More than a million insects and approximately 11,000 vertebrates utilize flapping wings to fly. However, flapping flight has only been studied in a few of these species, so many challenges remain in ...understanding this form of locomotion. Five key aerodynamic mechanisms have been identified for insect flight. Among these is the leading edge vortex, which is a convergent solution to avoid stall for insects, bats and birds. The roles of the other mechanisms - added mass, clap and fling, rotational circulation and wing-wake interactions - have not yet been thoroughly studied in the context of vertebrate flight. Further challenges to understanding bat and bird flight are posed by the complex, dynamic wing morphologies of these species and the more turbulent airflow generated by their wings compared with that observed during insect flight. Nevertheless, three dimensionless numbers that combine key flow, morphological and kinematic parameters - the Reynolds number, Rossby number and advance ratio - govern flapping wing aerodynamics for both insects and vertebrates. These numbers can thus be used to organize an integrative framework for studying and comparing animal flapping flight. Here, we provide a roadmap for developing such a framework, highlighting the aerodynamic mechanisms that remain to be quantified and compared across species. Ultimately, incorporating complex flight maneuvers, environmental effects and developmental stages into this framework will also be essential to advancing our understanding of the biomechanics, movement ecology and evolution of animal flight.
The aerodynamic performance of hovering insects is largely explained by the presence of a stably attached leading edge vortex (LEV) on top of their wings. Although LEVs have been visualized on real, ...physically modeled, and simulated insects, the physical mechanisms responsible for their stability are poorly understood. To gain fundamental insight into LEV stability on flapping fly wings we expressed the Navier-Stokes equations in a rotating frame of reference attached to the wing's surface. Using these equations we show that LEV dynamics on flapping wings are governed by three terms: angular, centripetal and Coriolis acceleration. Our analysis for hovering conditions shows that angular acceleration is proportional to the inverse of dimensionless stroke amplitude, whereas Coriolis and centripetal acceleration are proportional to the inverse of the Rossby number. Using a dynamically scaled robot model of a flapping fruit fly wing to systematically vary these dimensionless numbers, we determined which of the three accelerations mediate LEV stability. Our force measurements and flow visualizations indicate that the LEV is stabilized by the ;quasi-steady' centripetal and Coriolis accelerations that are present at low Rossby number and result from the propeller-like sweep of the wing. In contrast, the unsteady angular acceleration that results from the back and forth motion of a flapping wing does not appear to play a role in the stable attachment of the LEV. Angular acceleration is, however, critical for LEV integrity as we found it can mediate LEV spiral bursting, a high Reynolds number effect. Our analysis and experiments further suggest that the mechanism responsible for LEV stability is not dependent on Reynolds number, at least over the range most relevant for insect flight (100<Re<14,000). LEVs are stable and continue to augment force even when they burst. These and similar findings for propellers and wind turbines at much higher Reynolds numbers suggest that even large flying animals could potentially exploit LEV-based force augmentation during slow hovering flight, take-offs or landing. We calculated the Rossby number from single-wing aspect ratios of over 300 insects, birds, bats, autorotating seeds, and pectoral fins of fish. We found that, on average, wings and fins have a Rossby number close to that of flies (Ro=3). Theoretically, many of these animals should therefore be able to generate a stable LEV, a prediction that is supported by recent findings for several insects, one bat, one bird and one fish. This suggests that force augmentation through stably attached (leading edge) vortices could represent a convergent solution for the generation of high fluid forces over a quite large range in size.
The lift that animal wings generate to fly is typically considered a vertical force that supports weight, while drag is considered a horizontal force that opposes thrust. To determine how birds use ...lift and drag, here we report aerodynamic forces and kinematics of Pacific parrotlets (Forpus coelestis) during short, foraging flights. At takeoff they incline their wing stroke plane, which orients lift forward to accelerate and drag upward to support nearly half of their bodyweight. Upon landing, lift is oriented backward to contribute a quarter of the braking force, which reduces the aerodynamic power required to land. Wingbeat power requirements are dominated by downstrokes, while relatively inactive upstrokes cost almost no aerodynamic power. The parrotlets repurpose lift and drag during these flights with lift-to-drag ratios below two. Such low ratios are within range of proto-wings, showing how avian precursors may have relied on drag to take off with flapping wings.
Organisms that swim or fly with fins or wings physically interact with the surrounding water and air. The interactions are governed by the morphology and kinematics of the locomotory system that form ...boundary conditions to the Navier-Stokes (NS) equations. These equations represent Newton's law of motion for the fluid surrounding the organism. Several dimensionless numbers, such as the Reynolds number and Strouhal number, measure the influence of morphology and kinematics on the fluid dynamics of swimming and flight. There exists, however, no coherent theoretical framework that shows how such dimensionless numbers of organisms are linked to the NS equation. Here we present an integrated approach to scale the biological fluid dynamics of a wing that flaps, spins or translates. Both the morphology and kinematics of the locomotory system are coupled to the NS equation through which we find dimensionless numbers that represent rotational accelerations in the flow due to wing kinematics and morphology. The three corresponding dimensionless numbers are (1) the angular acceleration number, (2) the centripetal acceleration number, and (3) the Rossby number, which measures Coriolis acceleration. These dimensionless numbers consist of length scale ratios, which facilitate their geometric interpretation. This approach gives fundamental insight into the physical mechanisms that explain the differences in performance among flapping, spinning and translating wings. Although we derived this new framework for the special case of a model fly wing, the method is general enough to make it applicable to other organisms that fly or swim using wings or fins.
How wingless salamanders fly Lentink, David
Nature (London),
06/2022, Letnik:
606, Številka:
7913
Journal Article
Recenzirano
Parabolic-flight data1 indicate that some reptiles and amphibians floating in microgravity adopt skydiving postures, which is unexpected because most of these animals do not have body shapes adapted ...for aerial behaviour. To find out whether this was the case, Brown et al.4, writing in Current Biology, released five of these salamanders in a vertical wind tunnel. The need to survive if an animal fails to keep its footing and falls has driven the independent evolution of effective aerial behaviours in a colourful assembly of tree-dwelling animals - including ants, spiders, salamanders, frogs, lizards, snakes, squirrels, lemurs and marsupials - despite these animals not having wings or having only makeshift ones6.
Swifts are among the most aerodynamically refined gliding birds. However, the overlapping vanes and protruding shafts of their primary feathers make swift wings remarkably rough for their size. Wing ...roughness height is 1-2% of chord length on the upper surface--10,000 times rougher than sailplane wings. Sailplanes depend on extreme wing smoothness to increase the area of laminar flow on the wing surface and minimize drag for extended glides. To understand why the swift does not rely on smooth wings, we used a stethoscope to map laminar flow over preserved wings in a low-turbulence wind tunnel. By combining laminar area, lift, and drag measurements, we show that average area of laminar flow on swift wings is 69% (n = 3; std 13%) of their total area during glides that maximize flight distance and duration--similar to high-performance sailplanes. Our aerodynamic analysis indicates that swifts attain laminar flow over their rough wings because their wing size is comparable to the distance the air travels (after a roughness-induced perturbation) before it transitions from laminar to turbulent. To interpret the function of swift wing roughness, we simulated its effect on smooth model wings using physical models. This manipulation shows that laminar flow is reduced and drag increased at high speeds. At the speeds at which swifts cruise, however, swift-like roughness prolongs laminar flow and reduces drag. This feature gives small birds with rudimentary wings an edge during the evolution of glide performance.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Diurnal flying animals such as birds depend primarily on vision to coordinate their flight path during goal-directed flight tasks. To extract the spatial structure of the surrounding environment, ...birds are thought to use retinal image motion (optical flow) that is primarily induced by motion of their head. It is unclear what gaze behaviors birds perform to support visuomotor control during rapid maneuvering flight in which they continuously switch between flight modes. To analyze this, we measured the gaze behavior of rapidly turning lovebirds in a goal-directed task: take-off and fly away from a perch, turn on a dime, and fly back and land on the same perch. High-speed flight recordings revealed that rapidly turning lovebirds perform a remarkable stereotypical gaze behavior with peak saccadic head turns up to 2700 degrees per second, as fast as insects, enabled by fast neck muscles. In between saccades, gaze orientation is held constant. By comparing saccade and wingbeat phase, we find that these super-fast saccades are coordinated with the downstroke when the lateral visual field is occluded by the wings. Lovebirds thus maximize visual perception by overlying behaviors that impair vision, which helps coordinate maneuvers. Before the turn, lovebirds keep a high contrast edge in their visual midline. Similarly, before landing, the lovebirds stabilize the center of the perch in their visual midline. The perch on which the birds land swings, like a branch in the wind, and we find that retinal size of the perch is the most parsimonious visual cue to initiate landing. Our observations show that rapidly maneuvering birds use precisely timed stereotypic gaze behaviors consisting of rapid head turns and frontal feature stabilization, which facilitates optical flow based flight control. Similar gaze behaviors have been reported for visually navigating humans. This finding can inspire more effective vision-based autopilots for drones.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Swifts are aerodynamically sophisticated birds with a small arm and large hand wing that provides them with exquisite control over their glide performance. However, their hand wings have a seemingly ...unsophisticated surface roughness that is poised to disturb flow. This roughness of about 2% chord length is formed by the valleys and ridges of overlapping primary feathers with thick protruding rachides, which make the wing stiffer. An earlier flow study of laminar-turbulent boundary layer transition over prepared swift wings suggested that swifts can attain laminar flow at a low angle of attack. In contrast, aerodynamic design theory suggests that airfoils must be extremely smooth to attain such laminar flow. In hummingbirds, which have similarly rough wings, flow measurements on a 3D printed model suggest that the flow separates at the leading edge and becomes turbulent well above the rachis bumps in a detached shear layer. The aerodynamic function of wing roughness in small birds is, therefore, not fully understood. Here, we performed particle image velocimetry and force measurements to compare smooth versus rough 3D-printed models of the swift hand wing. The high-resolution boundary layer measurements show that the flow over rough wings is indeed laminar at a low angle of attack and a low Reynolds number, but becomes turbulent at higher values. In contrast, the boundary layer over the smooth wing forms open laminar separation bubbles that extend beyond the trailing edge. The boundary layer dynamics of the smooth surface varies non-linearly as a function of angle of attack and Reynolds number, whereas the rough surface boasts more consistent turbulent boundary layer dynamics. Comparison of the corresponding drag values, lift values and glide ratios suggests, however, that glide performance is equivalent. The increased structural performance, boundary layer robustness and equivalent aerodynamic performance of rough wings might have provided small (proto) birds with an evolutionary window to high glide performance.
Birds fly effectively and maneuver nimbly by dynamically changing the shape of their wings during each wingbeat. These shape changes have yet to be quantified automatically at high temporal and ...spatial resolution. Therefore, we developed a custom 3D surface reconstruction method, which uses a high-speed camera to identify spatially encoded binary striped patterns that are projected on a flying bird. This non-invasive structured-light method allows automated 3D reconstruction of each stand-alone frame and can be extended to multiple views. We demonstrate this new technique by automatically reconstructing the dorsal surface of a parrotlet wing at 3200 frames s
during flapping flight. From this shape we analyze key parameters such as wing twist and angle of attack distribution. While our binary 'single-shot' algorithm is demonstrated by quantifying dynamic shape changes of a flying bird, it is generally applicable to moving animals, plants and deforming objects.
Harnessing flight strategies refined by millions of years of evolution can help expedite the design of more efficient, manoeuvrable and robust flying robots. This review synthesizes recent advances ...and highlights remaining gaps in our understanding of how bird and bat wing adaptations enable effective flight. Included in this discussion is an evaluation of how current robotic analogues measure up to their biological sources of inspiration. Studies of vertebrate wings have revealed skeletal systems well suited for enduring the loads required during flight, but the mechanisms that drive coordinated motions between bones and connected integuments remain ill-described. Similarly, vertebrate flight muscles have adapted to sustain increased wing loading, but a lack of in vivo studies limits our understanding of specific muscular functions. Forelimb adaptations diverge at the integument level, but both bird feathers and bat membranes yield aerodynamic surfaces with a level of robustness unparalleled by engineered wings. These morphological adaptations enable a diverse range of kinematics tuned for different flight speeds and manoeuvres. By integrating vertebrate flight specializations—particularly those that enable greater robustness and adaptability—into the design and control of robotic wings, engineers can begin narrowing the wide margin that currently exists between flying robots and vertebrates. In turn, these robotic wings can help biologists create experiments that would be impossible in vivo.
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