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.

Olfactory sampling volume for pheromone capture by wing fanning of silkworm moth: a simulation-based study.

Odours used by insects for foraging and mating are carried by the air. Insects induce airflows around them by flapping their wings, and the distribution of these airflows may strongly influence odour source localisation. The flightless silkworm moth, Bombyx mori, has been a prominent insect model for olfactory research. However, although there have been numerous studies on antenna morphology and its fluid dynamics, neurophysiology, and localisation algorithms, the airflow manipulation of the B. mori by fanning has not been thoroughly investigated. In this study, we performed computational fluid dynamics (CFD) analyses of flapping B. mori to analyse this mechanism in depth. A three-dimensional simulation using reconstructed wing kinematics was used to investigate the effects of B. mori fanning on locomotion and pheromone capture. The fanning of the B. mori was found to generate an aerodynamic force on the scale of its weight through an aerodynamic mechanism similar to that of flying insects. Our simulations further indicate that the B. mori guides particles from its anterior direction within the ~ 60° horizontally by wing fanning. Hence, if it detects pheromones during fanning, the pheromone can be concluded to originate from the direction the head is pointing. The anisotropy in the sampling volume enables the B. mori to orient to the pheromone plume direction. These results provide new insights into insect behaviour and offer design guidelines for robots for odour source localisation.

Robotic flytrap with an ultra-sensitive 'trichome' and fast-response 'lobes'.

Nature abounds with examples of ultra-sensitive perception and agile body transformation for highly efficient predation as well as extraordinary adaptation to complex environments. Flytraps, as a representative example, could effectively detect the most minute physical stimulation of insects and respond instantly, inspiring numerous robotic designs and applications. However, current robotic flytraps face challenges in reproducing the ultra-sensitive insect-touch perception. In addition, fast and fully-covered capture of live insects with robotic flytraps remains elusive. Here we report a novel design of a robotic flytrap with an ultra-sensitive 'trichome' and bistable fast-response 'lobes'. Our results show that the 'trichome' of the proposed robotic flytrap could detect and respond to both the external stimulation of 0.45 mN and a tiny touch of a flying bee with a weight of 0.12 g. Besides, once the 'trichome' is triggered, the bistable 'lobes' could instantly close themselves in 0.2 s to form a fully-covered cage to trap the bees, and reopen to set them free after the tests. We introduce the design, modeling, optimization, and verification of the robotic flytrap, and envision broader applications of this technology in ultra-sensitive perception, fast-response grasping, and biomedical engineering studies.

Modelling take-off moment arms in an ornithocheiraean pterosaur.

Take-off is a vital part of powered flight which likely constrains the size of birds, yet extinct pterosaurs are known to have reached far larger sizes. Three different hypothesised take-off motions (bipedal burst launching, bipedal countermotion launching, and quadrupedal launching) have been proposed as explanations for how pterosaurs became airborne and circumvented this proposed morphological limit. We have constructed a computational musculoskeletal model of a 5 m wingspan ornithocheiraean pterosaur, reconstructing thirty-four key muscles to estimate the muscle moment arms throughout the three hypothesised take-off motions. Range of motion constrained hypothetical kinematic sequences for bipedal and quadrupedal take-off motions were modelled after extant flying vertebrates. Across our simulations we did not find higher hindlimb moment arms for bipedal take-off motions or noticeably higher forelimb moment arms in the forelimb for quadrupedal take-off motions. Despite this, in all our models we found the muscles utilised in the quadrupedal take-off have the largest total launch applicable moment arms throughout the entire take-off sequences and for the take-off pose. This indicates the potential availability of higher leverage for a quadrupedal take-off than hypothesised bipedal motions in pterosaurs pending further examination of muscle forces.

Turkey vultures tune their airspeed to changing air density.

Animals must tune their physical performance to changing environmental conditions, and the breadth of environmental tolerance may contribute to delineating the geographic range of a species. A common environmental challenge that flying animals face is the reduction of air density at high elevation and the reduction in the effectiveness of lift production that accompanies it. As a species, turkey vultures (Cathartes aura) inhabit a >3000 m elevation range, and fly considerably higher, necessitating that they accommodate for a 27% change in air density (0.890 to 1.227 kg m-3) through behavior, physiology or biomechanics. We predicted that birds flying at high elevation would maintain aerodynamic lift performance behaviorally via higher flight speeds, rather than increases in power output or local phenotypic adaptation. We used three-dimensional videography to track turkey vultures flying at three elevations, and data supported the hypothesized negative relationship between median airspeed and air density. Additionally, neither the ratio of horizontal speed to sinking speed nor flapping behavior varied with air density.

Generating controlled gust perturbations using vortex rings.

To understand the locomotory mechanisms of flying and swimming animals, it is often necessary to develop assays that enable us to measure their responses to external gust perturbations. Typically, such measurements have been carried out using a variety of gusts which are difficult to control or characterize owing to their inherently turbulent nature. Here, we present a method of generating discrete gusts under controlled laboratory conditions in the form of a vortex rings which are well-characterized and highly controllable. We also provide the theoretical guidelines underlying the design of gust generators for specific applications. As a case study, we tested the efficacy of this method to study the flight response of freely-flying soldier flies Hermetia illucens. The vortex ring based method can be used to generate controlled gusts to study diverse phenomena ranging from a natural flight in insects to the artificial flight of insect-sized drones and micro-aerial vehicles.

Leading-edge curvature effect on aerodynamic performance of flapping wings in hover and forward flight.

This study investigates the role of leading-edge (LE) curvature in flapping wing aerodynamics considering hovering and forward flight conditions. A scaled-up robotic model is towed along its longitudinal axis by a rack gear carriage system. The forward velocity of the robotic model is changed by varying the advance ratioJfrom 0 (hovering) to 1.0. The study reveals that the LE curvature has insignificant influence on the cycle-average aerodynamic lift and drag. However, the time-history lift coefficient shows that the curvature can enhance the lift around the middle of downstroke. This enhanced lift is reduced from 5% to 1.2% asJchanged from 0 to 1.0. Further flow examinations reveal that the LE curvature is beneficial by enhancing circulation only at the outboard wing sections. The enhanced outboard circulation is found to emanate from the less stretched leading-edge vortices (LEVs), weakened trailing-edge vortices (TEVs), and the coherent merging of the tip vortices (TVs) with the minor LEVs as observed from the phase-lock planar digital particle image velocimetry measurements. The far-wake observation shows that the LE curvature enhances the vorticity within the TV, helping to reduce the overall flow fluctuations in the far field. These findings can be extended to explain the predominantly straight LE wing shape with a small amount of curvature only observed near the wing tip for flapping fliers with Re from 103to 104.

Measurement of wing motion, deformation, and inertial forces of a biomimetic butterfly.

This paper introduces a method for measuring wing motion, deformation, and inertial forces in bio-inspired aircraft research using a camera motion capture system. The method involves placing markers on the wing surface and fitting rigid planes to determine the wing's spatial axis. This allows for describing the wing's rigid motion and obtaining deformation characteristics, such as deflection, twist angle, and gap distance of the forewing and hindwing. An image-based method is proposed for determining wing mass distribution, mass blocks, and mass points for inertial force measurement. The study addresses wing motion, deformation, and inertial force measurement in a real butterfly-like flapping wing vehicle and demonstrates the effectiveness of the approach. The results reveal that inertial forces play a negligible role in the generation of lift peaks and contribute minimal lift during the entire flapping cycle. Furthermore, a transitional phase between downstroke and upstroke is found in flexible wing motion, which has high lift production. This measurement approach offers a rapid and effective solution to experimental challenges in bio-inspired aircraft design and optimization.

Exploration-exploitation model of moth-inspired olfactory navigation.

Navigation of male moths towards females during the mating search offers a unique perspective on the exploration-exploitation (EE) model in decision-making. This study uses the EE model to explain male moth pheromone-driven flight paths. Wind tunnel measurements and three-dimensional tracking using infrared cameras have been leveraged to gain insights into male moth behaviour. During the experiments in the wind tunnel, disturbance to the airflow has been added and the effect of increased fluctuations on moth flights has been analysed, in the context of the proposed EE model. The exploration and exploitation phases are separated using a genetic algorithm to the experimentally obtained dataset of moth three-dimensional trajectories. First, the exploration-to-exploitation rate (EER) increases with distance from the source of the female pheromone is demonstrated, which can be explained in the context of the EE model. Furthermore, our findings reveal a compelling relationship between EER and increased flow fluctuations near the pheromone source. Using an olfactory navigation simulation and our moth-inspired navigation model, the phenomenon where male moths exhibit an enhanced EER as turbulence levels increase is explained. This research extends our understanding of optimal navigation strategies based on general biological EE models and supports the development of bioinspired navigation algorithms.

Wing deformation improves aerodynamic performance of forward flight of bluebottle flies flying in a flight mill.

Insect wings are flexible structures that exhibit deformations of complex spatiotemporal patterns. Existing studies on wing deformation underscore the indispensable role of wing deformation in enhancing aerodynamic performance. Here, we investigated forward flight in bluebottle flies, flying semi-freely in a magnetic flight mill; we quantified wing surface deformation using high-speed videography and marker-less surface reconstruction and studied the effects on aerodynamic forces, power and efficiency using computational fluid dynamics. The results showed that flies' wings exhibited substantial camber near the wing root and twisted along the wingspan, as they were coupled effects of deflection primarily about the claval flexion line. Such deflection was more substantial for supination during the upstroke when most thrust was produced. Compared with deformed wings, the undeformed wings generated 59-98% of thrust and 54-87% of thrust efficiency (i.e. ratio of thrust and power). Wing twist moved the aerodynamic centre of pressure proximally and posteriorly, likely improving aerodynamic efficiency.

Visual lateralization in the sky: Geese manifest visual lateralization when flying with pair mates.

The brain's sensory lateralization involves the processing of information from the sensory organs primarily in one hemisphere. This can improve brain efficiency by reducing interference and duplication of neural circuits. For species that rely on successful interaction among family partners, such as geese, lateralization can be advantageous. However, at the group level, one-sided biases in sensory lateralization can make individuals predictable to competitors and predators. We investigated lateral preferences in the positioning of pair mates of Greater white-fronted geese Anser albifrons albifrons. Using GPS-GSM trackers, we monitored individual geese in flight throughout the year. Our findings indicate that geese exhibit individual lateral biases when viewing their mate in flight, but the direction of these biases varies among individuals. We suggest that these patterns of visual lateralization could be an adaptive trait for the species with long-term social monogamy, high levels of interspecies communication and competition, and high levels of predator and hunting pressure.

A chemo-mechanical constitutive model for muscle activation in bat wing skins.

Birds, bats and insects have evolved unique wing structures to achieve a wide range of flight capabilities. Insects have relatively stiff and passive wings, birds have a complex and hierarchical feathered structure and bats have an articulated skeletal system integrated with a highly stretchable skin. The compliant skin of the wing distinguishes bats from all other flying animals and contributes to bats' remarkable, highly manoeuvrable flight performance and high energetic efficiency. The structural and functional complexity of the bat wing skin is one of the least understood although important elements of the bat flight anatomy. The wing skin has two unusual features: a discrete array of very soft elastin fibres and a discrete array of skeletal muscle fibres. The latter is intriguing because skeletal muscle is typically attached to bone, so the arrangement of intramembranous muscle in soft skin raises questions about its role in flight. In this paper, we develop a multi-scale chemo-mechanical constitutive model for bat wing skin. The chemo-mechanical model links cross-bridge cycling to a structure-based continuum model that describes the active viscoelastic behaviour of the soft anisotropic skin tissue. Continuum models at the tissue length-scale are valuable as they are easily implemented in commercial finite element codes to solve problems involving complex geometries, loading and boundary conditions. The constitutive model presented in this paper will be used in detailed finite element simulations to improve our understanding of the mechanics of bat flight in the context of wing kinematics and aerodynamic performance.

Intraspecific variation in the pterosaur Rhamphorhynchus muensteri-implications for flight and socio-sexual signaling.

Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus, which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals.

Transfer learning may explain pigeons' ability to detect cancer in histopathology.

Pigeons' unexpected competence in learning to categorize unseen histopathological images has remained an unexplained discovery for almost a decade (Levensonet al2015PLoS One10e0141357). Could it be that knowledge transferred from their bird's-eye views of the earth's surface gleaned during flight contributes to this ability? Employing a simulation-based verification strategy, we recapitulate this biological phenomenon with a machine-learning analog. We model pigeons' visual experience during flight with the self-supervised pre-training of a deep neural network on BirdsEyeViewNet; our large-scale aerial imagery dataset. As an analog of the differential food reinforcement performed in Levensonet al's study 2015PLoS One10e0141357), we apply transfer learning from this pre-trained model to the same Hematoxylin and Eosin (H&E) histopathology and radiology images and tasks that the pigeons were trained and tested on. The study demonstrates that pre-training neural networks with bird's-eye view data results in close agreement with pigeons' performance. These results support transfer learning as a reasonable computational model of pigeon representation learning. This is further validated with six large-scale downstream classification tasks using H&E stained whole slide image datasets representing diverse cancer types.

Effects of age on oxidative stress and locomotion in the pollinator, Megachile rotundata.

Despite numerous aging studies, the relationship between oxidative stress, aging, and decline in functions such as locomotion is still debated. Insects offer a promising model for analyzing the relationship between oxidative stress and aging, because they exhibit vast differences in lifespan that may be affected by the environment, social factors, levels of activity, and aging interventions. In this study, we explore the effects of aging on oxidative stress and locomotion using the pollinator, Megachile rotundata, a species that is very mobile and active in the adult stage. Across the adult lifespan of M. rotundata, we assessed changes in walking, flight, oxidative damage, and antioxidant defenses. Our results suggest that M. rotundata experience age-related declines in flight, but not walking. Additionally, we found that oxidative damage and antioxidant capacity initially increase with age and physical activity, but then levels are maintained. Overall, these data show that M. rotundata, like some other organisms, may not perfectly follow the free radical theory of aging.

Visually guided swarm motion coordination via insect-inspired small target motion reactions.

Despite progress developing experimentally-consistent models of insect in-flight sensing and feedback for individual agents, a lack of systematic understanding of the multi-agent and group performance of the resulting bio-inspired sensing and feedback approaches remains a barrier to robotic swarm implementations. This study introduces the small-target motion reactive (STMR) swarming approach by designing a concise engineering model of the small target motion detector (STMD) neurons found in insect lobula complexes. The STMD neuron model identifies the bearing angle at which peak optic flow magnitude occurs, and this angle is used to design an output feedback switched control system. A theoretical stability analysis provides bi-agent stability and state boundedness in group contexts. The approach is simulated and implemented on ground vehicles for validation and behavioral studies. The results indicate despite having the lowest connectivity of contemporary approaches (each agent instantaneously regards only a single neighbor), STMR achieves collective group motion. STMR group level metric analysis also highlights continuously varying polarization and decreasing heading variance.

Short communication: Maintained visual performance in birds under high altitude hypoxia.

Birds are highly dependent on their vision for orientation and navigation. The avian eye differs from the mammalian eye as the retina is avascular, leaving the inner, highly metabolically active layers with a very long diffusion distance to the oxygen supply. During flight at high altitudes, birds face a decrease in environmental oxygen partial pressure, which leads to a decrease in arterial oxygen levels. Since oxygen perfusion to the retina is already limited in birds, we hypothesize that visual function is impaired by low oxygen availability. However, the visual performance of birds exposed to hypoxia has not been evaluated before. Here, we assess the optomotor response (OMR) in zebra finches under simulated high-altitude hypoxia (10%) and show that the OMR is largely maintained under hypoxia with only a modest reduction in OMR, demonstrating that birds can largely maintain visual function at high altitudes. The method of our study does not provide insight into the mechanisms involved, but our findings suggest that birds have evolved physiological mechanisms for retinal function at low tissue oxygen levels.

Flock2: A model for orientation-based social flocking.

The aerial flocking of birds, or murmurations, has fascinated observers while presenting many challenges to behavioral study and simulation. We examine how the periphery of murmurations remain well bounded and cohesive. We also investigate agitation waves, which occur when a flock is disturbed, developing a plausible model for how they might emerge spontaneously. To understand these behaviors a new model is presented for orientation-based social flocking. Previous methods model inter-bird dynamics by considering the neighborhood around each bird, and introducing forces for avoidance, alignment, and cohesion as three dimensional vectors that alter acceleration. Our method introduces orientation-based social flocking that treats social influences from neighbors more realistically as a desire to turn, indirectly controlling the heading in an aerodynamic model. While our model can be applied to any flocking social bird we simulate flocks of starlings, Sturnus vulgaris, and demonstrate the possibility of orientation waves in the absence of predators. Our model exhibits spherical and ovoidal flock shapes matching observation. Comparisons of our model to Reynolds' on energy consumption and frequency analysis demonstrates more realistic motions, significantly less energy use in turning, and a plausible mechanism for emergent orientation waves.

Gliding toward an understanding of the origin of flight in bats.

Bats are the only mammals capable of powered flight and have correspondingly specialized body plans, particularly in their limb morphology. The origin of bat flight is still not fully understood due to an uninformative fossil record but, from the perspective of a functional transition, it is widely hypothesized that bats evolved from gliding ancestors. Here, we test predictions of the gliding-to-flying hypothesis of the origin of bat flight by using phylogenetic comparative methods to model the evolution of forelimb and hindlimb traits on a dataset spanning four extinct bats and 231 extant mammals with diverse locomotor modes. Our results reveal that gliders exhibit adaptive trait optima (1) toward relatively elongate forelimbs that are intermediate between those of bats and non-gliding arborealists, and (2) toward relatively narrower but not longer hindlimbs that are intermediate between those of non-gliders and bats. We propose an adaptive landscape based on limb length and width optimal trends derived from our modeling analyses. Our results support a hypothetical evolutionary pathway wherein glider-like postcranial morphology precedes a bat-like morphology adapted to powered-flight, setting a foundation for future developmental, biomechanical, and evolutionary research to test this idea.

Insect ecology: Tracking migratory butterflies through pollen.

Painted ladies are well-known migratory butterflies, but confirmation of the details of their swarming flights through Europe has evaded scientists until now. It was their role as pollinators, carrying pollen grains on their flights, that helped unlock the secrets of their migrations.