. In addition, the angular position of the stroke reversal points became asymmetrical. When the wingbeat amplitude and frequency of the three wings were used as input in a simple aerodynamic model, the estimation of total aerodynamic force was not significantly different (paired t-test, p = 0.73) from the force produced by the four wings during normal flight. Thus, the removal of one wing resulted in adjustments of the motions of the remaining three wings, exemplifying the precision and plasticity of coordination between the operational wings. Such coordination is vital for precise maneuvering during normal flight but it also provides the means to maintain flight when some of the wings are severely damaged.     

Aerodynamic performance of a hovering hawkmoth with flexible wings: a computational approach

Insect wings are deformable structures that change shape passively and dynamically owing to inertial and aerodynamic forces during flight. It is still unclear how the three-dimensional and passive change of wing kinematics owing to inherent wing flexibility contributes to unsteady aerodynamics and energetics in insect flapping flight. Here, we perform a systematic fluid-structure interaction based analysis on the aerodynamic performance of a hovering hawkmoth, Manduca, with an integrated computational model of a hovering insect with rigid and flexible wings. Aerodynamic performance of flapping wings with passive deformation or prescribed deformation is evaluated in terms of aerodynamic force, power and efficiency. Our results reveal that wing flexibility can increase downwash in wake and hence aerodynamic force: first, a dynamic wing bending is observed, which delays the breakdown of leading edge vortex near the wing tip, responsible for augmenting the aerodynamic force-production; second, a combination of the dynamic change of wing bending and twist favourably modifies the wing kinematics in the distal area, which leads to the aerodynamic force enhancement immediately before stroke reversal. Moreover, an increase in hovering efficiency of the flexible wing is achieved as a result of the wing twist. An extensive study of wing stiffness effect on aerodynamic performance is further conducted through a tuning of Young's modulus and thickness, indicating that insect wing structures may be optimized not only in terms of aerodynamic performance but also dependent on many factors, such as the wing strength, the circulation capability of wing veins and the control of wing movements.     

How Could Beetle＇s Elytra Support Their Own Weight during Forward Flight？

The aerodynamic role of the elytra during a beetle＇s flapping motion is not well-elucidated, although it is well-recognized that the evolution of elytra has been a key in the success of coleopteran insects due to their protective function. An experimental study on wing kinematics reveals that for almost concurrent flapping with the hind wings, the flapping angle of the elytra is 5 times smaller than that of the hind wings. Then, we explore the aerodynamic forces on elytra in free forward flight with and without an effect of elytron-hind wing interaction by three-dimensional numerical simulation. The numerical results show that vertical force generated by the elytra without interaction is not sufficient to support even its own weight. However, the elytron-hind wing interaction improves the vertical force on the elytra up to 80%; thus, the total vertical force could fully support its own weight. The interaction slightly increases the vertical force on the hind wind by 6% as well.     

Wing flexibility enhances load-lifting capacity in bumblebees

The effect of wing flexibility on aerodynamic force production has emerged as a central question in insect flight research. However, physical and computational models have yielded conflicting results regarding whether wing deformations enhance or diminish flight forces. By experimentally stiffening the wings of live bumblebees, we demonstrate that wing flexibility affects aerodynamic force production in a natural behavioural context. Bumblebee wings were artificially stiffened in vivo by applying a micro-splint to a single flexible vein joint, and the bees were subjected to load-lifting tests. Bees with stiffened wings showed an 8.6 per cent reduction in maximum vertical aerodynamic force production, which cannot be accounted for by changes in gross wing kinematics, as stroke amplitude and flapping frequency were unchanged. Our results reveal that flexible wing design and the resulting passive deformations enhance vertical force production and load-lifting capacity in bumblebees, locomotory traits with important ecological implications.     

Numerical investigation of the aerodynamic characteristics of a hovering Coleopteran insect

The aerodynamic characteristics of the Coleopteran beetle species Epilachna quadricollis, a species with flexible hind wings and stiff elytra (fore wings), are investigated in terms of hovering flight. The flapping wing kinematics of the Coleopteran insect are modeled through experimental observations with a digital high-speed camera and curve fitting from an ideal harmonic kinematics model. This model numerically simulates flight by estimating a cross section of the wing as a two-dimensional elliptical plane. There is currently no detailed study on the role of the elytron or how the elytron-hind wing interaction affects aerodynamic performance. In the case of hovering flight, the relatively small vertical or horizontal forces generated by the elytron suggest that the elytron makes no significant contribution to aerodynamic force.     

Muscle function in avian flight: achieving power and control

Flapping flight places strenuous requirements on the physiological performance of an animal. Bird flight muscles, particularly at smaller body sizes, generally contract at high frequencies and do substantial work in order to produce the aerodynamic power needed to support the animal's weight in the air and to overcome drag. This is in contrast to terrestrial locomotion, which offers mechanisms for minimizing energy losses associated with body movement combined with elastic energy savings to reduce the skeletal muscles' work requirements. Muscles also produce substantial power during swimming, but this is mainly to overcome body drag rather than to support the animal's weight. Here, I review the function and architecture of key flight muscles related to how these muscles contribute to producing the power required for flapping flight, how the muscles are recruited to control wing motion and how they are used in manoeuvring. An emergent property of the primary flight muscles, consistent with their need to produce considerable work by moving the wings through large excursions during each wing stroke, is that the pectoralis and supracoracoideus muscles shorten over a large fraction of their resting fibre length (33-42%). Both muscles are activated while being lengthened or undergoing nearly isometric force development, enhancing the work they perform during subsequent shortening. Two smaller muscles, the triceps and biceps, operate over a smaller range of contractile strains (12-23%), reflecting their role in controlling wing shape through elbow flexion and extension. Remarkably, pigeons adjust their wing stroke plane mainly via changes in whole-body pitch during take-off and landing, relative to level flight, allowing their wing muscles to operate with little change in activation timing, strain magnitude and pattern.     

Time-Varying Wing-Twist Improves Aerodynamic Efficiency of Forward Flight in Butterflies

Insect wings can undergo significant chordwise (camber) as well as spanwise (twist) deformation during flapping flight but the effect of these deformations is not well understood. The shape and size of butterfly wings leads to particularly large wing deformations, making them an ideal test case for investigation of these effects. Here we use computational models derived from experiments on free-flying butterflies to understand the effect of time-varying twist and camber on the aerodynamic performance of these insects. High-speed videogrammetry is used to capture the wing kinematics, including deformation, of a Painted Lady butterfly (Vanessa cardui) in untethered, forward flight. These experimental results are then analyzed computationally using a high-fidelity, three-dimensional, unsteady Navier-Stokes flow solver. For comparison to this case, a set of non-deforming, flat-plate wing (FPW) models of wing motion are synthesized and subjected to the same analysis along with a wing model that matches the time-varying wing-twist observed for the butterfly, but has no deformation in camber. The simulations show that the observed butterfly wing (OBW) outperforms all the flat-plate wings in terms of usable force production as well as the ratio of lift to power by at least 29% and 46%, respectively. This increase in efficiency of lift production is at least three-fold greater than reported for other insects. Interestingly, we also find that the twist-only-wing (TOW) model recovers much of the performance of the OBW, demonstrating that wing-twist, and not camber is key to forward flight in these insects. The implications of this on the design of flapping wing micro-aerial vehicles are discussed.     

Aerial locomotion in flies and robots: kinematic control and aerodynamics of oscillating wings

Flight in flies results from a feedback cascade in which the animal converts mechanical power produced by the flight musculature into aerodynamic forces. A major goal of flight research is to understand the functional significance of the various components in this cascade ranging from the generation of the neural code, the control of muscle mechanical power output, wing kinematics and unsteady aerodynamic mechanisms. Here, I attempted to draw a broad outline on fluid dynamic mechanisms found in flapping insect wings such as leading edge vorticity, rotational circulation and wake capture momentum transfer, as well as on the constraints of flight force control by the neuromuscular system of the fruit fly Drosophila. This system-level perspective on muscle control and aerodynamic mechanisms is thought to be a fundamental bridge in any attempt to link the function and performance of the various flight components with their particular role for wing motion and aerodynamic control in the behaving animal. Eventually, this research might facilitate the development of man-made biomimetic autonomous micro air vehicles using flapping wing motion for propulsion that are currently under construction by engineers.     

Aerodynamic Interactions Between Wing and Body of a Model Insect in Forward Flight and Maneuvers

The aerodynamic interactions between the body and the wings of a model insect in forward flight and maneuvers are studied using the method of numerically solving the Navier-Stokes equations over moving overset grids. Three cases are considered, including a complete insect, wing pair only and body only. By comparing the results of these cases, the interaction effect between the body and the wing pair can be identified. The changes in the force and moment coefficients of the wing pair due to the presence of the body are less than 4.5% of the mean vertical force coefficient of the model insect; the changes in the aerodynamic force coefficients of the body due to the presence of the wings are less than 5.0% of the mean vertical force coefficient of the model insect. The results of this paper indicate that in studying the aerodynamics and flight dynamics of a flapping insect in forward flight or maneuver, separately computing (or measuring) the aerodynamic forces and moments on the wing pair and on the body could be a good approximation.     

Aerodynamics of a bio-inspired flexible flapping-wing micro air vehicle

MAVs (micro air vehicles) with a maximal dimension of 15 cm and nominal flight speeds of around 10 m s?1, operate in a Reynolds number regime of 10? or lower, in which most natural flyers including insects, bats and birds fly. Furthermore, due to their light weight and low flight speed, the MAVs' flight characteristics are substantially affected by environmental factors such as wind gust. Like natural flyers, the wing structures of MAVs are often flexible and tend to deform during flight. Consequently, the aero/fluid and structural dynamics of these flyers are closely linked to each other, making the entire flight vehicle difficult to analyze. We have recently developed a hummingbird-inspired, flapping flexible wing MAV with a weight of 2.4-3.0 g and a wingspan of 10-12 cm. In this study, we carry out an integrated study of the flexible wing aerodynamics of this flapping MAV by combining an in-house computational fluid dynamic (CFD) method and wind tunnel experiments. A CFD model that has a realistic wing planform and can mimic realistic flexible wing kinematics is established, which provides a quantitative prediction of unsteady aerodynamics of the four-winged MAV in terms of vortex and wake structures and their relationship with aerodynamic force generation. Wind tunnel experiments further confirm the effectiveness of the clap and fling mechanism employed in this bio-inspired MAV as well as the importance of the wing flexibility in designing small flapping-wing MAVs.     

Two-and Three-Dimensional Simulations of Beetle Hind Wing Flapping during Free Forward Flight

Aerodynamic characteristic of the beetle, Trypoxylus dichotomus, which has a pair of elytra (forewings) and hind wings, is numerically investigated. Based on the experimental results of wing kinematics, two-dimensional (2D) and three-dimensional (3D) computational fluid dynamic simulations were carried out to reveal aerodynamic performance of the hind wing. The roles of the spiral Leading Edge Vortex (LEV) and the spanwise flow were clarified by comparing 2D and 3D simulations. Mainly due to pitching down of chord line during downstroke in highly inclined stroke plane, relatively high averaged thrust was produced in the free forward flight of the beetle. The effects of the local corrugation and the camber variation were also investigated for the beetle's hind wings. Our results show that the camber variation plays a significant role in improving both lift and thrust in the flapping. On the other hand, the local corrugation pattern has no significant effect on the aerodynamic force due to large angle of attack during flapping.     

The Role of Soft Vein Joints in Dragonfly Flight

Dragonflies are excellent flyers among insects and their flight ability is closely related to the architecture and material properties of their wings.The veins are main structure components of a dragonfly wing,which are found to be connected by resilin with high elasticity at some joints.A three-dimensional (3D) finite element model of dragonfly wing considering the soft vein joints is developed,with some simplifications.Passive deformation under aerodynamic loads and active flapping motion of the wing are both studied.The functions of soft vein joints in dragonfly flight are concluded.In passive deformation,the chordwise flexibility is improved by soft vein joints and the wing is cambered under loads,increasing the action area with air.In active flapping,the wing rigidity in spanwise direction is maintained to achieve the required amplitude.As a result,both the passive deformation and the active control of flapping work well in dragonfly flight.The present study may also inspire the design of biomimetic Flapping Micro Air Vehicles (FMAVs).     

Analysis of Maneuvering Flight of an Insect

Wing motion of a dragonfly in the maneuvering flight, which was measured by Wang et al. was investigated. Equations of motion for a maneuvering flight of an insect were derived. These equations were applied for analyzing the maneuvering flight. Inertial forces and moments acting on a body and wings were estimated by using these equations and the measured motions of the body and the wings. The results indicated the following characteristics of this flight: ( 1 ) The phase difference in flapping motion between the two fore wings and two hind wings, and the phase difference between the flapping motion and the feathering motion of the four wings are equal to those in a steady forward flight with the maximum efficiency. (2)The camber change and the feathering motion were mainly controlled by muscles at the wing bases.     

Modulation of leading edge vorticity and aerodynamic forces in flexible flapping wings

In diverse biological flight systems, the leading edge vortex has been implicated as a flow feature of key importance in the generation of flight forces. Unlike fixed wings, flapping wings can translate at higher angles of attack without stalling because their leading edge vorticity is more stable than the corresponding fixed wing case. Hence, the leading edge vorticity has often been suggested as the primary determinant of the high forces generated by flapping wings. To test this hypothesis, it is necessary to modulate the size and strength of the leading edge vorticity independently of the gross kinematics while simultaneously monitoring the forces generated by the wing. In a recent study, we observed that forces generated by wings with flexible trailing margins showed a direct dependence on the flexural stiffness of the wing. Based on that study, we hypothesized that trailing edge flexion directly influences leading edge vorticity, and thereby the magnitude of aerodynamic forces on the flexible flapping wings. To test this hypothesis, we visualized the flows on wings of varying flexural stiffness using a custom 2D digital particle image velocimetry system, while simultaneously monitoring the magnitude of the aerodynamic forces. Our data show that as flexion decreases, the magnitude of the leading edge vorticity increases and enhances aerodynamic forces, thus confirming that the leading edge vortex is indeed a key feature for aerodynamic force generation in flapping flight. The data shown here thus support the hypothesis that camber influences instantaneous aerodynamic forces through modulation of the leading edge vorticity.     

Flight of the dragonflies and damselflies

This work is a synthesis of our current understanding of the mechanics, aerodynamics and visually mediated control of dragonfly and damselfly flight, with the addition of new experimental and computational data in several key areas. These are: the diversity of dragonfly wing morphologies, the aerodynamics of gliding flight, force generation in flapping flight, aerodynamic efficiency, comparative flight performance and pursuit strategies during predatory and territorial flights. New data are set in context by brief reviews covering anatomy at several scales, insect aerodynamics, neuromechanics and behaviour. We achieve a new perspective by means of a diverse range of techniques, including laser-line mapping of wing topographies, computational fluid dynamics simulations of finely detailed wing geometries, quantitative imaging using particle image velocimetry of on-wing and wake flow patterns, classical aerodynamic theory, photography in the field, infrared motion capture and multi-camera optical tracking of free flight trajectories in laboratory environments. Our comprehensive approach enables a novel synthesis of datasets and subfields that integrates many aspects of flight from the neurobiology of the compound eye, through the aeromechanical interface with the surrounding fluid, to flight performance under cruising and higher-energy behavioural modes.This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.     

Structural dynamics and aerodynamics measurements of biologically inspired flexible flapping wings

Flapping wing flight as seen in hummingbirds and insects poses an interesting unsteady aerodynamic problem: coupling of wing kinematics, structural dynamics and aerodynamics. There have been numerous studies on the kinematics and aerodynamics in both experimental and computational cases with both natural and artificial wings. These studies tend to ignore wing flexibility; however, observation in nature affirms that passive wing deformation is predominant and may be crucial to the aerodynamic performance. This paper presents a multidisciplinary experimental endeavor in correlating a flapping micro air vehicle wing's aeroelasticity and thrust production, by quantifying and comparing overall thrust, structural deformation and airflow of six pairs of hummingbird-shaped membrane wings of different properties. The results show that for a specific spatial distribution of flexibility, there is an effective frequency range in thrust production. The wing deformation at the thrust-productive frequencies indicates the importance of flexibility: both bending and twisting motion can interact with aerodynamic loads to enhance wing performance under certain conditions, such as the deformation phase and amplitude. By measuring structural deformations under the same aerodynamic conditions, beneficial effects of passive wing deformation can be observed from the visualized airflow and averaged thrust. The measurements and their presentation enable observation and understanding of the required structural properties for a thrust effective flapping wing. The intended passive responses of the different wings follow a particular pattern in correlation to their aerodynamic performance. Consequently, both the experimental technique and data analysis method can lead to further studies to determine the design principles for micro air vehicle flapping wings.     

枯叶蛱蝶扑翼飞行的力学分析

模仿昆虫扑翼飞行的飞行器具有重量轻、质量小、噪音低、效率高、隐蔽性好等优点,在军用、民用领域被广泛地关注与应用.枯叶蛱蝶是典型的扑翼昆虫,在连续上升飞行过程中会出现停顿和跃升的现象.为了研究停顿和跃升现象的产生原因,对枯叶蛱蝶的翅型和扑翼行为进行了力学分析.通过测量鳞翅结构参数,记录飞行行为,运用能量守恒与动量守恒原理,考虑生物能的作用,视空气为不可压缩颗粒,建立了数学模型模拟枯叶蛱蝶飞行情况.结果表明,扑翼行为通过改变飞行动力的动量和分力大小来影响枯叶蛱蝶的飞行轨迹,鳞翅形状则通过改变飞行动力的大小来影响枯叶蛱蝶的飞行轨迹,扑翼行为导致停顿和跃升现象的产生.本文为设计扑翼型飞行器提供了力学仿生学基础与生物学模型,为进一步设计出更优化的仿生飞行器提供科学依据.     

Pitching Moment Generation in an Insect-Mimicking Flapping-Wing System

Unlike birds, insects lack control surfaces at the tail and hence most insects modify their wing kinematics to produce control forces or moments while flapping their wings. Change of the flapping angle range is one of the ways to modify wing kinematics, resulting in relocation of the mean Aerodynamic force Center （mean AC） and finally creating control moments. In an attempt to mimic this feature, we developed a flapping-wing system that generates a desired pitching moment during flap- ping-wing motion. The system comprises a flapping mechanism that creates a large and symmetric flapping motion in a pair of wings, a flapping angle change mechanism that modifies the flapping angle range, artificial wings, and a power source. From the measured wing kinematics, we have found that the flapping-wing system can properly modify the flapping angle ranges. The measured pitching moments show that the flapping-wing system generates a pitching moment in a desired direction by shifting the flapping angle range. We also demonstrated that the system can in practice change the longitudinal attitude by generating a nonzero pitching moment.     

Falling with Style: Bats Perform Complex Aerial Rotations by Adjusting Wing Inertia

The remarkable maneuverability of flying animals results from precise movements of their highly specialized wings. Bats have evolved an impressive capacity to control their flight, in large part due to their ability to modulate wing shape, area, and angle of attack through many independently controlled joints. Bat wings, however, also contain many bones and relatively large muscles, and thus the ratio of bats’ wing mass to their body mass is larger than it is for all other extant flyers. Although the inertia in bat wings would typically be associated with decreased aerial maneuverability, we show that bat maneuvers challenge this notion. We use a model-based tracking algorithm to measure the wing and body kinematics of bats performing complex aerial rotations. Using a minimal model of a bat with only six degrees of kinematic freedom, we show that bats can perform body rolls by selectively retracting one wing during the flapping cycle. We also show that this maneuver does not rely on aerodynamic forces, and furthermore that a fruit fly, with nearly massless wings, would not exhibit this effect. Similar results are shown for a pitching maneuver. Finally, we combine high-resolution kinematics of wing and body movements during landing and falling maneuvers with a 52-degree-of-freedom dynamical model of a bat to show that modulation of wing inertia plays the dominant role in reorienting the bat during landing and falling maneuvers, with minimal contribution from aerodynamic forces. Bats can, therefore, use their wings as multifunctional organs, capable of sophisticated aerodynamic and inertial dynamics not previously observed in other flying animals. This may also have implications for the control of aerial robotic vehicles.     

Animal flight dynamics II. Longitudinal stability in flapping flight

Stability is essential to flying and is usually assumed to be especially problematic in flapping flight. If so, problems of stability may have presented a particular hurdle to the evolution of flapping flight. In spite of this, the stability of flapping flight has never been properly analysed. Here we use quasi-static and blade element approaches to analyse the stability provided by a flapping wing. By using reduced order approximations to the natural modes of motion, we show that wing beat frequencies are generally high enough compared to the natural frequencies of motion for a quasi-static approach to be valid as a first approximation. Contrary to expectations, we find that there is noting inherently destabilizing about flapping: beating the wings faster simply amplifies any existing stability or instability, and flapping can even enhance stability compared to gliding at the same air speed. This suggests that aerodynamic stability may not have been a particular hurdle in the evolution of flapping flight. Hovering animals, like hovering helicopters, are predicted to possess neutral static stability. Flapping animals, like fixed wing aircraft, are predicted to be stable in forward flight if the mean flight force acts above and/or behind the centre of gravity. In this case, the downstroke will always be stabilizing. The stabilizing contribution may be diminished by an active upstroke with a low advance ratio and more horizontal stroke plane; other forms of the upstroke may make a small positive contribution to stability. An active upstroke could, therefore, be used to lower stability and enhance manoeuvrability. Translatory mechanisms of unsteady lift production are predicted to amplify the stability predicted by a quasi-static analysis. Non-translatory mechanisms will make little or no contribution to stability. This may be one reason why flies, and other animals which rely upon non-translatory aerodynamic mechanisms, often appear inherently unstable.