Coordination of wingbeat and respiration in birds. II. "Fictive" flight.

To determine whether an interaction between central respiratory and locomotor networks may be involved in the observed coordination of wingbeat and respiratory rhythms during free flight in birds, we examined the relationship between wingbeat and respiratory activity in decerebrate Canada geese and Pekin ducks before and after paralysis. Locomotor activity was induced through electrical stimulation of brain stem locomotor regions. Respiratory frequency (fv) was monitored via pneumotachography and intercostal electromyogram recordings before paralysis and via intercostal and cranial nerve IX electroneurogram recordings after paralysis. Wingbeat frequency (fW) was monitored using pectoralis major electromyogram recordings before, and electroneurogram recordings after, paralysis. Respiratory and cardiovascular responses of decerebrate birds during active (nonparalyzed) and "fictive" (paralyzed) wing activity were qualitatively similar to those of a variety of vertebrate species to exercise. As seen during free flight, wingbeat and respiratory rhythms were always coordinated during electrically induced wing activity. Before paralysis during active wing flapping, coupling ratios (fW/fv) of 1:1, 2:1, 3:1, and 4:1 (wingbeats per breath) were observed. After paralysis, fW and fv remained coupled; however, 1:1 coordination predominated. All animals tested (n = 9) showed 1:1 coordination. Two animals also showed brief periods of 2:1 coupling. It is clear that locomotor and respiratory networks interact on a central level to produce a synchronized output. The observation that the coordination between fW and fv differs in paralyzed and nonparalyzed birds suggests that peripheral feedback is involved in the modulation of a centrally derived coordination.     

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.     

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.     

Morphology, Velocity, and Intermittent Flight in Birds

Body size, pectoralis composition, aspect ratio of the wing,and forward speed affect the use of intermittent flight in birds.During intermittent non-flapping phases, birds extend theirwings and glide or flex their wings and bound. The pectoralismuscle is active during glides but not during bounds; activityin other primary flight muscles is variable. Mechanical power,altitude, and velocity vary among wingbeats in flapping phases;associated with this variation are changes in neuromuscularrecruitment, wingbeat frequency, amplitude, and gait. Speciesof intermediate body mass (35–158 g) tend to flap-glideat slower speeds and flap-bound at faster speeds, regardlessof the aspect ratio of their wings. Such behavior may reducemechanical power output relative to continuous flapping. Smallerspecies (<20 g) with wings of low aspect ratio may flap-boundat all speeds, yet existing models do not predict an aerodynamicadvantage for the flight style at slow speeds. The behaviorof these species appears to be due to wing shape rather thanpectoralis physiology. As body size increases among species,percent time spent flapping increases, and birds much largerthan 300 g do not flap-bound. This pattern may be explainedby adverse scaling of mass-specific power or lift per unit poweroutput available from flight muscles. The size limit for theability to bound intermittently may be offset somewhat by thescaling of pectoralis composition. The percentage of time spentflapping during intermittent flight also varies according toflight speed.     

A wing-assisted running robot and implications for avian flight evolution

DASH+Wings is a small hexapedal winged robot that uses flapping wings to increase its locomotion capabilities. To examine the effects of flapping wings, multiple experimental controls for the same locomotor platform are provided by wing removal, by the use of inertially similar lateral spars, and by passive rather than actively flapping wings. We used accelerometers and high-speed cameras to measure the performance of this hybrid robot in both horizontal running and while ascending inclines. To examine consequences of wing flapping for aerial performance, we measured lift and drag forces on the robot at constant airspeeds and body orientations in a wind tunnel; we also determined equilibrium glide performance in free flight. The addition of flapping wings increased the maximum horizontal running speed from 0.68 to 1.29 m s?1, and also increased the maximum incline angle of ascent from 5.6° to 16.9°. Free flight measurements show a decrease of 10.3° in equilibrium glide slope between the flapping and gliding robot. In air, flapping improved the mean lift:drag ratio of the robot compared to gliding at all measured body orientations and airspeeds. Low-amplitude wing flapping thus provides advantages in both cursorial and aerial locomotion. We note that current support for the diverse theories of avian flight origins derive from limited fossil evidence, the adult behavior of extant flying birds, and developmental stages of already volant taxa. By contrast, addition of wings to a cursorial robot allows direct evaluation of the consequences of wing flapping for locomotor performance in both running and flying.     

Strouhal number for flying and swimming in rhinoceros auklets Cerorhinca monocerata

Alcids propel themselves by flapping wings in air and water that have vastly different densities. We hypothesized that alcids change wing kinematics and maintain Strouhal numbers (St = fA/U, where f is wingbeat frequency, A is the wingbeat amplitude, and U is forward speed) within a certain range, to achieve efficient locomotion during both flying and swimming. We used acceleration and GPS loggers to measure the wingbeat frequency and forward speed of free‐ranging rhinoceros auklets Cerorhinca monocerata during both flying and swimming. We also measured wingbeat amplitude from video footage taken in the wild. On average, wingbeat frequency, forward speed, and wingbeat amplitude were 8.9 Hz, 15.3 m s^?1, and 0.39 m, respectively, during flying, and 2.6 Hz, 1.3 m s^?1, and 0.18 m, respectively, during swimming. The smaller wingbeat amplitude during swimming was achieved by partially folding the wings, while maintaining the dorso‐ventral wingbeat angle. Mean St was 0.23 during flying and 0.36 during swimming. The higher St value for swimming might be related to the higher thrust force required for propulsion in water. Our results suggest that rhinoceros auklets maintain St for both flying and swimming within the range (0.2–0.4) that propulsive efficiency is known to be high and St in both flying specialists and swimming specialists are known to converge.     

Measurement on Camber Deformation of Wings of Free-flying Dragonflies and Beating-flying Dragonflies

1　IntroductionNumerouskinematicparameters,includingwing beatfrequency ,wingorientation ,andbothspan andchord wisedeformation ,arerelevanttotheaerodynam icanalysisofinsectflight[1,2 ] .Althoughnearlyalltherecentstudiesofinsectflightaerodynamics[3,4 ] haveidentifiedthatthemechanismsrequireflowseparationattheleadingedge ,andcamberisnotexpectedtohaveanysignificantinfluenceonthemagnitudeoftheforcecoefficient,someinsects ,suchasdragonfliesandbut terflies,frequently glideusinglowanglesofattack ,lead…     

Hovering and intermittent flight in birds

Two styles of bird locomotion, hovering and intermittent flight, have great potential to inform future development of autonomous flying vehicles. Hummingbirds are the smallest flying vertebrates, and they are the only birds that can sustain hovering. Their ability to hover is due to their small size, high wingbeat frequency, relatively large margin of mass-specific power available for flight and a suite of anatomical features that include proportionally massive major flight muscles (pectoralis and supracoracoideus) and wing anatomy that enables them to leave their wings extended yet turned over (supinated) during upstroke so that they can generate lift to support their weight. Hummingbirds generate three times more lift during downstroke compared with upstroke, with the disparity due to wing twist during upstroke. Much like insects, hummingbirds exploit unsteady mechanisms during hovering including delayed stall during wing translation that is manifest as a leading-edge vortex (LEV) on the wing and rotational circulation at the end of each half stroke. Intermittent flight is common in small- and medium-sized birds and consists of pauses during which the wings are flexed (bound) or extended (glide). Flap-bounding appears to be an energy-saving style when flying relatively fast, with the production of lift by the body and tail critical to this saving. Flap-gliding is thought to be less costly than continuous flapping during flight at most speeds. Some species are known to shift from flap-gliding at slow speeds to flap-bounding at fast speeds, but there is an upper size limit for the ability to bound (~0.3 kg) and small birds with rounded wings do not use intermittent glides.     

Wingbeat frequency and flap-pause ratio during natural migratory flight in thrushes

Powered flapping flight has evolved independently in many differenttaxa. For flapping fliers, wingbeat parameters such as frequencyand amplitude are the primary determinants of these animals’energetic expenditure during flight. Here we present data onwingbeat frequency and amplitude for three New World thrushspecies during 15 entire nocturnal migratory flights over theMidwestern United States. Using continuous (non-pulsing) radiotransmitters, we were able to measure wingbeat frequency andrelative amplitude of wingbeats as well as the characteristicsof flap-pauses. Contrary to previous telemetric findings, allof the individuals we followed used both flapping-only and flap-pauseflight. During migratory flights, wingbeat frequency, effectivewingbeat frequency, and amplitude were highest during initialascent. Effective wingbeat frequency and amplitude were lowestduring final descent. We show that identification of speciesbased solely on characteristics of the wingbeat e.g., duringradar studies, can be difficult because variables such as wingbeatfrequency and amplitude, wingbeat pausing, and pattern of beatsand pauses vary between individuals of the same species andeven within individual flights. We also show that observed wingbeatfrequencies were lower than those predicted by theoretical models.We speculate that this may be because theoretical predictionsare generally based on (1) data from larger birds and (2) datafrom diurnal flights. We found that diurnal wingbeat frequenciesof thrushes were generally higher than were those during nocturnalmigratory flight. Finally, we suggest that rather than remainingat a single altitude during flight or climbing slightly as theoreticalmodels predict, thrushes often moved up and down in the aircolumn, perhaps searching for favorable atmospheric conditions.     

Characteristics of a Beetle's Free Flight and a Flapping-Wing System that Mimics Beetle Flight

In this work, we first present a method to experimentally capture the free flight of a beetle (Allomyrina dichotoma), which is not an active flyer. The beetle is suspended in the air by a hanger to induce the free flight. This flight is filmed using two high-speed cameras. The high speed images are then examined to obtain flapping angle, flapping frequency, and wing rotation of the hind wing. The acquired data of beetle free flight are used to design a motor-driven flapper that can approximately mimic the beetle in terms of size, flapping frequency and wing kinematics. The flapper can create a large flapping angle over 140° with a large passive wing rotation angle. Even though the flapping frequency of the flapper is not high enough compared to that of a real beetle due to the limited motor torque, the flapper could produce positive average vertical force. This work will provide important experience for future development of a beetle-mimicking Flapping-Wing Micro Air Vehicle (FWMAV).     

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).     

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.     

Effect of outer wing separation on lift and thrust generation in a flapping wing system

We explore the implementation of wing feather separation and lead-lagging motion to a flapping wing. A biomimetic flapping wing system with separated outer wings is designed and demonstrated. The artificial wing feather separation is implemented in the biomimetic wing by dividing the wing into inner and outer wings. The features of flapping, lead-lagging, and outer wing separation of the flapping wing system are captured by a high-speed camera for evaluation. The performance of the flapping wing system with separated outer wings is compared to that of a flapping wing system with closed outer wings in terms of forward force and downward force production. For a low flapping frequency ranging from 2.47 to 3.90 Hz, the proposed biomimetic flapping wing system shows a higher thrust and lift generation capability as demonstrated by a series of experiments. For 1.6 V application (lower frequency operation), the flapping wing system with separated wings could generate about 56% higher forward force and about 61% less downward force compared to that with closed wings, which is enough to demonstrate larger thrust and lift production capability of the separated outer wings. The experiments show that the outer parts of the separated wings are able to deform, resulting in a smaller amount of drag production during the upstroke, while still producing relatively greater lift and thrust during the downstroke.     

Dipteran flight motor pattern: Invariabilities and changes during postlarval development

For Calliphora the wingbeat frequency and the underlying motoneuronal activity were recorded during adult life. Wingbeat frequency increases during the ten days following last molt. The activity of motoneurons serving four selected flight muscles (nonfibrillar and fibrillar ones) also increases with age. The motoneuronal activity of young and old flies was analyzed statistically (serial and cross-correlograms, latency and phase histograms). In addition, several wing manipulations were carried out to evaluate the significance of sensory feedback on pattern generation during maturation. These ontogenetic studies suggest a centrally generated motor pattern that (1) is essentially complete with the molt to adulthood, (2) shows a progressive increase in intrinsic activity, and (3) is modulated by sensory feedback from the wing region by the same amount irrespective of age. Similarities in the postlarval development of the flight pattern of neurogenic and myogenic flyers are discussed.     

Upstroke wing flexion and the inertial cost of bat flight

Flying vertebrates change the shapes of their wings during the upstroke, thereby decreasing wing surface area and bringing the wings closer to the body than during downstroke. These, and other wing deformations, might reduce the inertial cost of the upstroke compared with what it would be if the wings remained fully extended. However, wing deformations themselves entail energetic costs that could exceed any inertial energy savings. Using a model that incorporates detailed three-dimensional wing kinematics, we estimated the inertial cost of flapping flight for six bat species spanning a 40-fold range of body masses. We estimate that folding and unfolding comprises roughly 44 per cent of the inertial cost, but that the total inertial cost is only approximately 65 per cent of what it would be if the wing remained extended and rigid throughout the wingbeat cycle. Folding and unfolding occurred mostly during the upstroke; hence, our model suggests inertial cost of the upstroke is not less than that of downstroke. The cost of accelerating the metacarpals and phalanges accounted for around 44 per cent of inertial costs, although those elements constitute only 12 per cent of wing weight. This highlights the energetic benefit afforded to bats by the decreased mineralization of the distal wing bones.     

Artificial Cambered-Wing for a Beetle-Mimicking Flapper

In this paper, we have attempted to improve the aerodynamic force generation ability of an artificial wing by implementing initial wing camber in the flexible artificial wing. This initial camber is used to create passive wing camber during flapping motion. We modified original artificial wing by removing many minor vein structures in the wing and then placed the initial camber between two major veins. Stiffness measurements for the original artificial wing and the present wing with initial camber were conducted to compare the stiffnesses of the two artificial wings, and the similarities of the two wings are discussed. A flapping test was carried out using a previously-built flapper that can flap at higher than 25 Hz flapping frequency to verify the wing camber effect. Finally, a performance comparison between uncambered- and cambered-wings was also undertaken based on observations using a high-speed camera and force measurements from wired-flight tests and swing tests. The comparison showed that the cambered-wing could produce about 10% higher thrust than the uncambered-wing.     

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.     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).     

Wing structure and neural encoding jointly determine sensing strategies in insect flight

Animals rely on sensory feedback to generate accurate, reliable movements. In many flying insects, strain-sensitive neurons on the wings provide rapid feedback that is critical for stable flight control. While the impacts of wing structure on aerodynamic performance have been widely studied, the impacts of wing structure on sensing are largely unexplored. In this paper, we show how the structural properties of the wing and encoding by mechanosensory neurons interact to jointly determine optimal sensing strategies and performance. Specifically, we examine how neural sensors can be placed effectively on a flapping wing to detect body rotation about different axes, using a computational wing model with varying flexural stiffness. A small set of mechanosensors, conveying strain information at key locations with a single action potential per wingbeat, enable accurate detection of body rotation. Optimal sensor locations are concentrated at either the wing base or the wing tip, and they transition sharply as a function of both wing stiffness and neural threshold. Moreover, the sensing strategy and performance is robust to both external disturbances and sensor loss. Typically, only five sensors are needed to achieve near-peak accuracy, with a single sensor often providing accuracy well above chance. Our results show that small-amplitude, dynamic signals can be extracted efficiently with spatially and temporally sparse sensors in the context of flight. The demonstrated interaction of wing structure and neural encoding properties points to the importance of understanding each in the context of their joint evolution.     

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.