Steady as they hover: kinematics of kestrel wing and tail morphing during hovering flights.

Wind-hovering birds exhibit remarkable steadiness in flight, achieved through the morphing of their wings and tail. We analysed the kinematics of two nankeen kestrels (Falco cenchroides) engaged in steady wind-hovering flights in a smooth flow wind tunnel. Motion-tracking cameras were used to capture the movements of the birds as they maintained their position. The motion of the birds' head and body, and the morphing motions of their wings and tail were tracked and analysed using correlation methods. The results revealed that wing sweep, representing the flexion/extension movement of the wing, played a significant role in wing motion. Additionally, correlations between different independent degrees of freedom (DoF), including wing and tail coupling, were observed. These kinematic couplings indicate balancing of forces and moments necessary for steady wind hovering. Variation in flight behaviour between the two birds highlighted the redundancy of DoF and the versatility of wing morphing in achieving control. This study provides insights into fixed-wing craft flight control from the avian world and may inspire novel flight control strategies for future fixed-wing aircraft.

Applying the "SOTEC" framework of sociotechnical risk analysis to the development of an autonomous robot swarm for a public cloakroom.

The past decade has seen efforts to develop new forms of autonomous systems with varying applications in different domains, from underwater search and rescue to clinical diagnosis. All of these applications require risk analyses, but such analyses often focus on technical sources of risk without acknowledging its wider systemic and organizational dimensions. In this article, we illustrate this deficit and a way of redressing it by offering a more systematic analysis of the sociotechnical sources of risk in an autonomous system. To this end, the article explores the development, deployment, and operation of an autonomous robot swarm for use in a public cloakroom in light of Macrae's structural, organizational, technological, epistemic, and cultural framework of sociotechnical risk. We argue that this framework provides a useful tool for capturing the complex "nontechnical" dimensions of risk in this domain that might otherwise be overlooked in the more conventional risk analyses that inform regulation and policymaking.

Dynamics of hinged wings in strong upward gusts.

A bird's wings are articulated to its body via highly mobile shoulder joints. The joints confer an impressive range of motion, enabling the wings to make broad, sweeping movements that can modulate quite dramatically the production of aerodynamic load. This is enormously useful in challenging flight environments, especially the gusty, turbulent layers of the lower atmosphere. In this study, we develop a dynamics model to examine how a bird-scale gliding aircraft can use wing-root hinges (analogous to avian shoulder joints) to reject the initial impact of a strong upward gust. The idea requires that the spanwise centre of pressure and the centre of percussion of the hinged wing start, and stay, in good initial alignment (the centre of percussion here is related to the idea of a 'sweet spot' on a bat, as in cricket or baseball). We propose a method for achieving this rejection passively, for which the essential ingredients are (i) appropriate lift and mass distributions; (ii) hinges under constant initial torque; and (iii) a wing whose sections stall softly. When configured correctly, the gusted wings will first pivot on their hinges without disturbing the fuselage of the aircraft, affording time for other corrective actions to engage. We expect this system to enhance the control of aircraft that fly in gusty conditions.

Active breeding seabirds prospect alternative breeding colonies.

Compared to other animal movements, prospecting by adult individuals for a future breeding site is commonly overlooked. Prospecting influences the decision of where to breed and has consequences on fitness and lifetime reproductive success. By analysing movements of 31 satellite- and GPS-tracked gull and tern populations belonging to 14 species in Europe and North America, we examined the occurrence and factors explaining prospecting by actively breeding birds. Prospecting in active breeders occurred in 85.7% of studied species, across 61.3% of sampled populations. Prospecting was more common in populations with frequent inter-annual changes of breeding sites and among females. These results contradict theoretical models which predict that prospecting is expected to evolve in relatively predictable and stable environments. More long-term tracking studies are needed to identify factors affecting patterns of prospecting in different environments and understand the consequences of prospecting on fitness at the individual and population level.

A method for continuous study of soaring and windhovering birds.

Avian flight continues to inspire aircraft designers. Reducing the scale of autonomous aircraft to that of birds and large insects has resulted in new control challenges when attempting to hold steady flight in turbulent atmospheric wind. Some birds, however, are capable of remarkably stable hovering flight in the same conditions. This work describes the development of a wind tunnel configuration that facilitates the study of flapless windhovering (hanging) and soaring bird flight in wind conditions replicating those in nature. Updrafts were generated by flow over replica "hills" and turbulence was introduced through upstream grids, which had already been developed to replicate atmospheric turbulence in prior studies. Successful flight tests with windhovering nankeen kestrels (Falco cenchroides) were conducted, verifying that the facility can support soaring and wind hovering bird flight. The wind tunnel allows the flow characteristics to be carefully controlled and measured, providing great advantages over outdoor flight tests. Also, existing wind tunnels may be readily configured using this method, providing a simpler alternative to the development of dedicated bird flight wind tunnels such as tilting wind tunnels, and the large test section allows for the replication of orographic soaring. This methodology holds promise for future testing investigating the flight behaviour and control responses employed by soaring and windhovering birds.

Quantifying avian inertial properties using calibrated computed tomography.

Estimating centre of mass and mass moments of inertia is an important aspect of many studies in biomechanics. Characterising these parameters accurately in three dimensions is challenging with traditional methods requiring dissection or suspension of cadavers. Here, we present a method to quantify the three-dimensional centre of mass and inertia tensor of birds of prey using calibrated computed tomography (CT) scans. The technique was validated using several independent methods, providing body segment mass estimates within approximately 1% of physical dissection measurements and moment of inertia measurements with a 0.993 R2 correlation with conventional trifilar pendulum measurements. Calibrated CT offers a relatively straightforward, non-destructive approach that yields highly detailed mass distribution data that can be used for three-dimensional dynamics modelling in biomechanics. Although demonstrated here with birds, this approach should work equally well with any animal or appendage capable of being CT scanned.

Dynamic shapes of the zygote and two-cell mouse and human.

Mouse zygote morphokinetics were measured during interphase, the mitotic period, cytokinesis, and two-cell stage. Sequences of rounder-distorted-rounder shapes were revealed, as were changing patterns of cross section area. A calcium chelator and an actin-disrupting agent inhibited the area changes that occurred between pronuclear envelope breakdown and cytokinesis. During cell division, two vortices developed in each nascent cell and they rotated in opposite directions at each end of the cell, a pattern that sometimes persisted for up to 10 h. Exchange with the environment may have been promoted by these shape and area cycles and persisting circulation in the cytoplasm may have a similar function between a cell's interior and periphery. Some of these movements were sporadically also seen in human zygotes with abnormal numbers of pronuclei and the two-cell stages that developed from these compromised human zygotes.

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.

Urban gulls adapt foraging schedule to human-activity patterns.

Numerous animals are able to adapt to temporal patterns in natural food availability, but whether species living in relatively novel environments such as cities can adapt to anthropogenic activity cycles is less well understood. We aimed to assess the extent to which urban gulls have adapted their foraging schedule to anthropogenic food source fluctuations related to human activity by combining field observations at three distinct urban feeding grounds (park, school and waste centre) with global positioning system (GPS) tracking data of gulls visiting similar types of feeding grounds throughout the same city. We found that the birds' foraging patterns closely matched the timing of school breaks and the opening and closing times of the waste centre, but gull activity in the park appeared to correspond to the availability of natural food sources. Overall, this suggests that gulls may have the behavioural flexibility to adapt their foraging behaviour to human time schedules when beneficial and that this trait could potentially enable them to thrive in cities.

Bird wings act as a suspension system that rejects gusts.

Musculoskeletal systems cope with many environmental perturbations without neurological control. These passive preflex responses aid animals to move swiftly through complex terrain. Whether preflexes play a substantial role in animal flight is uncertain. We investigated how birds cope with gusty environments and found that their wings can act as a suspension system, reducing the effects of vertical gusts by elevating rapidly about the shoulder. This preflex mechanism rejected the gust impulse through inertial effects, diminishing the predicted impulse to the torso and head by 32% over the first 80 ms, before aerodynamic mechanisms took effect. For each wing, the centre of aerodynamic loading aligns with the centre of percussion, consistent with enhancing passive inertial gust rejection. The reduced motion of the torso in demanding conditions simplifies crucial tasks, such as landing, prey capture and visual tracking. Implementing a similar preflex mechanism in future small-scale aircraft will help to mitigate the effects of gusts and turbulence without added computational burden.

High aerodynamic lift from the tail reduces drag in gliding raptors.

Many functions have been postulated for the aerodynamic role of the avian tail during steady-state flight. By analogy with conventional aircraft, the tail might provide passive pitch stability if it produced very low or negative lift. Alternatively, aeronautical principles might suggest strategies that allow the tail to reduce inviscid, induced drag: if the wings and tail act in different horizontal planes, they might benefit from biplane-like aerodynamics; if they act in the same plane, lift from the tail might compensate for lift lost over the fuselage (body), reducing induced drag with a more even downwash profile. However, textbook aeronautical principles should be applied with caution because birds have highly capable sensing and active control, presumably reducing the demand for passive aerodynamic stability, and, because of their small size and low flight speeds, operate at Reynolds numbers two orders of magnitude below those of light aircraft. Here, by tracking up to 20,000, 0.3 mm neutrally buoyant soap bubbles behind a gliding barn owl, tawny owl and goshawk, we found that downwash velocity due to the body/tail consistently exceeds that due to the wings. The downwash measured behind the centreline is quantitatively consistent with an alternative hypothesis: that of constant lift production per planform area, a requirement for minimizing viscous, profile drag. Gliding raptors use lift distributions that compromise both inviscid induced drag minimization and static pitch stability, instead adopting a strategy that reduces the viscous drag, which is of proportionately greater importance to lower Reynolds number fliers.

Habitat use of urban-nesting lesser black-backed gulls during the breeding season.

Increasing urbanisation is detrimental for some animal species and potentially advantageous for others. Urban-nesting populations of gulls have undergone rapid population increases worldwide, which has resulted in an increase in human-gull conflicts. In order to inform management and conservation decisions in relation to these populations, more information is needed about the behaviour of these birds in urban settings and how they utilise their environment. This study combined Global Positioning System (GPS) tracking data of 12 urban-nesting lesser black-backed gulls, Larus fuscus, with habitat and behaviour data over three breeding seasons (2016-2018). Despite the proximity of marine areas (~10 km), the birds only made significant use of terrestrial environments, spending two-thirds of their time away from the nest in suburban and urban areas, and one-third in rural green areas. The gulls utilised suburban and urban areas more as their chicks grew and appeared to use diverse foraging strategies to suit different habitats. These results indicate that the range of potential foraging areas available needs to be considered in management decisions and that urban bird populations may not use the resources they are expected to.

Avian surface reconstruction in free flight with application to flight stability analysis of a barn owl and peregrine falcon.

Birds primarily create and control the forces necessary for flight through changing the shape and orientation of their wings and tail. Their wing geometry is characterised by complex variation in parameters such as camber, twist, sweep and dihedral. To characterise this complexity, a multi-view stereo-photogrammetry setup was developed for accurately measuring surface geometry in high resolution during free flight. The natural patterning of the birds was used as the basis for phase correlation-based image matching, allowing indoor or outdoor use while being non-intrusive for the birds. The accuracy of the method was quantified and shown to be sufficient for characterising the geometric parameters of interest, but with a reduction in accuracy close to the wing edge and in some localised regions. To demonstrate the method's utility, surface reconstructions are presented for a barn owl (Tyto alba) and peregrine falcon (Falco peregrinus) during three instants of gliding flight per bird. The barn owl flew with a consistent geometry, with positive wing camber and longitudinal anhedral. Based on flight dynamics theory, this suggests it was longitudinally statically unstable during these flights. The peregrine falcon flew with a consistent glide angle, but at a range of air speeds with varying geometry. Unlike the barn owl, its glide configuration did not provide a clear indication of longitudinal static stability/instability. Aspects of the geometries adopted by both birds appeared to be related to control corrections and this method would be well suited for future investigations in this area, as well as for other quantitative studies into avian flight dynamics.

Head movements quadruple the range of speeds encoded by the insect motion vision system in hawkmoths.

Flying insects use compensatory head movements to stabilize gaze. Like other optokinetic responses, these movements can reduce image displacement, motion and misalignment, and simplify the optic flow field. Because gaze is imperfectly stabilized in insects, we hypothesized that compensatory head movements serve to extend the range of velocities of self-motion that the visual system encodes. We tested this by measuring head movements in hawkmoths Hyles lineata responding to full-field visual stimuli of differing oscillation amplitudes, oscillation frequencies and spatial frequencies. We used frequency-domain system identification techniques to characterize the head's roll response, and simulated how this would have affected the output of the motion vision system, modelled as a computational array of Reichardt detectors. The moths' head movements were modulated to allow encoding of both fast and slow self-motion, effectively quadrupling the working range of the visual system for flight control. By using its own output to drive compensatory head movements, the motion vision system thereby works as an adaptive sensor, which will be especially beneficial in nocturnal species with inherently slow vision. Studies of the ecology of motion vision must therefore consider the tuning of motion-sensitive interneurons in the context of the closed-loop systems in which they function.

Fine-scale flight strategies of gulls in urban airflows indicate risk and reward in city living.

Birds modulate their flight paths in relation to regional and global airflows in order to reduce their travel costs. Birds should also respond to fine-scale airflows, although the incidence and value of this remains largely unknown. We resolved the three-dimensional trajectories of gulls flying along a built-up coastline, and used computational fluid dynamic models to examine how gulls reacted to airflows around buildings. Birds systematically altered their flight trajectories with wind conditions to exploit updraughts over features as small as a row of low-rise buildings. This provides the first evidence that human activities can change patterns of space-use in flying birds by altering the profitability of the airscape. At finer scales still, gulls varied their position to select a narrow range of updraught values, rather than exploiting the strongest updraughts available, and their precise positions were consistent with a strategy to increase their velocity control in gusty conditions. Ultimately, strategies such as these could help unmanned aerial vehicles negotiate complex airflows. Overall, airflows around fine-scale features have profound implications for flight control and energy use, and consideration of this could lead to a paradigm-shift in the way ecologists view the urban environment.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

Vision-based flight control in the hawkmoth Hyles lineata.

Vision is a key sensory modality for flying insects, playing an important role in guidance, navigation and control. Here, we use a virtual-reality flight simulator to measure the optomotor responses of the hawkmoth Hyles lineata, and use a published linear-time invariant model of the flight dynamics to interpret the function of the measured responses in flight stabilization and control. We recorded the forces and moments produced during oscillation of the visual field in roll, pitch and yaw, varying the temporal frequency, amplitude or spatial frequency of the stimulus. The moths' responses were strongly dependent upon contrast frequency, as expected if the optomotor system uses correlation-type motion detectors to sense self-motion. The flight dynamics model predicts that roll angle feedback is needed to stabilize the lateral dynamics, and that a combination of pitch angle and pitch rate feedback is most effective in stabilizing the longitudinal dynamics. The moths' responses to roll and pitch stimuli coincided qualitatively with these functional predictions. The moths produced coupled roll and yaw moments in response to yaw stimuli, which could help to reduce the energetic cost of correcting heading. Our results emphasize the close relationship between physics and physiology in the stabilization of insect flight.

Phospholipase C-ζ-induced Ca2+ oscillations cause coincident cytoplasmic movements in human oocytes that failed to fertilize after intracytoplasmic sperm injection.

OBJECTIVE: To evaluate the imaging of cytoplasmic movements in human oocytes as a potential method to monitor the pattern of Ca(2+) oscillations during activation. DESIGN: Test of a laboratory technique. SETTING: University medical school research laboratory. PATIENT(S): Donated unfertilized human oocytes from intracytoplasmic sperm injection (ICSI) cycles. INTERVENTION(S): Microinjection of oocytes with phospholipase C (PLC) zeta (ζ) cRNA and a Ca(2+)-sensitive fluorescent dye. MAIN OUTCOME MEASURE(S): Simultaneous detection of oocyte cytoplasmic movements using particle image velocimetry (PIV) and of Ca(2+) oscillations using a Ca(2+)-sensitive fluorescent dye. RESULT(S): Microinjection of PLCζ cRNA into human oocytes that had failed to fertilize after ICSI resulted in the appearance of prolonged Ca(2+) oscillations. Each transient Ca(2+) concentration change was accompanied by a small coordinated movement of the cytoplasm that could be detected using PIV analysis. CONCLUSION(S): The occurrence and frequency of cytoplasmic Ca(2+) oscillations, a critical parameter in activating human zygotes, can be monitored by PIV analysis of cytoplasmic movements. This simple method provides a novel, noninvasive approach to determine in real time the occurrence and frequency of Ca(2+) oscillations in human zygotes.

Rhythmic actomyosin-driven contractions induced by sperm entry predict mammalian embryo viability.

Fertilization-induced cytoplasmic flows are a conserved feature of eggs in many species. However, until now the importance of cytoplasmic flows for the development of mammalian embryos has been unknown. Here, by combining a rapid imaging of the freshly fertilized mouse egg with advanced image analysis based on particle image velocimetry, we show that fertilization induces rhythmical cytoplasmic movements that coincide with pulsations of the protrusion forming above the sperm head. We find that these movements are caused by contractions of the actomyosin cytoskeleton triggered by Ca(2+) oscillations induced by fertilization. Most importantly, the relationship between the movements and the events of egg activation makes it possible to use the movements alone to predict developmental potential of the zygote. In conclusion, this method offers, thus far, the earliest and fastest, non-invasive way to predict the viability of eggs fertilized in vitro and therefore can potentially improve greatly the prospects for IVF treatment.

No role for direct touch using the pectoral fins, as an information gathering strategy in a blind fish.

Blind Mexican cave fish (Astyanax fasciatus) lack a functional visual system and have been shown to sense their environment using a technique called hydrodynamic imaging, whereby nearby objects are detected by sensing distortions in the flow field of water around the body using the mechanosensory lateral line. This species has also been noted to touch obstacles, mainly with the pectoral fins, apparently using this tactile information alongside hydrodynamic imaging to sense their surroundings. This study aimed to determine the relative contributions of hydrodynamic and tactile information during wall following behaviour in blind Mexican cave fish. A wall was custom built with a 'netted' region in its centre, which provided very similar tactile information to a solid tank wall, but was undetectable using hydrodynamic imaging. The fish swam significantly closer to and collided more frequently with the netted region of this wall than the solid regions, indicating that the fish did not perceive the netted region as a solid obstacle despite being able to feel it as such with their pectoral fins. We conclude that the touching of objects with the pectoral fins may be an artefact of the intrinsic link between pectoral fin extensions and tail beating whilst swimming, and does not function to gather information. During wall following, hydrodynamic information appears to be used strongly in preference to tactile information in this non-visual system.

The flow fields involved in hydrodynamic imaging by blind Mexican cave fish (Astyanax fasciatus). Part I: open water and heading towards a wall.

Blind Mexican cave fish (Astyanax fasciatus) sense the presence of nearby objects by sensing changes in the water flow around their body. The information available to the fish using this hydrodynamic imaging ability depends on the properties of the flow field it generates while gliding and how this flow field is altered by the presence of objects. Here, we used particle image velocimetry to measure the flow fields around gliding blind cave fish as they moved through open water and when heading towards a wall. These measurements, combined with computational fluid dynamics models, were used to estimate the stimulus to the lateral line system of the fish. Our results showed that there was a high-pressure region around the nose of the fish, low-pressure regions corresponding to accelerated flow around the widest part of the body and a thick laminar boundary layer down the body. When approaching a wall head-on, the changes in the stimulus to the lateral line were confined to approximately the first 20% of the body. Assuming that the fish are sensitive to a certain relative change in lateral line stimuli, it was found that swimming at higher Reynolds numbers slightly decreased the distance at which the fish could detect a wall when approaching head-on, which is the opposite to what has previously been expected. However, when the effects of environmental noise are considered, swimming at higher speed may improve the signal to noise ratio of the stimulus to the lateral line.