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.

Tailbeat perturbations improve swimming efficiency by reducing the phase lag between body motion and the resulting fluid response.

Understanding how animals swim efficiently and generate high thrust in complex fluid environments is of considerable interest to researchers in various fields, including biology, physics, and engineering. However, the influence of often-overlooked perturbations on swimming fish remains largely unexplored. Here, we investigate the propulsion generated by oscillating tailbeats with superimposed rhythmic perturbations of high frequency and low amplitude. We reveal, using a combination of experiments in a biomimetic fish-like robotic platform, computational fluid dynamics simulations, and theoretical analysis, that rhythmic perturbations can significantly increase both swimming efficiency and thrust production. The introduction of perturbations increases pressure-induced thrust, while reduced phase lag between body motion and the subsequent fluid dynamics response improves swimming efficiency. Moreover, our findings suggest that beneficial perturbations are sensitive to kinematic parameters, resolving previous conflicts regarding the effects of such perturbations. Our results highlight the potential benefits of introducing perturbations in propulsion generators, providing potential hypotheses for living systems and inspiring the design of artificial flapping-based propulsion systems.

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.

Controlling a bio-inspired miniature blimp using a depth sensing neural-network camera.

Miniature blimps are lighter-than-air vehicles which have become an increasingly common unmanned aerial system research platform due to their extended endurance and collision tolerant design. The UNSW-C bio-inspired miniature blimp consists of a 0.5 m spherical mylar envelope filled with helium. Four fins placed along the equator provide control over the three translatory axes and yaw rotations. A gondola attached to the bottom of the blimp contains all the electronics and flight controller. Here, we focus on using the UNSW-C blimp as a platform to achieve autonomous flight in GPS-denied environments. The majority of unmanned flying systems rely on GPS or multi-camera motion capture systems for position and orientation estimation. However, such systems are expensive, difficult to set up and not compact enough to be deployed in real environments. Instead, we seek to achieve basic flight autonomy for the blimp using a low-priced and portable solution. We make use of a low-cost embedded neural network stereoscopic camera (OAK-D-PoE) for detecting and positioning the blimp while an onboard inertia measurement unit was used for orientation estimation. Flight tests and analysis of trajectories revealed that 3D position hold as well as basic waypoint navigation could be achieved with variance (<0.1 m). This performance was comparable to that when a conventional multi-camera positioning system (VICON) was used for localizing the blimp. Our results highlight the potentially favorable tradeoffs offered by such low-cost positioning systems in extending the operational domain of unmanned flight systems when direct line of sight is available.

Functional anatomy of the worker honeybee stinger (Apis mellifera).

The honeybee stinger is a powerful defense mechanism that combines painful venom, a subcutaneous delivery system, and the ability to autotomize. It is a complex organ and to function autonomously it must carry with it all the anatomical components required to operate. In this study, we combined high-speed filming, SEM imagery, and micro-CT for volumetric rendering of the stinger with a synthesis of existing literature. We present a comprehensive description of all components, including cuticular elements, musculature, nervous and glandular tissue using updated imagery. We draw from the Hymenoptera literature to make interspecific comparisons where relevant. The use of 3D reconstruction allows us to separate stinger components and present the first 3D renders of the bee stinger including the terminal abdominal ganglion and its projections. It also clarifies the in-situ geometry of the valves within the bulb and the spatial relationships among the accessory plates and accompanying musculature.

Passive dynamics regulates aperiodic transitions in flapping wing systems.

Natural and artificial flapping wing flyers generally do not exhibit chaos or aperiodic dynamic modes, though several experimental and numerical studies with canonical models of flapping foils have reported inevitable chaotic transition at high ranges of dynamic plunge velocity ( κ h ). Here we considered the idealized case of a pitching-plunging flapping foil and numerically investigated the effects of passive pitching dynamics on the fluid forces and dynamical states, and compared it with a fully actuated wing. We found that in comparison to fully actuated foils, aperiodic transition can be avoided even for high κ h when passive oscillations are allowed. Passive pitching modulated the relative foil orientation with respect to the incoming free stream to maintain a lower effective angle-of-attack throughout the stroke and reduced the leading-edge-vortex (LEV) strength. Absence of aperiodic triggers such as flow separation and strong LEVs keep the wake periodic, and chaotic transition is averted. In the presence of fluctuating inflow conditions, passive pitching attenuated the fluid loads experienced by the airfoil thus improving the wing's gust mitigating potential. These findings highlight the favorable properties of passive dynamics in regularizing aerodynamic loads on flapping wing systems and presents viable solutions for artificial flying platforms.

Bumblebees display characteristics of active vision during robust obstacle avoidance flight.

Insects are remarkable flyers and capable of navigating through highly cluttered environments. We tracked the head and thorax of bumblebees freely flying in a tunnel containing vertically oriented obstacles to uncover the sensorimotor strategies used for obstacle detection and collision avoidance. Bumblebees presented all the characteristics of active vision during flight by stabilizing their head relative to the external environment and maintained close alignment between their gaze and flightpath. Head stabilization increased motion contrast of nearby features against the background to enable obstacle detection. As bees approached obstacles, they appeared to modulate avoidance responses based on the relative retinal expansion velocity (RREV) of obstacles and their maximum evasion acceleration was linearly related to RREVmax. Finally, bees prevented collisions through rapid roll manoeuvres implemented by their thorax. Overall, the combination of visuo-motor strategies of bumblebees highlights elegant solutions developed by insects for visually guided flight through cluttered environments.

Effects of uniform vertical inflow perturbations on the performance of flapping wings.

Flapping wings have attracted significant interest for use in miniature unmanned flying vehicles. Although numerous studies have investigated the performance of flapping wings under quiescent conditions, effects of freestream disturbances on their performance remain under-explored. In this study, we experimentally investigated the effects of uniform vertical inflows on flapping wings using a Reynolds-scaled apparatus operating in water at Reynolds number ≈ 3600. The overall lift and drag produced by a flapping wing were measured by varying the magnitude of inflow perturbation from J Vert = -1 (downward inflow) to J Vert = 1 (upward inflow), where J Vert is the ratio of the inflow velocity to the wing's velocity. The interaction between flapping wing and downward-oriented inflows resulted in a steady linear reduction in mean lift and drag coefficients, C ¯ L and C ¯ D , with increasing inflow magnitude. While a steady linear increase in C ¯ L and C ¯ D was noted for upward-oriented inflows between 0 < J Vert < 0.3 and J Vert > 0.7, a significant unsteady wing-wake interaction occurred when 0.3 ≤ J Vert < 0.7, which caused large variations in instantaneous forces over the wing and led to a reduction in mean performance. These findings highlight asymmetrical effects of vertically oriented perturbations on the performance of flapping wings and pave the way for development of suitable control strategies.

Performance of passively pitching flapping wings in the presence of vertical inflows.

The successful implementation of passively pitching flapping wings strongly depends on their ability to operate efficiently in wind disturbances. In this study, we experimentally investigated the interaction between a uniform vertical inflow perturbation and a passive-pitching flapping wing using a Reynolds-scaled apparatus operating in water at Reynolds number ≈3600. A parametric study was performed by systematically varying the Cauchy number (Ch) of the wings from 0.09 to 11.52. The overall lift and drag, and pitch angle of the wing were measured by varying the magnitude of perturbation fromJVert= -0.6 (downward inflow) toJVert= 0.6 (upward inflow) at eachCh, whereJVertis the ratio of the inflow velocity to the wing's velocity. We found that the lift and drag had remarkably different characteristics in response to bothChandJVert. Across allCh, while mean lift tended to increase as the inflow perturbation varied from -0.6 to 0.6, drag was significantly less sensitive to the perturbation. However effect of the vertical inflow on drag was dependent onCh, where it tended to vary from an increasing to a decreasing trend asChwas changed from 0.09 to 11.52. The differences in the lift and drag with perturbation magnitude could be attributed to the reorientation of the net force over the wing as a result of the interaction with the perturbation. These results highlight the complex interactions between passively pitching flapping wings and freestream perturbations and will guide the design of miniature flying crafts with such architectures.

Visual and movement memories steer foraging bumblebees along habitual routes.

One persistent question in animal navigation is how animals follow habitual routes between their home and a food source. Our current understanding of insect navigation suggests an interplay between visual memories, collision avoidance and path integration, the continuous integration of distance and direction travelled. However, these behavioural modules have to be continuously updated with instantaneous visual information. In order to alleviate this need, the insect could learn and replicate habitual movements ('movement memories') around objects (e.g. a bent trajectory around an object) to reach its destination. We investigated whether bumblebees, Bombus terrestris, learn and use movement memories en route to their home. Using a novel experimental paradigm, we habituated bumblebees to establish a habitual route in a flight tunnel containing 'invisible' obstacles. We then confronted them with conflicting cues leading to different choice directions depending on whether they rely on movement or visual memories. The results suggest that they use movement memories to navigate, but also rely on visual memories to solve conflicting situations. We investigated whether the observed behaviour was due to other guidance systems, such as path integration or optic flow-based flight control, and found that neither of these systems was sufficient to explain the behaviour.

Using a robotic platform to study the influence of relative tailbeat phase on the energetic costs of side-by-side swimming in fish.

A potential benefit of swimming together in coordinated schools is to allow fish to extract energy from vortices shed by their neighbours, thus reducing the costs of locomotion. This hypothesis has been very hard to test in real fish schools, and it has proven very difficult to replicate the complex hydrodynamics at relevant Reynolds numbers using computational simulations. A complementary approach, and the one we adopt here, is to develop and analyse the performance of biomimetic autonomous robotic models that capture the salient kinematics of fish-like swimming, and also interact via hydrodynamic forces. We developed bio-inspired robotic fish which perform sub-carangiform locomotion, and measured the speed and power consumption of robots when swimming in isolation and when swimming side-by-side in pairs. We found that swimming side-by-side confers a substantial increase in both the speed and efficiency of locomotion of both fish regardless of the relative phase relationship of their body undulations. However, we also find that each individual can slightly increase their own power efficiency if they change relative tailbeat phase by approximately 0.25π with respect to, and at the energetic expense of, their neighbour. This suggests the possibility of a competitive game-theoretic dynamic between individuals in swimming groups. Our results also demonstrate the potential applicability of our platform, and provide a natural connection between the biology and robotics of collective motion.

Bumblebees perceive the spatial layout of their environment in relation to their body size and form to minimize inflight collisions.

Animals that move through complex habitats must frequently contend with obstacles in their path. Humans and other highly cognitive vertebrates avoid collisions by perceiving the relationship between the layout of their surroundings and the properties of their own body profile and action capacity. It is unknown whether insects, which have much smaller brains, possess such abilities. We used bumblebees, which vary widely in body size and regularly forage in dense vegetation, to investigate whether flying insects consider their own size when interacting with their surroundings. Bumblebees trained to fly in a tunnel were sporadically presented with an obstructing wall containing a gap that varied in width. Bees successfully flew through narrow gaps, even those that were much smaller than their wingspans, by first performing lateral scanning (side-to-side flights) to visually assess the aperture. Bees then reoriented their in-flight posture (i.e., yaw or heading angle) while passing through, minimizing their projected frontal width and mitigating collisions; in extreme cases, bees flew entirely sideways through the gap. Both the time that bees spent scanning during their approach and the extent to which they reoriented themselves to pass through the gap were determined not by the absolute size of the gap, but by the size of the gap relative to each bee's own wingspan. Our findings suggest that, similar to humans and other vertebrates, flying bumblebees perceive the affordance of their surroundings relative their body size and form to navigate safely through complex environments.

Kinematics of gecko climbing: the lateral undulation pattern.

Geckos are exceptional at terrestrial locomotion and can move on diverse terrains and surface orientations. Geckos employ cyclical lateral bending of their flexible trunk and tail to coordinate their limb movements. In this study, using an optical motion capture system, we measured the kinematics of this lateral undulation pattern of geckos (Gekko gecko) at increasing locomotion velocity on horizontal plane, 45° inclined plane and vertical plane, respectively. We observed that geckos increased their stride frequency and stride length to increase the locomotion velocity; the effect of stride frequency on the locomotion velocity was greater than that of stride length. With increasing speed, the lateral undulation pattern changed from standing to travelling. The waveform of the trunk movement appeared as single-peak curves in a standing wave at low speeds and was propagated from head to tail in a travelling wave at high speeds. Analysis of the anatomical characteristics and axial angular kinematics of the two patterns revealed that the lateral undulation pattern results from girdle rotation and axial muscle activity. Thus, the travelling wave is the combined effect of the lateral trunk bending and deflection of the body relative to the motion direction.

CFD based parameter tuning for motion control of robotic fish.

After millions of years of evolution, fishes have been endowed with agile swimming ability to accomplish various behaviourally relevant tasks. In comparison, robotic fish are still quite poor swimmers. One of the unique challenges facing robotic fish is the difficulty in tuning the motion control parameters on the robot directly. This is mainly due to the complex fluid environment robotic fish need to contend with and endurance limitations (i.e. battery capacity limitations). To overcome these limitations, we propose a computational fluid dynamics (CFD) simulation platform to first tune the motion control parameters for the computational robotic fish and then refine the parameters by experiments on robotic fish. Within the simulation platform, the body morphology and gait control of the computational robotic fish are designed according to a robotic fish. The gait control is implemented by a central pattern generator (CPG); The CFD model is solved by using a hydrodynamic-kinematics strong-coupling method. We tested our simulation platform with three basic tasks under active disturbance rejection control (ADRC) and try-and-error-based parameter tuning. Trajectory comparisons between the computational robotic fish and robotic fish verify the effectiveness of our simulation platform. Moreover, power costs and swimming efficiency under the motion control are also analyzed based on the outputs from the simulation platform. Our results indicate that the CFD based simulation platform is powerful and robust, and shed new light on the efficient design and parameter optimization of the motion control of robotic fish.

Modulation of Flight Muscle Recruitment and Wing Rotation Enables Hummingbirds to Mitigate Aerial Roll Perturbations.

Both biological and artificial fliers must contend with aerial perturbations that are ubiquitous in the outdoor environment. Flapping fliers are generally least stable but also most maneuverable around the roll axis, yet our knowledge of roll control in biological fliers remains limited. Hummingbirds are suitable models for linking aerodynamic perturbations to flight control strategies, as these small, powerful fliers are capable of remaining airborne even in adverse wind conditions. We challenged hummingbirds to fly within a steady, longitudinally (streamwise) oriented vortex that imposed a continuous roll perturbation, measured wing kinematics and neuromotor activation of the flight muscles with synchronized high-speed video and electromyography and used computational fluid dynamics (CFD) to estimate the aerodynamic forces generated by observed wing motions. Hummingbirds responded to the perturbation with bilateral differences in activation of the main flight muscles while maintaining symmetry in most major aspects of wing motion, including stroke amplitude, stroke plane angle, and flapping frequency. Hummingbirds did display consistent bilateral differences in subtler wing kinematic traits, including wing rotation and elevation. CFD modeling revealed that asymmetric wing rotation was critical for attenuating the effects of the perturbation. The birds also augmented flight stabilization by adjusting body and tail posture to expose greater surface area to upwash than to the undesirable downwash. Our results provide insight into the remarkable capacity of hummingbirds to maintain flight control, as well as bio-inspiration for simple yet effective control strategies that could allow robotic fliers to contend with unfamiliar and challenging real-world aerial conditions.

Bottom-level motion control for robotic fish to swim in groups: modeling and experiments.

Moving in groups is an amazing spectacle of collective behaviour in fish and has attracted considerable interest from many fields, including biology, physics and engineering. Although robotic fish have been well studied, including algorithms to simulate group swimming, experiments that demonstrate multiple robotic fish as a stable group are yet to be achieved. One of the challenges is the lack of a robust bottom-level motion control system for robotic fish platforms. Here we seek to overcome this challenge by focusing on the design and implementation of a motion controller for robotic fish that allows multiple individuals to swim in groups. As direction control is essential in motion control, we first propose a high-accuracy controller which can control a sub-carangiform robotic fish from one arbitrary position/pose (position and direction) to another. We then develop a hydrodynamic-model-based simulation platform to expedite the process of the parameter tuning of the controller. The accuracy of the simulation platform was assessed by comparing the results from experiments on a robotic fish using speeding and turning tests. Subsequently, extensive simulations and experiments with robotic fish were used to verify the accuracy and robustness of the bottom-level motion control. Finally, we demonstrate the efficacy of our controller by implementing group swimming using three robotic fish swimming freely in prescribed trajectories. Although the fluid environment can be complex during group swimming, our bottom-level motion control remained nominally accurate and robust. This motion control strategy lays a solid foundation for further studies of group swimming with multiple robotic fish.

Gap perception in bumblebees.

A number of insects fly over long distances below the natural canopy, where the physical environment is highly cluttered consisting of obstacles of varying shape, size and texture. While navigating within such environments, animals need to perceive and disambiguate environmental features that might obstruct their flight. The most elemental aspect of aerial navigation through such environments is gap identification and 'passability' evaluation. We used bumblebees to seek insights into the mechanisms used for gap identification when confronted with an obstacle in their flight path and behavioral compensations employed to assess gap properties. Initially, bumblebee foragers were trained to fly though an unobstructed flight tunnel that led to a foraging chamber. After the bees were familiar with this situation, we placed a wall containing a gap that unexpectedly obstructed the flight path on a return trip to the hive. The flight trajectories of the bees as they approached the obstacle wall and traversed the gap were analyzed in order to evaluate their behavior as a function of the distance between the gap and a background wall that was placed behind the gap. Bumblebees initially decelerated when confronted with an unexpected obstacle. Deceleration was first noticed when the obstacle subtended around 35 deg on the retina but also depended on the properties of the gap. Subsequently, the bees gradually traded off their longitudinal velocity to lateral velocity and approached the gap with increasing lateral displacement and lateral velocity. Bumblebees shaped their flight trajectory depending on the salience of the gap, indicated in our case by the optic flow contrast between the region within the gap and on the obstacle, which decreased with decreasing distance between the gap and the background wall. As the optic flow contrast decreased, the bees spent an increasing amount of time moving laterally across the obstacles. During these repeated lateral maneuvers, the bees are probably assessing gap geometry and passability.

Bees with attitude: the effects of directed gusts on flight trajectories.

Flight is a complicated task at the centimetre scale particularly due to unsteady air fluctuations which are ubiquitous in outdoor flight environments. Flying organisms deal with these difficulties using active and passive control mechanisms to steer their body motion. Body attitudes of flapping organisms are linked with their resultant flight trajectories and performance, yet little is understood about how isolated unsteady aerodynamic phenomena affect the interlaced dynamics of such systems. In this study, we examined freely flying bumblebees subject to a single isolated gust to emulate aerodynamic disturbances encountered in nature. Bumblebees are expert commanders of the aerial domain as they persistently forage within complex terrain elements. By tracking the three-dimensional dynamics of bees flying through gusts, we determined the sequences of motion that permit flight in three disturbance conditions: sideward, upward and downward gusts. Bees executed a series of passive impulsive maneuvers followed by active recovery maneuvers. Impulsive motion was unique in each gust direction, maintaining control by passive manipulation of the body. Bees pitched up and slowed down at the beginning of recovery in every disturbance, followed by corrective maneuvers which brought body attitudes back to their original state. Bees were displaced the most by the sideward gust, displaying large lateral translations and roll deviations. Upward gusts were easier for bees to fly through, causing only minor flight changes and minimal recovery times. Downward gusts severely impaired the control response of bees, inflicting strong adverse forces which sharply upset trajectories. Bees used a variety of control strategies when flying in each disturbance, offering new insights into insect-scale flapping flight and bio-inspired robotic systems.This article has an associated First Person interview with the first author of the paper.

Bumblebees minimize control challenges by combining active and passive modes in unsteady winds.

The natural wind environment that volant insects encounter is unsteady and highly complex, posing significant flight-control and stability challenges. It is critical to understand the strategies insects employ to safely navigate in natural environments. We combined experiments on free flying bumblebees with high-fidelity numerical simulations and lower-order modeling to identify the mechanics that mediate insect flight in unsteady winds. We trained bumblebees to fly upwind towards an artificial flower in a wind tunnel under steady wind and in a von Kármán street formed in the wake of a cylinder. Analysis revealed that at lower frequencies in both steady and unsteady winds the bees mediated lateral movement with body roll - typical casting motion. Numerical simulations of a bumblebee in similar conditions permitted the separation of the passive and active components of the flight trajectories. Consequently, we derived simple mathematical models that describe these two motion components. Comparison between the free-flying live and modeled bees revealed a novel mechanism that enables bees to passively ride out high-frequency perturbations while performing active maneuvers at lower frequencies. The capacity of maintaining stability by combining passive and active modes at different timescales provides a viable means for animals and machines to tackle the challenges posed by complex airflows.