Dynamics of in vivo power output and efficiency of Nasonia asynchronous flight muscle

By simultaneously measuring aerodynamic performance, wing kinematics, and metabolic activity, we have estimated the in vivo limits of mechanical power production and efficiency of the asynchronous flight muscle (IFM) in three species of ectoparasitoid wasps genus Nasonia (N. giraulti, N. longicornis, and N. vitripennis). The 0.6 mg animals were flown under tethered flight conditions in a flight simulator that allowed modulation of power production by employing an open-loop visual stimulation technique. At maximum locomotor capacity, flight muscles of Nasonia are capable to sustain 72.2 +/- 18.3 W kg(-1) muscle mechanical power at a chemo-mechanical conversion efficiency of approximately 9.8 +/- 0.9%. Within the working range of the locomotor system, profile power requirement for flight dominates induced power requirement suggesting that the cost to overcome wing drag places the primary limit on overall flight performance. Since inertial power is only approximately 25% of the sum of induced and profile power requirements, Nasonia spp. may not benefit from elastic energy storage during wing deceleration phases. A comparison between wing size-polymorphic males revealed that wing size reduction is accompanied by a decrease in total flight muscle volume, muscle mass-specific mechanical power production, and total flight efficiency. In animals with small wings maximum total flight efficiency is below 0.5%. The aerodynamic and power estimates reported here for Nasonia are comparable to values reported previously for the fruit fly Drosophila flying under similar experimental conditions, while muscle efficiency of the tiny wasp is more at the lower end of values published for various other insects.     

Aerodynamic aspects of formation flight in birds

In formation flight each wing flies in an upwash field generated by all other wings of the formation. This leads to a reduction in flight power demand for each wing as well as for the whole formation.Methods of theoretical aerodynamics are used to calculate the flight power reduction for arbitrarily shaped flight formations with any number of birds. These methods are applied to homogeneous and inhomogeneous flight formations in which birds of the same kind or birds of different span, aspect ratio and weight may be present.The total flight power reduction of the whole formation strongly depends on the lateral distance of the wings. A longitudinal displacement of the wings in flight direction has no influence on the total flight power reduction but only on their distribution on the involved individuals. The local flight power reduction is highest in the inner parts of the formation and decreases towards the apex and towards the side edges of the formation. Small and light individuals are automatically favoured by larger and heavier birds. It is shown that some minor portion of twist is necessary to fly in a formation without a rolling moment. In addition it turns out that the optimum flight speed of a formation is slightly lower than the optimum flight speed of single individuals.     

Wing morphology and flight development in the short-nosed fruit bat Cynopterus sphinx

Postnatal changes in wing morphology, flight development and aerodynamics were studied in captive free-flying short-nosed fruit bats, Cynopterus sphinx. Pups were reluctant to move until 25 days of age and started fluttering at the mean age of 40 days. The wingspan and wing area increased linearly until 45 days of age by which time the young bats exhibited clumsy flight with gentle turns. At birth, C. sphinx had less-developed handwings compared to armwings; however, the handwing developed faster than the armwing during the postnatal period. Young bats achieved sustained flight at 55 days of age. Wing loading decreased linearly until 35 days of age and thereafter increased to a maximum of 12.82 Nm(-2) at 125 days of age. The logistic equation fitted the postnatal changes in wingspan and wing area better than the Gompertz and von Bertalanffy equations. The predicted minimum power speed (V(mp)) and maximum range speed (V(mr)) decreased until the onset of flight and thereafter the V(mp) and V(mr) increased linearly and approached 96.2% and 96.4%, respectively, of the speed of postpartum females at the age of 125 days. The requirement of minimum flight power (P(mp)) and maximum range power (P(mr)) increased until 85 days of age and thereafter stabilised. The minimum theoretical radius of banked turn (r(min)) decreased until 35 days of age and thereafter increased linearly and attained 86.5% of the r(min) of postpartum females at the age of 125 days.     

Wing kinematics of avian flight across speeds

To test whether wing shape affects the kinematics of wing motion during bird flight, we recorded high-speed video (250 Hz) of four species flying in a variable-speed wind tunnel. The birds flew at intervals of 2 m s⁻¹, ranging from 1 m s⁻¹ up to their respective maximum flight speed, which varied from 14 to 17 m s⁻¹ depending on the species. Kinematic data obtained from two synchronized, high-speed video cameras were analyzed using 3D reconstruction. Three species with relatively pointed, high-aspect ratio wings changed wingbeat styles according to flight speed (budgerigar, Melopsittacus undulatus ; cockatiel, Nymphicus hollandicus ; ringed turtle dove, Streptopelia risoria ). These species used a wing-tip reversal upstroke, characterized by supination of the distal wing at mid-upstroke, at equivalent airspeeds ≤7 to 9 m s⁻¹. In faster flight, they used a swept-wing upstroke, without distal wing supination. At mid-upstroke at any speed, wingspan in these species was greater than wrist span. In contrast, at all steady flight speeds, the black-billed magpie Pica hudsonia with relatively broad, low-aspect ratio wings, used a flexed-wing, feathered upstroke in which wrist spans were equal to or greater than wingspans. Our results demonstrate that wing kinematics vary gradually as a function of flight speed, and that the patterns of variation are strongly influenced by external wing shape.     

Biomechanics and biomimetics in insect-inspired flight systems

Insect- and bird-size drones—micro air vehicles (MAV) that can perform autonomous flight in natural and man-made environments are now an active and well-integrated research area. MAVs normally operate at a low speed in a Reynolds number regime of 10⁴–10⁵ or lower, in which most flying animals of insects, birds and bats fly, and encounter unconventional challenges in generating sufficient aerodynamic forces to stay airborne and in controlling flight autonomy to achieve complex manoeuvres. Flying insects that power and control flight by flapping wings are capable of sophisticated aerodynamic force production and precise, agile manoeuvring, through an integrated system consisting of wings to generate aerodynamic force, muscles to move the wings and a control system to modulate power output from the muscles. In this article, we give a selective review on the state of the art of biomechanics in bioinspired flight systems in terms of flapping and flexible wing aerodynamics, flight dynamics and stability, passive and active mechanisms in stabilization and control, as well as flapping flight in unsteady environments. We further highlight recent advances in biomimetics of flapping-wing MAVs with a specific focus on insect-inspired wing design and fabrication, as well as sensing systems.This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.     

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.     

The Function of Dipteran Flight Muscle

The flight muscles of flies are separated into two physiologically, anatomically, and functionally distinct classes: power muscles and control muscles. The large indirect power muscles sustain the high level of mechanical energy required to flap the wings up and down during flight. The contractions in the asynchronous power muscles are initiated by stretch, and their slow presynaptic motor drive serves only to maintain a tonic level of cytosolic calcium. Although providing the mechanical energy for flight, the power muscles are not directly attached to the wings. Instead, their mechanical energy is transmitted to the base of the wings through the complex linkage system of the wing hinge. In contrast, the small control muscles insert directly onto the skeletal elements at the base of the wing. Through their mechanical effects on the hinge, the control muscles act collectively as a transmission system that determines how the mechanical energy produced by the power muscles is transformed into wing motion. The control muscles are activated by motor spikes in the conventional one-for-one fashion. Thus, although the control muscles can generate little mechanical power, they provide the means by which the nervous system can rapidly alter wing kinematics during sophisticated aerial maneuvers.     

Genetic Variability of Flight Metabolism in DROSOPHILA MELANOGASTER. I. Characterization of Power Output during Tethered Flight

The mechanical power imparted to the wings during tethered flight of Drosophila melanogaster is estimated from wing-beat frequency, wing-stroke amplitude and various aspects of wing morphology by applying the steady-state aerodynamics model of insect flight developed by Weis-Fogh (1972, 1973). Wing-beat frequency, the major determinant of power output, is highly correlated with the rate of oxygen consumption. Estimates of power generated during flight should closely reflect rates of ATP production in the flight muscles, since flies do not acquire an oxygen debt or accumulate ATP during flight. In an experiment using 21 chromosome 2 substitution lines, lines were a significant source of variation for all flight parameters measured. Broadsense heritabilities ranged from 0.16 for wing-stroke amplitude to 0.44 for inertial power. The variation among lines is not explained by variation in total body size (i.e., live weight). Line differences in flight parameters are robust with respect to age, ambient temperature and duration of flight. These results indicate that characterization of the power output during tethered flight will provide a sensitive experimental system for detecting the physiological effects of variation in the structure or quantity of the enzymes involved in flight metabolism.     

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.     

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.     

Thermal physiological ecology of Colias butterflies in flight

Summary As a comparison to the many studies of larger flying insects, we carried out an initial study of heat balance and thermal dependence of flight of a small butterfly (Colias) in a wind tunnel and in the wild.Unlike many larger, or facultatively endothermic insects, Colias do not regulate heat loss by altering hemolymph circulation between thorax and abdomen as a function of body temperature. During flight, thermal excess of the abdomen above ambient temperature is weakly but consistently coupled to that of the thorax. Total heat loss is best expressed as the sum of heat loss from the head and thorex combined plus heat loss from the abdomen because the whole body is not isothermal. Convective cooling is a simple linear function of the square root of air speed from 0.2 to 2.0 m/s in the wind tunnel. Solar heat flux is the main source of heat gain in flight, just as it is the exclusive source for warmup at rest. The balance of heat gain from sunlight versus heat loss from convection and radiation does not appear to change by more than a few percent between the wings-closed basking posture and the variable opening of wings in flight, although several aspects require further study. Heat generation by action of the flight muscles is small (on the order of 100 m W/g tissue) compared to values reported for other strongly flying insects. Colias appears to have only very limited capacity to modulate flight performance. Wing beat frequency varies from 12–19 Hz depending on body mass, air speed, and thoracic temperature. At suboptimal flight temperatures, wing beat frequency increases significantly with thoracic temperature and body mass but is independent of air speed. Within the reported thermal optimum of 35–39°C, wing beat frequency is negatively dependent on air speed at values above 1.5 m/s, but independent of mass and body temperature. Flight preference of butterflies in the wind tunnel is for air speeds of 0.5–1.5 m/s, and no flight occurs at or above 2.5 m/s. Voluntary flight initiation in the wild occurs only at air speeds 1.4 m/s.In the field, Colias fly just above the vegetation at body temperatures of 1–2°C greater than when basking at the top of the vegetation. These measurements are consistent with our findings on low heat gain from muscular activity during flight. Basking temperatures of butterflies sheltered from the wind within the vegetation were 1–2°C greater than flight temperatures at vegetation height.     

Reliability assessment of ballistic jump squats and bench throws

The purpose of this investigation was to determine the test-retest reliability and coefficient of variation of 2 novel physical performance tests. Ten healthy men (22.0 +/- 3.0 years, 87.0 +/- 8.0 kg, 20.0 +/- 5.0% body fat) performed 30 continuous and dynamic jump squats (JS) and bench throws (BT) on 4 separate occasions. The movements were performed under loaded conditions utilizing 30% of subject's predetermined 1 repetition maximum in the back squat and bench press. Mean power (MP; W), peak power (PP; W), mean velocity (MV; m.s(-1)), peak velocity (PV; m.s(-1)), and total work (TW; J) were assessed using a ballistic measurement system (Innervations Inc., Muncie, IN). Data were analyzed using repeated measures analysis of variance with Duncan's post hoc test when mean differences were p < or = 0.05. Intraclass correlation coefficient (ICC) and within-subject coefficient of variation (CV%) were also calculated. All values are presented as mean +/- SE. BT variables were statistically similar across the 4 sessions: MP (350.0 +/- 13.9 W), PP (431.4 +/- 18.5 W) MV (1.6 +/- 0.03 m.s(-1)), PV (2.0 +/- 0.03 m.s(-1)), and TW (199.1 +/- 7.2 J). For JS, session 3 PP (1,669.8 +/- 111.2 W) was significantly greater vs. sessions 1, 2, and 4 (1,601.2 +/- 58.4 W). Session 4 MP (1,403.2 +/- 88.6 W) and MV (1.9 +/- 0.1 m.s(-1)) for JS were significantly lower during sessions 1, 2, and 3 (MP: 1,479.4.5 +/- 44.8 W, MV: 2.0 +/- 0.05 m.s(-1)). TW (834.7 +/- 24.3 J) and PV (2.2 +/- 0.04 m.s(-1)) were statistically similar during all sessions for JS. The CVs ranged from 3.0 to 7.6% for the BT and 3.2 to 5.7% for the JS. ICCs for MP, PP, MV, PV, and TW were 0.92, 0.95, 0.94, 0.91, and 0.95, respectively, during BT. ICCs during JS for MP, PP, MV, PV, and TW were 0.96, 0.98, 0.94, 0.94, and 0.89, respectively. The results of the current study support the use of a 30 continuous and dynamic BT protocol as a reliable upper-body physical performance test, which can be administered with minimal practice. Slightly greater variability for JS was observed, although the test had high reliability.     

Scientific approach to the 1-h cycling world record: a case study.

The purpose of this study was to describe the physiological and aerodynamic characteristics and the preparation for a successful attempt to break the 1-h cycling world record. An elite professional road cyclist (30 yr, 188 cm, 81 kg) performed an incremental laboratory test to assess maximal power output (W(max)) and power output (W(OBLA)), estimated speed (V(OBLA)), and heart rate (HR(OBLA)) at the onset of blood lactate accumulation (OBLA). He also completed an incremental velodrome (cycling track) test (VT1), during which V(OBLAVT1) and HR(OBLAVT1) were measured and W(OBLAVT1) was estimated. W(max) was 572 W, W(OBLA) 505 W, V(OBLA) 52.88 km/h, and HR(OBLA) 183 beats/min. V(OBLAVT1), HR(OBLAVT1), and W(OBLAVT1) were 52.7 km/h, 180 beats/min, and 500.6 W, respectively. Drag coefficient and shape coefficient, measured in a wind tunnel, were 0. 244 and 0.65 m(2), respectively. The cyclist set a world record of 53,040 m, with an estimated average power output of 509.5 W. Based on direct laboratory data of the power vs. oxygen uptake relationship for this cyclist, this is slightly higher than the 497. 25 W corresponding to his oxygen uptake at OBLA (5.65 l/min). In conclusion, 1) the 1-h cycling world record is the result of the interaction between physiological and aerodynamic characteristics; and 2) performance in this event can be predicted using mathematical models that integrate the principal performance-determining variables.     

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.     

Skipping flights in Ypthima butterflies (Lepidoptera: Nymphalidae)

The skipping flight patterns of three species of Ypthima (Lepidoptera: Nymphalidae) were analyzed using high‐speed video recordings to clarify how wings move and how driving forces are produced. All three species showed a flight pattern that includes a pause that accounts for about 50% of a flap cycle when their wings completely close after each upstroke. The observed pause causes the “skipping” flight trajectory based on the clap–fling mechanism. Pause duration was correlated with upstroke wing motion, suggesting the contribution of the latter to a long pause duration. This is also supported by the temporal relationship between the wing and body motions. The aerodynamic power necessary for the pause flight was calculated for the three species.     

Alternative N-terminal regions of Drosophila myosin heavy chain tune muscle kinetics for optimal power output

We assessed the influence of alternative versions of a region near the N-terminus of Drosophila myosin heavy chain on muscle mechanical properties. Previously, we exchanged N-terminal regions (encoded by alternative exon 3s) between an embryonic (EMB) isoform and the indirect flight muscle isoform (IFI) of myosin, and demonstrated that it influences solution ATPase rates and in vitro actin sliding velocity. Because each myosin is expressed in Drosophila indirect flight muscle, in the absence of other myosin isoforms, this allows for muscle mechanical and whole organism locomotion assays. We found that exchanging the flight muscle specific exon 3 region into the embryonic isoform (EMB-3b) increased maximum power generation (P(max)) and optimal frequency of power generation (f(max)) threefold and twofold compared to fibers expressing EMB, whereas exchanging the embryonic exon 3 region into the flight muscle isoform (IFI-3a) decreased P(max) and f(max) to approximately 80% of IFI fiber values. Drosophila expressing IFI-3a exhibited a reduced wing beat frequency compared to flies expressing IFI, which optimized power generation from their kinetically slowed flight muscle. However, the slower wing beat frequency resulted in a substantial loss of aerodynamic power as manifest in decreased flight performance of IFI-3a compared to IFI. Thus the N-terminal region is important in tuning myosin kinetics to match muscle speed for optimal locomotory performance.     

Effect of exercise intensity on the slow component of oxygen uptake in decremental work load exercise.

The paper sought to determine the exercise intensity where the slow component of oxygen uptake (Vo(2)) first appears in decremental work load exercise (DLE). Incremental work load exercise (ILE) was performed with an increment rate of 15 watts (W) per minute. In DLE, power outputs were decreased by 15 W per minute, from 120 (DLE(120)), 160 (DLE(160)), 200 (DLE(200)) and 240 (DLE(240)) W, respectively. The slopes of Vo(2) against the power output were obtained in the lower section from 0 to 50 W in all DLEs, and in the upper section from 80 to 120 W in DLE(160) and from 100 to 150 W in DLE(200) and DLE(240). The power output at exhaustion in ILE was 274 +/- 20 W. The power output at the ventilatory threshold (VT) obtained in ILE was 167 +/- 22 W. The initial power output in DLE(160) was near the power output at VT. The slopes obtained in the upper sections were 11.4 +/- 0.9 ml x min(-1) x W(-1)1 in DLE(160), 12.8 +/- 0.8 ml x min(-1) x W(-1) in DLE(200), and 14.8 +/- 1.1 ml x min(-1) x W(-1) in DLE(240). The slope obtained in DLE(120) was 10.9 +/- 0.6 ml x min(-1). There were no differences in slope between the upper and lower sections in DLE(160) but there were significant differences in slopes between the upper and lower sections in DLE(200) and DLE(240). Thus, the slow component, which could be observed as a steeper slope in DLE, began to increase when the initial power output in DLE was near to VT.     

Morphometrical and front wing abrasion analysis of a Hungarian cotton bollworm <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> (Lepidoptera: Noctuidae) population

To determine the cotton bollworm migrating population rate in Hungary, we examined the weights and the front wing morphological feautures of trapped moths. We used sex pheromone traps to monitor field populations during the maize vegetation cycle period in 2008. We examined moths trapped at various times, and measured their body mass (m) and morphological features, namely the front wing quotient (fWQ = quotient of length of front wing/width of front wing), modified wing loading (WL = weight of moth/surface of front wing), and the relative thorax size (RTS = width of thorax/width of head). The data were analysed by Student t-test, anterior wing abrasion and darkness were analysed by a Adobe Photoshop 7.0 software. The Hungarian appearance of three cottom bollworm generations in 2008 was also observed. Based on the examined morphological features we found regularity in body mass, front wing quotient and modified wing loading changes during the flight period. The specimens trapped in the first and third part of the flight period had lower body mass, larger wing surface, longer wings and more favourable modified wing loading than the specimens trapped in the middle of the flight period. The abrasion and colour of the anterior wings of cotton bollworms were concordant to morphometric investigations. The abrasion in darker spots E1 and E3 clearly showed a more intensive usage of the wings in case of specimens trapped at the beginning and at the end of the flight period.     

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.     

Energetics of foraging and locomotion in the platypus Ornithorhynchus anatinus

We measured the energy requirements of platypuses foraging, diving and resting in a swim tank using flow-through respirometry. Also, walking metabolic rates were obtained from platypuses walking on a conventional treadmill. Energy requirements while foraging were found to depend on water temperature, body weight and dive duration and averaged 8.48 W kg(-1). Rates for subsurface swimming averaged 6.71 W kg(-1). Minimal cost of transport for subsurface swimming platypuses was 1.85 J N(-1)m(-1) at a speed of 0.4 m s(-1). Aerobic dive limit of the platypus amounted to 59 s. Metabolic rate of platypuses resting on the water surface was minimal with 3.91 W kg(-1) while minimal RMR on land was 2.08 W kg(-1). The metabolic rate for walking was 8.80 W kg(-1) and 10.56 W kg(-1) at speeds of 0.2 m s(-1) and 0.3 m s(-1), respectively. A formula was derived, which allows prediction of power requirements of platypuses in the wild from measurements of body weight, dive duration and water temperature. Platypuses were found to expend energy at only half the rate of semiaquatic eutherians of comparable body sizes during both walking and diving. However, costs of transport at optimal speed were in line with findings for eutherians. These patterns suggest that underwater locomotion of semiaquatic mammals have converged on very similar efficiencies despite differences in phylogeny and locomotor mode.