Hovering hummingbird wing aerodynamics during the annual cycle. II. Implications of wing feather moult.
ABSTRACT: Birds usually moult their feathers in a particular sequence which may incur aerodynamic, physiological and behavioural implications. Among birds, hummingbirds are unique species in their sustained hovering flight. Because hummingbirds frequently hover-feed, they must maintain sufficiently high flight capacities even when moulting their flight feathers. A hummingbird wing consists of 10 primary flight feathers whose absence during moult may strongly affect wing performance. Using dynamic similarity rules, we compared time-accurate aerodynamic loads and flow field measurements over several wing geometries that follow the natural feather moult sequence of Calypte anna, a common hummingbird species in western North America. Our results suggest a drop of more than 20% in lift production during the early stages of the moult sequence in which mid-wing flight feathers are moulted. We also found that the wing's ability to generate lift strongly depended on the morphological integrity of the outer primaries and leading-edge. These findings may explain the evolution of wing morphology and moult attributes. Specifically, the high overlap between adjacent wing feathers, especially at the wing tip, and the slow sequential replacement of the wing feathers result in a relatively small reduction in wing surface area during moult with limited aerodynamic implications. We present power and efficiency analyses for hover flight during moult under several plausible scenarios, suggesting that body mass reduction could be a compensatory mechanism that preserves the energetic costs of hover flight.
Project description:The diverse hummingbird family (Trochilidae) has unique adaptations for nectarivory, among which is the ability to sustain hover-feeding. As hummingbirds mainly feed while hovering, it is crucial to maintain this ability throughout the annual cycle-especially during flight-feather moult, in which wing area is reduced. To quantify the aerodynamic characteristics and flow mechanisms of a hummingbird wing throughout the annual cycle, time-accurate aerodynamic loads and flow field measurements were correlated over a dynamically scaled wing model of Anna's hummingbird (Calypte anna). We present measurements recorded over a model of a complete wing to evaluate the baseline aerodynamic characteristics and flow mechanisms. We found that the vorticity concentration that had developed from the wing's leading-edge differs from the attached vorticity structure that was typically found over insects' wings; firstly, it is more elongated along the wing chord, and secondly, it encounters high levels of fluctuations rather than a steady vortex. Lift characteristics resemble those of insects; however, a 20% increase in the lift-to-torque ratio was obtained for the hummingbird wing model. Time-accurate aerodynamic loads were also used to evaluate the time-evolution of the specific power required from the flight muscles, and the overall wingbeat power requirements nicely matched previous studies.
Project description:Hummingbirds are the only birds that can sustain hovering. This unique flight behaviour comes, however, at high energetic cost. Based on helicopter and aeroplane design theory, we expect that hummingbird wing aspect ratio (AR), which ranges from about 3.0 to 4.5, determines aerodynamic efficacy. Previous quasi-steady experiments with a wing spinner set-up provide no support for this prediction. To test this more carefully, we compare the quasi-steady hover performance of 26 wings, from 12 hummingbird taxa. We spun the wings at angular velocities and angles of attack that are representative for every species and measured lift and torque more precisely. The power (aerodynamic torque × angular velocity) required to lift weight depends on aerodynamic efficacy, which is measured by the power factor. Our comparative analysis shows that AR has a modest influence on lift and drag forces, as reported earlier, but interspecific differences in power factor are large. During the downstroke, the power required to hover decreases for larger AR wings at the angles of attack at which hummingbirds flap their wings (p < 0.05). Quantitative flow visualization demonstrates that variation in hover power among hummingbird wings is driven by similar stable leading edge vortices that delay stall during the down- and upstroke. A side-by-side aerodynamic performance comparison of hummingbird wings and an advanced micro helicopter rotor shows that they are remarkably similar.
Project description:To maintain the quality of the feathers, birds regularly undergo moult. It is widely accepted that moult affects flight performance, but the specific aerodynamic consequences have received relatively little attention. Here we measured the components of aerodynamic drag from the wake behind a gliding jackdaw (Corvus monedula) at different stages of its natural wing moult. We found that span efficiency was reduced (lift induced drag increased) and the wing profile drag coefficient was increased. Both effects best correlated with the corresponding reduction in spanwise camber. The negative effects are partially mitigated by adjustments of wing posture to minimize gaps in the wing, and by weight loss to reduce wing loading. By studying the aerodynamic consequences of moult, we can refine our understanding of the emergence of various moulting strategies found among birds.
Project description:Hummingbirds and nectar bats are the only vertebrates that are specialized for hovering in front of flowers to forage nectar. How their aerodynamic performance compares is, however, unclear. To hover, hummingbirds consistently generate about a quarter of the vertical aerodynamic force required to support their body weight during the upstroke. In contrast, generalist birds in slow hovering flight generate little upstroke weight support. We report that nectar bats also generate elevated weight support during the upstroke compared to generalist bats. Comparing 20 Neotropical species, we show how nectarivorous birds and bats converged on this ability by inverting their respective feathered and membrane wings more than species with other diets. However, while hummingbirds converged on an efficient horizontal wingbeat to mostly generate lift, bats rely on lift and drag during the downstroke to fully support their body weight. Furthermore, whereas the ability of nectar bats to aerodynamically support their body weight during the upstroke is elevated, it is much smaller than that of hummingbirds. Bats compensate by generating more aerodynamic weight support during their extended downstroke. Although, in principle, it requires more aerodynamic power to hover using this method, bats have adapted by evolving much larger wings for their body weight. Therefore, the net aerodynamic induced power required to hover is similar among hummingbirds and bats per unit body mass. This mechanistic insight into how feathered wings and membrane wings ultimately require similar aerodynamic power to hover may inform analogous design trade-offs in aerial robots.
Project description:Flight at high elevation is energetically demanding because of parallel reductions in air density and oxygen availability. The hovering flight of hummingbirds is one of the most energetically expensive forms of animal locomotion, but hummingbirds are nonetheless abundant at high elevations throughout the Americas. Two mechanisms enhance aerodynamic performance in high-elevation hummingbirds: increase in wing size and wing stroke amplitude during hovering. How do these changes in morphology, kinematics, and physical properties of air combine to influence the aerodynamic power requirements of flight across elevations? Here, we present data on the flight performance of 43 Andean hummingbird species as well as a 76-taxon multilocus molecular phylogeny that served as the historical framework for comparative analyses. Along a 4,000-m elevational transect, hummingbird body mass increased systematically, placing further aerodynamic demands on high-elevation taxa. However, we found that the minimum power requirements for hovering flight remain constant with respect to elevation because hummingbirds compensate sufficiently through increases in wing size and stroke amplitude. Thus, high-elevation hummingbirds are not limited in their capacity for hovering flight despite the challenges imposed by hypobaric environments. Other flight modes including vertical ascent and fast forward flight are more mechanically and energetically demanding, and we accordingly also tested for the maximum power available to hummingbirds by using a load-lifting assay. In contrast to hovering, excess power availability decreased substantially across elevations, thereby reducing the biomechanical potential for more complex flight such as competitive and escape maneuvers.
Project description:Botanical samaras spin about their centre of mass and create vertical aerodynamic forces which slow their rate of descent. Descending autorotation of animal wings, however, has never been documented. We report here that isolated wings from Anna's hummingbirds, and also from 10 species of insects, can stably autorotate and achieve descent speeds and aerodynamic performance comparable to those of samaras. A hummingbird wing loaded at its base with the equivalent of 50% of the bird's body mass descended only twice as fast as an unloaded wing, and rotated at frequencies similar to those of the wings in flapping flight. We found that even entire dead insects could stably autorotate depending on their wing postures. Feather removal trials showed no effect on descent velocity when the secondary feathers were removed from hummingbird wings. By contrast, partial removal of wing primaries substantially improved performance, except when only the outer primary was present. A scaling law for the aerodynamic performance of autorotating wings is well supported if the wing aspect ratio and the relative position of the spinning axis from the wing base are included. Autorotation is a useful and practical method that can be used to explore the aerodynamics of wing design.
Project description:The aerodynamic forces acting on a revolving dried pigeon wing and a flat card replica were measured with a propeller rig, effectively simulating a wing in continual downstroke. Two methods were adopted: direct measurement of the reaction vertical force and torque via a forceplate, and a map of the pressures along and across the wing measured with differential pressure sensors. Wings were tested at Reynolds numbers up to 108,000, typical for slow-flying pigeons, and considerably above previous similar measurements applied to insect and hummingbird wing and wing models. The pigeon wing out-performed the flat card replica, reaching lift coefficients of 1.64 compared with 1.44. Both real and model wings achieved much higher maximum lift coefficients, and at much higher geometric angles of attack (43°), than would be expected from wings tested in a windtunnel simulating translating flight. It therefore appears that some high-lift mechanisms, possibly analogous to those of slow-flying insects, may be available for birds flapping with wings at high angles of attack. The net magnitude and orientation of aerodynamic forces acting on a revolving pigeon wing can be determined from the differential pressure maps with a moderate degree of precision. With increasing angle of attack, variability in the pressure signals suddenly increases at an angle of attack between 33° and 38°, close to the angle of highest vertical force coefficient or lift coefficient; stall appears to be delayed compared with measurements from wings in windtunnels.
Project description:A theoretical model of avian flight is developed which simulates wing motion through a class of methods known as predictive simulation. This approach uses numerical optimization to predict power-optimal kinematics of avian wings in hover, cruise, climb and descent. The wing dynamics capture both aerodynamic and inertial loads. The model is used to simulate the flight of the pigeon, Columba livia, and the results are compared with previous experimental measurements. In cruise, the model unearths a vast range of kinematic modes that are capable of generating the required forces for flight. The most efficient mode uses a near-vertical stroke-plane and a flexed-wing upstroke, similar to kinematics recorded experimentally. In hover, the model predicts that the power-optimal mode uses an extended-wing upstroke, similar to hummingbirds. In flexing their wings, pigeons are predicted to consume 20% more power than if they kept their wings full extended, implying that the typical kinematics used by pigeons in hover are suboptimal. Predictions of climbing flight suggest that the most energy-efficient way to reach a given altitude is to climb as steeply as possible, subjected to the availability of power.
Project description:The shape and function of insect wings tremendously vary between insect species. This review is engaged in how wing design determines the aerodynamic mechanisms with which wings produce an air momentum for body weight support and flight control. We work out the tradeoffs associated with aerodynamic key parameters such as vortex development and lift production, and link the various components of wing structure to flight power requirements and propulsion efficiency. A comparison between rectangular, ideal-shaped and natural-shaped wings shows the benefits and detriments of various wing shapes for gliding and flapping flight. The review expands on the function of three-dimensional wing structure, on the specific role of wing corrugation for vortex trapping and lift enhancement, and on the aerodynamic significance of wing flexibility for flight and body posture control. The presented comparison is mainly concerned with wings of flies because these animals serve as model systems for both sensorimotor integration and aerial propulsion in several areas of biology and engineering.
Project description:An aerodynamic structure ubiquitous in Aves is the alula; a small collection of feathers muscularized near the wrist joint. New research into the aerodynamics of this structure suggests that its primary function is to induce leading-edge vortex (LEV) flow over bird's outer hand-wing to enhance wing lift when manuevering at slow speeds. Here, we explore scaling trends of the alula's spanwise position and its connection to this function. Specifically, we test the hypothesis that the relative distance of the alula from the wing tip is that which maximizes LEV-lift when the wing is spread and operated in a deep-stall flight condition. To test this, we perform experiments on model wings in a wind tunnel to approximate this distance and compare our results to positional measurements of the alula on spread-wing specimens. We found the position of the alula on non-aquatic birds selected for alula optimization to be located at or near the lift-maximizing position predicted by wind tunnel experiments. These findings shed new light on avian wing anatomy and the role of unconventional aerodynamics in shaping it.