Interaction of barn owl leading edge serrations with freestream turbulence.

The silent flight of barn owls is associated with wing and feather specialisations. Three special features are known: a serrated leading edge that is formed by free-standing barb tips which appears as a comb-like structure, a soft dorsal surface, and a fringed trailing edge. We used a model of the leading edge comb with 3D-curved serrations that was designed based on 3D micro-scans of rows of barbs from selected barn-owl feathers. The interaction of the flow with the serrations was measured with Particle-Image-Velocimetry in a flow channel at uniform steady inflow and was compared to the situation of inflow with freestream turbulence, generated from the turbulent wake of a cylinder placed upstream. In steady uniform flow, the serrations caused regular velocity streaks and a flow turning effect. When vortices of different size impacted the serrations, the serrations reduced the flow fluctuations downstream in each case, exemplified by a decreased root-mean-square value of the fluctuations in the wake of the serrations. This attenuation effect was stronger for the spanwise velocity component, leading to an overall flow homogenization. Our findings suggest that the serrations of the barn owl provide a passive flow control leading to reduced leading-edge noise when flying in turbulent environments.

Animal behavior: Mosquitos ride the wave.

Some insects have a frustrating knack for avoiding a swatter. A new study shows that mosquitos not only evade the visual image of the looming threat, they also surf the wave of air the swatter creates.

Machine learning reveals the control mechanics of an insect wing hinge.

Insects constitute the most species-rich radiation of metazoa, a success that is due to the evolution of active flight. Unlike pterosaurs, birds and bats, the wings of insects did not evolve from legs1, but are novel structures that are attached to the body via a biomechanically complex hinge that transforms tiny, high-frequency oscillations of specialized power muscles into the sweeping back-and-forth motion of the wings2. The hinge consists of a system of tiny, hardened structures called sclerites that are interconnected to one another via flexible joints and regulated by the activity of specialized control muscles. Here we imaged the activity of these muscles in a fly using a genetically encoded calcium indicator, while simultaneously tracking the three-dimensional motion of the wings with high-speed cameras. Using machine learning, we created a convolutional neural network3 that accurately predicts wing motion from the activity of the steering muscles, and an encoder-decoder4 that predicts the role of the individual sclerites on wing motion. By replaying patterns of wing motion on a dynamically scaled robotic fly, we quantified the effects of steering muscle activity on aerodynamic forces. A physics-based simulation incorporating our hinge model generates flight manoeuvres that are remarkably similar to those of free-flying flies. This integrative, multi-disciplinary approach reveals the mechanical control logic of the insect wing hinge, arguably among the most sophisticated and evolutionarily important skeletal structures in the natural world.

An open platform for visual stimulation of insects.

To study how the nervous system processes visual information, experimenters must record neural activity while delivering visual stimuli in a controlled fashion. In animals with a nearly panoramic field of view, such as flies, precise stimulation of the entire visual field is challenging. We describe a projector-based device for stimulation of the insect visual system under a microscope. The device is based on a bowl-shaped screen that provides a wide and nearly distortion-free field of view. It is compact, cheap, easy to assemble, and easy to operate using the included open-source software for stimulus generation. We validate the virtual reality system technically and demonstrate its capabilities in a series of experiments at two levels: the cellular, by measuring the membrane potential responses of visual interneurons; and the organismal, by recording optomotor and fixation behavior of Drosophila melanogaster in tethered flight. Our experiments reveal the importance of stimulating the visual system of an insect with a wide field of view, and we provide a simple solution to do so.

Joint extension speed dictates bio-inspired morphing trajectories for optimal longitudinal flight dynamics.

Avian wing morphing allows dynamic, active control of complex flight manoeuvres. Previous linear time-invariant (LTI) models have quantified the effect of varying fixed wing configurations but the time-dependent effects of morphing between different configurations is not well understood. To fill this gap, I implemented a linear parameter-varying (LPV) model for morphing wing gull flight. This approach models the wing joint angles as scheduled parameters and accounts for nonlinear kinematic and gravitational effects while interpolating between LTI models at discrete trim points. With the resulting model, I investigated the longitudinal response associated with various joint extension trajectories. By optimizing the extension trajectory for four independent objectives (speed and pitch angle overshoot, speed rise time and pitch angle settling time), I found that the extension trajectory inherent to the gull wing does not guarantee an optimal response but may provide a sufficient response with a simpler mechanical implementation. Furthermore, the results indicated that gulls likely require extension speed feedback. This morphing LPV model provides insights into underlying control mechanisms, which may allow for avian-like flight in future highly manoeuvrable uncrewed aerial vehicles.

Palaeoatmosphere facilitates a gliding transition to powered flight in the Eocene bat, Onychonycteris finneyi.

The evolutionary transition to powered flight remains controversial in bats, the only flying mammals. We applied aerodynamic modeling to reconstruct flight in the oldest complete fossil bat, the archaic Onychonycteris finneyi from the early Eocene of North America. Results indicate that Onychonycteris was capable of both gliding and powered flight either in a standard normodense aerial medium or in the hyperdense atmosphere that we estimate for the Eocene from two independent palaeogeochemical proxies. Aerodynamic continuity across a morphological gradient is further demonstrated by modeled intermediate forms with increasing aspect ratio (AR) produced by digital elongation based on chiropteran developmental data. Here a gliding performance gradient emerged of decreasing sink rate with increasing AR that eventually allowed applying available muscle power to achieve level flight using flapping, which is greatly facilitated in hyperdense air. This gradient strongly supports a gliding (trees-down) transition to powered flight in bats.

Flying fast improves aerodynamic economy of heavier birds.

A paradox of avian long-distance migrations is that birds must greatly increase their body mass prior to departure, yet this is presumed to substantially increase their energy cost of flight. However, here we show that when homing pigeons flying in a flock are loaded with ventrally located weight, both their heart rate and estimated energy expenditure rise by a remarkably small amount. The net effect is that costs per unit time increase only slightly and per unit mass they decrease. We suggest that this is because these homing flights are relatively fast, and consequently flight costs associated with increases in body parasite drag dominate over those of weight support, leading to an improvement in mass-specific flight economy. We propose that the relatively small absolute aerodynamic penalty for carrying enlarged fuel stores and flight muscles during fast flight has helped to select for the evolution of long-distance migration.

Landing and take-off capabilities of bioinspired aerial vehicles: a review.

Bioinspired flapping-wing micro aerial vehicles (FWMAVs) have emerged over the last two decades as a promising new type of robot. Their high thrust-to-weight ratio, versatility, safety, and maneuverability, especially at small scales, could make them more suitable than fixed-wing and multi-rotor vehicles for various applications, especially in cluttered, confined environments and in close proximity to humans, flora, and fauna. Unlike natural flyers, however, most FWMAVs currently have limited take-off and landing capabilities. Natural flyers are able to take off and land effortlessly from a wide variety of surfaces and in complex environments. Mimicking such capabilities on flapping-wing robots would considerably enhance their practical usage. This review presents an overview of take-off and landing techniques for FWMAVs, covering different approaches and mechanism designs, as well as dynamics and control aspects. The special case of perching is also included. As well as discussing solutions investigated for FWMAVs specifically, we also present solutions that have been developed for different types of robots but may be applicable to flapping-wing ones. Different approaches are compared and their suitability for different applications and types of robots is assessed. Moreover, research and technology gaps are identified, and promising future work directions are identified.

A thermal performance curve perspective explains decades of disagreements over how air temperature affects the flight metabolism of honey bees.

While multiple studies have shown that honey bees and some other flying insects lower their flight metabolic rates when flying at high air temperatures, critics have suggested such patterns result from poor experimental methods as, theoretically, air temperature should not appreciably affect aerodynamic force requirements. Here, we show that apparently contradictory studies can be reconciled by considering the thermal performance curve of flight muscle. We show that prior studies that found no effects of air temperature on flight metabolism of honey bees achieved flight muscle temperatures that were near or on equal, opposite sides of the thermal performance curve. Honey bees vary their wing kinematics and metabolic heat production to thermoregulate, and how air temperature affects the flight metabolic rate of honey bees is predictable using a non-linear thermal performance perspective of honey bee flight muscle.

Evolutionary change in flight-to-light response in urban moths comes with changes in wing morphology.

Moths and other insects are attracted by artificial light sources. This flight-to-light behaviour disrupts their general activity focused on finding resources, such as mating partners, and increases predation risk. It thus has substantial fitness costs. In illuminated urban areas, spindle ermine moths Yponomeuta cagnagella were reported to have evolved a reduced flight-to-light response. Yet, the specific mechanism remained unknown, and was hypothesized to involve either changes in visual perception or general flight ability or overall mobility traits. Here, we test whether spindle ermine moths from urban and rural populations-with known differences in flight-to-light responses-differ in flight-related morphological traits. Urban individuals were found to have on average smaller wings than rural moths, which in turn correlated with a lower probability of being attracted to an artificial light source. Our finding supports the reduced mobility hypothesis, which states that reduced mobility in urban areas is associated with specific morphological changes in the flight apparatus.

Turbulence causes kinematic and behavioural adjustments in a flapping flier.

Turbulence is a widespread phenomenon in the natural world, but its influence on flapping fliers remains little studied. We assessed how freestream turbulence affected the kinematics, flight effort and track properties of homing pigeons (Columba livia), using the fine-scale variations in flight height as a proxy for turbulence levels. Birds showed a small increase in their wingbeat amplitude with increasing turbulence (similar to laboratory studies), but this was accompanied by a reduction in mean wingbeat frequency, such that their flapping wing speed remained the same. Mean kinematic responses to turbulence may therefore enable birds to increase their stability without a reduction in propulsive efficiency. Nonetheless, the most marked response to turbulence was an increase in the variability of wingbeat frequency and amplitude. These stroke-to-stroke changes in kinematics provide instantaneous compensation for turbulence. They will also increase flight costs. Yet pigeons only made small adjustments to their flight altitude, likely resulting in little change in exposure to strong convective turbulence. Responses to turbulence were therefore distinct from responses to wind, with the costs of high turbulence being levied through an increase in the variability of their kinematics and airspeed. This highlights the value of investigating the variability in flight parameters in free-living animals.

Analysis of collision avoidance in honeybee flight.

Insects are excellent at flying in dense vegetation and navigating through other complex spatial environments. This study investigates the strategies used by honeybees (Apis mellifera) to avoid collisions with an obstacle encountered frontally during flight. Bees were trained to fly through a tunnel that contained a solitary vertically oriented cylindrical obstacle placed along the midline. Flight trajectories of bees were recorded for six conditions in which the diameter of the obstructing cylinder was systematically varied from 25 mm to 160 mm. Analysis of salient events during the bees' flight, such as the deceleration before the obstacle, and the initiation of the deviation in flight path to avoid collisions, revealed a strategy for obstacle avoidance that is based on the relative retinal expansion velocity generated by the obstacle when the bee is on a collision course. We find that a quantitative model, featuring a controller that extracts specific visual cues from the frontal visual field, provides an accurate characterization of the geometry and the dynamics of the manoeuvres adopted by honeybees to avoid collisions. This study paves the way for the design of unmanned aerial systems, by identifying the visual cues that are used by honeybees for performing robust obstacle avoidance flight.

Aerodynamic response of a red-tailed hawk to discrete transverse gusts.

A limiting factor in the design of smaller size uncrewed aerial vehicles is their inability to navigate through gust-laden environments. As a result, engineers have turned towards bio-inspired engineering approaches for gust mitigation techniques. In this study, the aerodynamics of a red-tailed hawk's response to variable-magnitude discrete transverse gusts was investigated. The hawk was flown in an indoor flight arena instrumented by multiple high-speed cameras to quantify the 3D motion of the bird as it navigated through the gust. The hawk maintained its flapping motion across the gust in all runs; however, it encountered the gust at different points in the flapping pattern depending on the run and gust magnitude. The hawk responded with a downwards pitching motion of the wing, decreasing the wing pitch angle to between -20∘and -5∘, and remained in this configuration until gust exit. The wing pitch data was then applied to a lower-order aerodynamic model that estimated lift coefficients across the wing. In gusts slower than the forward flight velocity (low gust ratio), the lift coefficient increases at a low-rate, to a maximum of around 2-2.5. In gusts faster than the forward flight velocity (high gust ratio), the lift coefficient initially increased rapidly, before increasing at a low-rate to a value around 4-5. In both regimes, the hawk's observed height change due to gust interaction was similar (and small), despite larger estimated lift coefficients over the high gust regime. This suggests another mitigation factor apart from the wing response is present. One potential factor is the tail pitching response observed here, which prior work has shown serves to mitigate pitch disturbances from gusts.

Evolution, types, and distribution of flight control devices on wings and elytra in bark beetles.

Gaining the ability to fly actively was a ground-breaking moment in insect evolution, providing an unprecedented advantage over other arthropods. Nevertheless, active flight was a costly innovation, requiring the development of wings and flight muscles, the provision of sufficient energetic resources, and a complex flight control system. Although wings, flight muscles, and the energetic budget of insects have been intensively studied in the last decades, almost nothing is known regarding the flight-control devices of many crucial insect groups, especially beetles (Coleoptera). Here, we conducted a phylogenetic-informed analysis of flight-related mechanosensors in 28 species of bark beetles (Curculionidae: Scolytinae, Platypodinae), an economically and ecologically important group of insects characterized by striking differences in dispersal abilities. The results indicated that beetle flight apparatus is equipped with different functional types of mechanosensors, including strain- and flow-encoding sensilla. We found a strong effect of allometry on the number of mechanosensors, while no effect of relative wing size (a proxy of flight investment) was identified. Our study constitutes the first step to understanding the drivers and constraints of the evolution of flight-control devices in Coleoptera, including bark beetles. More research, including a quantitative neuroanatomical analysis of beetle wings, should be conducted in the future.

Bumblebees compensate for the adverse effects of sidewind during visually guided landings.

Flying animals often encounter winds during visually guided landings. However, how winds affect their flight control strategy during landing is unknown. Here, we investigated how sidewind affects the landing performance and sensorimotor control of foraging bumblebees (Bombus terrestris). We trained bumblebees to forage in a wind tunnel, and used high-speed stereoscopic videography to record 19,421 landing maneuvers in six sidewind speeds (0 to 3.4 m s-1), which correspond to winds encountered in nature. Bumblebees landed less often in higher windspeeds, but the landing durations from free flight were not increased by wind. By testing how bumblebees adjusted their landing control to compensate for adverse effects of sidewind on landing, we showed that the landing strategy in sidewind resembled that in still air, but with important adaptations. Bumblebees landing in a sidewind tended to drift downwind, which they controlled for by performing more hover maneuvers. Surprisingly, the increased hover prevalence did not increase the duration of free-flight landing maneuvers, as these bumblebees flew faster towards the landing platform outside the hover phases. Hence, by alternating these two flight modes along their flight path, free-flying bumblebees negated the adverse effects of high windspeeds on landing duration. Using control theory, we hypothesize that bumblebees achieve this by integrating a combination of direct aerodynamic feedback and a wind-mediated mechanosensory feedback control, with their vision-based sensorimotor control loop. The revealed landing strategy may be commonly used by insects landing in windy conditions, and may inspire the development of landing control strategies onboard autonomously flying robots.

Dynamic antennal positioning allows honeybee followers to decode the dance.

The honeybee waggle dance has been widely studied as a communication system, yet we know little about how nestmates assimilate the information needed to navigate toward the signaled resource. They are required to detect the dancer's orientation relative to gravity and duration of the waggle phase and translate this into a flight vector with a direction relative to the sun1 and distance from the hive.2,3 Moreover, they appear capable of doing so from varied, dynamically changing positions around the dancer. Using high-speed, high-resolution video, we have uncovered a previously unremarked correlation between antennal position and the relative body axes of dancer and follower bees. Combined with new information about antennal inputs4,5 and spatial encoding in the insect central complex,6,7 we show how a neural circuit first proposed to underlie path integration could be adapted to decoding the dance and acquiring the signaled information as a flight vector that can be followed to the resource. This provides the first plausible account of how the bee brain could support the interpretation of its dance language.

Proximity to the water surface markedly enhances the force production on underwater flapping wings.

The potential application of flapping wings in micro-aerial vehicles is gaining interest due to their ability to generate high lift even in confined spaces. Most studies in the past have investigated hovering wings as well as those flapping near solid surfaces. However, the presence of surface tension at the water-air interface and the ability of the water surface to move might differentiate its response to the proximity of wings, compared to that of solid surfaces. Motivated by underwater, amphibian robots and several underwater experimental studies on flapping wings, our study investigated the effects of the proximity of flapping wings to the water surface at low Reynolds numbers (Re = 3400). Experiments were performed on a rectangular wing in a water tank with prescribed flapping kinematics and the aerodynamic forces were measured. The effects of surface proximity on the wing in its both upright and inverted orientations were studied. Broadly, the mean lift and drag coefficients in both orientations decreased significantly (by up to 60%) as the distance from the water surface was increased. In the case of the upright orientation, the mean lift coefficient was slightly decreased very close to the water surface with its peak being observed at the normalized clearance of [Formula: see text]. Overall, the study revealed an enhancement in the aerodynamic forces closer to the water surface.

Small deviations in kinematics and body form dictate muscle performances in the finely tuned avian downstroke.

Avian takeoff requires peak pectoralis muscle power to generate sufficient aerodynamic force during the downstroke. Subsequently, the much smaller supracoracoideus recovers the wing during the upstroke. How the pectoralis work loop is tuned to power flight is unclear. We integrate wingbeat-resolved muscle, kinematic, and aerodynamic recordings in vivo with a new mathematical model to disentangle how the pectoralis muscle overcomes wing inertia and generates aerodynamic force during takeoff in doves. Doves reduce the angle of attack of their wing mid-downstroke to efficiently generate aerodynamic force, resulting in an aerodynamic power dip, that allows transferring excess pectoralis power into tensioning the supracoracoideus tendon to assist the upstroke-improving the pectoralis work loop efficiency simultaneously. Integrating extant bird data, our model shows how the pectoralis of birds with faster wingtip speed need to generate proportionally more power. Finally, birds with disproportionally larger wing inertia need to activate the pectoralis earlier to tune their downstroke.

Energy consumption during insect flight and bioinspiration for MAV design: A review.

The excellent biological characteristics of insects provide an important source of inspiration for designing micro air vehicles (MAVs). Insect flight is an incredibly complex and energy-intensive process. Unique insect flight muscles and contraction mechanisms enable flapping at high frequencies. Moreover, the metabolic rate during flight can reach hundreds of times the resting state. Understanding energy consumption during flight is crucial for designing efficient biomimetic aircraft. This paper summarizes the structures and contraction mechanisms of insect flight muscles, explores the underlying metabolic processes, and identifies methods for energy substrate identification and detection, and discusses inspiration for biomimetic MAV design. This paper reviews energy consumption during insect flight, promotes the understanding of insect bioenergetics, and applies this information to the design of MAVs.