Systematic characterization of wing mechanosensors that monitor airflow and wing deformations.

Animal wings deform during flight in ways that can enhance lift, facilitate flight control, and mitigate damage. Monitoring the structural and aerodynamic state of the wing is challenging because deformations are passive, and the flow fields are unsteady; it requires distributed mechanosensors that respond to local airflow and strain on the wing. Without a complete map of the sensor arrays, it is impossible to model control strategies underpinned by them. Here, we present the first systematic characterization of mechanosensors on the dragonfly's wings: morphology, distribution, and wiring. By combining a cross-species survey of sensor distribution with quantitative neuroanatomy and a high-fidelity finite element analysis, we show that the mechanosensors are well placed to perceive features of the wing dynamics relevant to flight. This work describes the wing sensory apparatus in its entirety and advances our understanding of the sensorimotor loop that facilitates exquisite flight control in animals with highly deformable wings.

Kinematics and hydrodynamics analyses of swimming penguins: wing bending improves propulsion performance.

Penguins are adapted to underwater life and have excellent swimming abilities. Although previous motion analyses revealed their basic swimming characteristics, the details of the 3D wing kinematics, wing deformation and thrust generation mechanism of penguins are still largely unknown. In this study, we recorded the forward and horizontal swimming of gentoo penguins (Pygoscelis papua) at an aquarium with multiple underwater action cameras and then performed a 3D motion analysis. We also conducted a series of water tunnel experiments with a 3D printed rigid wing to obtain lift and drag coefficients in the gliding configuration. Using these coefficients, the thrust force during flapping was calculated in a quasi-steady manner, where the following two wing models were considered: (1) an 'original' wing model reconstructed from 3D motion analysis including bending deformation and (2) a 'flat' wing model obtained by flattening the original wing model. The resultant body trajectory showed that the penguin accelerated forward during both upstroke and downstroke. The motion analysis of the two wing models revealed that considerable bending occurred in the original wing, which reduced its angle of attack during the upstroke in particular. Consequently, the calculated stroke-averaged thrust was larger for the original wing than for the flat wing during the upstroke. In addition, the propulsive efficiency for the original wing was estimated to be 1.8 times higher than that for the flat wing. Our results unveil a detailed mechanism of lift-based propulsion in penguins and underscore the importance of wing bending.

Raptor wing morphing with flight speed.

In gliding flight, birds morph their wings and tails to control their flight trajectory and speed. Using high-resolution videogrammetry, we reconstructed accurate and detailed three-dimensional geometries of gliding flights for three raptors (barn owl, Tyto alba; tawny owl, Strix aluco, and goshawk, Accipiter gentilis). Wing shapes were highly repeatable and shoulder actuation was a key component of reconfiguring the overall planform and controlling angle of attack. The three birds shared common spanwise patterns of wing twist, an inverse relationship between twist and peak camber, and held their wings depressed below their shoulder in an anhedral configuration. With increased speed, all three birds tended to reduce camber throughout the wing, and their wings bent in a saddle-shape pattern. A number of morphing features suggest that the coordinated movements of the wing and tail support efficient flight, and that the tail may act to modulate wing camber through indirect aeroelastic control.

Mechanical power curve measured in the wake of pied flycatchers indicates modulation of parasite power across flight speeds.

How aerodynamic power required for animal flight varies with flight speed determines optimal speeds during foraging and migratory flight. Despite its relevance, aerodynamic power provides an elusive quantity to measure directly in animal flight. Here, we determine the aerodynamic power from wake velocity fields, measured using tomographical particle image velocimetry, of pied flycatchers flying freely in a wind tunnel. We find a shallow U-shaped power curve, which is flatter than expected by theory. Based on how the birds vary body angle with speed, we speculate that the shallow curve results from increased body drag coefficient and body frontal area at lower flight speeds. Including modulation of body drag in the model results in a more reasonable fit with data than the traditional model. From the wake structure, we also find a single starting vortex generated from the two wings during the downstroke across flight speeds (1-9 m s-1). This is accomplished by the arm wings interacting at the beginning of the downstroke, generating a unified starting vortex above the body of the bird. We interpret this as a mechanism resulting in a rather uniform downwash and low induced power, which can help explain the higher aerodynamic performance in birds compared with bats.

Quantifying the dynamic wing morphing of hovering hummingbird.

Animal wings are lightweight and flexible; hence, during flapping flight their shapes change. It has been known that such dynamic wing morphing reduces aerodynamic cost in insects, but the consequences in vertebrate flyers, particularly birds, are not well understood. We have developed a method to reconstruct a three-dimensional wing model of a bird from the wing outline and the feather shafts (rachides). The morphological and kinematic parameters can be obtained using the wing model, and the numerical or mechanical simulations may also be carried out. To test the effectiveness of the method, we recorded the hovering flight of a hummingbird (Amazilia amazilia) using high-speed cameras and reconstructed the right wing. The wing shape varied substantially within a stroke cycle. Specifically, the maximum and minimum wing areas differed by 18%, presumably due to feather sliding; the wing was bent near the wrist joint, towards the upward direction and opposite to the stroke direction; positive upward camber and the 'washout' twist (monotonic decrease in the angle of incidence from the proximal to distal wing) were observed during both half-strokes; the spanwise distribution of the twist was uniform during downstroke, but an abrupt increase near the wrist joint was found during upstroke.

Aerodynamic Ground Effect in Fruitfly Sized Insect Takeoff.

Aerodynamic ground effect in flapping-wing insect flight is of importance to comparative morphologies and of interest to the micro-air-vehicle (MAV) community. Recent studies, however, show apparently contradictory results of either some significant extra lift or power savings, or zero ground effect. Here we present a numerical study of fruitfly sized insect takeoff with a specific focus on the significance of leg thrust and wing kinematics. Flapping-wing takeoff is studied using numerical modelling and high performance computing. The aerodynamic forces are calculated using a three-dimensional Navier-Stokes solver based on a pseudo-spectral method with volume penalization. It is coupled with a flight dynamics solver that accounts for the body weight, inertia and the leg thrust, while only having two degrees of freedom: the vertical and the longitudinal horizontal displacement. The natural voluntary takeoff of a fruitfly is considered as reference. The parameters of the model are then varied to explore possible effects of interaction between the flapping-wing model and the ground plane. These modified takeoffs include cases with decreased leg thrust parameter, and/or with periodic wing kinematics, constant body pitch angle. The results show that the ground effect during natural voluntary takeoff is negligible. In the modified takeoffs, when the rate of climb is slow, the difference in the aerodynamic forces due to the interaction with the ground is up to 6%. Surprisingly, depending on the kinematics, the difference is either positive or negative, in contrast to the intuition based on the helicopter theory, which suggests positive excess lift. This effect is attributed to unsteady wing-wake interactions. A similar effect is found during hovering.