Submicron drops from flapping bursting bubbles.

Tiny water drops produced from bubble bursting play a critical role in forming clouds, scattering sunlight, and transporting pathogens from water to the air. Bubbles burst by nucleating a hole at their cap foot and may produce jets or film drops. The latter originate from the fragmentation of liquid ligaments formed by the centripetal destabilization of the opening hole rim. They constitute a major fraction of the aerosols produced from bubbles with cap radius of curvature (R) > ∼0.4 × capillary length (a). However, our present understanding of the corresponding mechanisms does not explain the production of most submicron film drops, which represent the main number fraction of sea spray aerosols. In this study, we report observations showing that bursting bubbles with R < ∼0.4a are actually mainly responsible for submicron film drop production, through a mechanism involving the flapping shear instability of the cap with the outer environment. With this proposed pathway, the complex relations between bubble size and number of drops produced per bubble can be better explained, providing a fundamental framework for understanding the production flux of aerosols and the transfer of substances mediated by bubble bursting through the air-water interface and the sensitivity of the process to the nature of the environment.

Musculoskeletal wing-actuation model of hummingbirds predicts diverse effects of primary flight muscles in hovering flight.

Hummingbirds have evolved to hover and manoeuvre with exceptional flight control. This is enabled by their musculoskeletal system that successfully exploits the agile motion of flapping wings. Here, we synthesize existing empirical and modelling data to generate novel hypotheses for principles of hummingbird wing actuation. These may help guide future experimental work and provide insights into the evolution and robotic emulation of hummingbird flight. We develop a functional model of the hummingbird musculoskeletal system, which predicts instantaneous, three-dimensional torque produced by primary (pectoralis and supracoracoideus) and combined secondary muscles. The model also predicts primary muscle contractile behaviour, including stress, strain, elasticity and work. Results suggest that the primary muscles (i.e. the flight 'engine') function as diverse effectors, as they do not simply power the stroke, but also actively deviate and pitch the wing with comparable actuation torque. The results also suggest that the secondary muscles produce controlled-tightening effects by acting against primary muscles in deviation and pitching. The diverse effects of the pectoralis are associated with the evolution of a comparatively enormous bicipital crest on the humerus.

Kinematic and hydrodynamic analyses of turning manoeuvres in penguins: body banking and wing upstroke generate centripetal force.

Penguins perform lift-based swimming by flapping their wings. Previous kinematic and hydrodynamic studies have revealed the basics of wing motion and force generation in penguins. Although these studies have focused on steady forward swimming, the mechanism of turning manoeuvres is not well understood. In this study, we examined the horizontal turning of penguins via 3D motion analysis and quasi-steady hydrodynamic analysis. Free swimming of gentoo penguins (Pygoscelis papua) at an aquarium was recorded, and body and wing kinematics were analysed. In addition, quasi-steady calculations of the forces generated by the wings were performed. Among the selected horizontal swimming manoeuvres, turning was distinguished from straight swimming by the body trajectory for each wingbeat. During the turns, the penguins maintained outward banking through a wingbeat cycle and utilized a ventral force during the upstroke as a centripetal force to turn. Within a single wingbeat during the turns, changes in the body heading and bearing also mainly occurred during the upstroke, while the subsequent downstroke accelerated the body forward. We also found contralateral differences in the wing motion, i.e. the inside wing of the turn became more elevated and pronated. Quasi-steady calculations of the wing force confirmed that the asymmetry of the wing motion contributes to the generation of the centripetal force during the upstroke and the forward force during the downstroke. The results of this study demonstrate that the hydrodynamic force of flapping wings, in conjunction with body banking, is actively involved in the mechanism of turning manoeuvres in penguins.

Reconstruction of Flight Parameters of a Bat for Flapping Robots.

The flight of bats is comparatively less documented and understood than birds and insects and may provide novel inspiration for the design of flapping flight robots. This study captured the natural flight of short-nosed fruit bats (Cynopterus sphinx) by an optical motion capture system, "OptiTrack", with pasted markers on the wings and body to reconstruct the flight parameters. Due to the self-occlusion at some moments, points on the membrane wings cannot be captured by any cameras. To draw a smooth trajectory, it is desired to reconstruct all missing data. Therefore, an algorithm is proposed by using numerical techniques, accompanied by modern mathematical and computational tools, to envisage the missing data from the captured flight. The least-square fitted polynomial engendered the parameter equations for x-, y-, and z-coordinates of marked points which were used to reconstruct the trajectory of the flight. The parameter equations of position coordinates were also used to compute the morphological and aerodynamic characteristics of the flight. The most outstanding contribution of the work is that not only the trajectory, velocity, and velocity field but also the morphing areas of the membrane wings were recreated using the reconstructed data. These data and reconstructed curves of trajectory and velocity field will be used for the further aerodynamic analysis and mechanism design of the flapping robot. This method can also be generalized to reconstruct the performance parameters of any other animals for bionic design.

Flow interactions between uncoordinated flapping swimmers give rise to group cohesion.

Many species of fish and birds travel in groups, yet the role of fluid-mediated interactions in schools and flocks is not fully understood. Previous fluid-dynamical models of these collective behaviors assume that all individuals flap identically, whereas animal groups involve variations across members as well as active modifications of wing or fin motions. To study the roles of flapping kinematics and flow interactions, we design a minimal robotic "school" of two hydrofoils swimming in tandem. The flapping kinematics of each foil are independently prescribed and systematically varied, while the forward swimming motions are free and result from the fluid forces. Surprisingly, a pair of uncoordinated foils with dissimilar kinematics can swim together cohesively-without separating or colliding-due to the interaction of the follower with the wake left by the leader. For equal flapping frequencies, the follower experiences stable positions in the leader's wake, with locations that can be controlled by flapping amplitude and phase. Further, a follower with lower flapping speed can defy expectation and keep up with the leader, whereas a faster-flapping follower can be buffered from collision and oscillate in the leader's wake. We formulate a reduced-order model which produces remarkable agreement with all experimentally observed modes by relating the follower's thrust to its flapping speed relative to the wake flow. These results show how flapping kinematics can be used to control locomotion within wakes, and that flow interactions provide a mechanism which promotes group cohesion.

Physiological Signatures of Changes in Honeybee's Central Complex During Wing Flapping.

Many kinds of locomotion abilities of insects-including flight control, spatial orientation memory, position memory, angle information integration, and polarized light guidance are considered to be related to the central complex. However, evidence was still not sufficient to support those conclusions from the aspect of neural basis. For the locomotion form of wing flapping, little is known about the patterns of changes in brain activity of the central complex during movement. Here, we analyze the changes in honeybees' neuronal population firing activity of central complex and optic lobes with the perspectives of energy and nonlinear changes. Although the specific function of the central complex remains unknown, evidence suggests that its neural activities change remarkably during wing flapping and its delta rhythm is dominative. Together, our data reveal that the firing activity of some of the neuronal populations of the optic lobe shows reduction in complexity during wing flapping. Elucidating the brain activity changes during a flapping period of insects promotes our understanding of the neuro-mechanisms of insect locomotor control, thus can inspire the fine control of insect cyborgs.

Kinematics and hydrodynamics analyses of swimming penguins: wing bending improves propulsion performance.

Penguins are adapted to underwater life and have excellent swimming abilities. Although previous motion analyses revealed their basic swimming characteristics, the details of the 3D wing kinematics, wing deformation and thrust generation mechanism of penguins are still largely unknown. In this study, we recorded the forward and horizontal swimming of gentoo penguins (Pygoscelis papua) at an aquarium with multiple underwater action cameras and then performed a 3D motion analysis. We also conducted a series of water tunnel experiments with a 3D printed rigid wing to obtain lift and drag coefficients in the gliding configuration. Using these coefficients, the thrust force during flapping was calculated in a quasi-steady manner, where the following two wing models were considered: (1) an 'original' wing model reconstructed from 3D motion analysis including bending deformation and (2) a 'flat' wing model obtained by flattening the original wing model. The resultant body trajectory showed that the penguin accelerated forward during both upstroke and downstroke. The motion analysis of the two wing models revealed that considerable bending occurred in the original wing, which reduced its angle of attack during the upstroke in particular. Consequently, the calculated stroke-averaged thrust was larger for the original wing than for the flat wing during the upstroke. In addition, the propulsive efficiency for the original wing was estimated to be 1.8 times higher than that for the flat wing. Our results unveil a detailed mechanism of lift-based propulsion in penguins and underscore the importance of wing bending.

Direct Measurements of the Wing Kinematics of a Bat in Straight Flight.

Bat is the only mammal in the nature that can fly. Compared with birds and insects, bats are quite special in that their wings are formed by an elastic membrane, which renders that the airfoil deforms greatly during downstroke and upstroke. Due to the compliant skin of a bat, the movements of its wings are three-dimensionally complex during diverse flight behaviors. To understand the maneuverability and flight performance, three-dimensional reconstruction of the flight kinematics is essential. This study focuses on the reconstruction of the wing kinematics of the bat and identifies the primary relationship of parameters of aerodynamics in straight flight. With markers pasted on the wings and body of a bat, the motions of these points are recorded by a computerized optical motion capture system. The kinematic analysis shows that the motion of wings is very intricate. The digits of the wing display the sign of coupled motion. A novel approach was developed to measure the angle of attack and flapping angle of the wing. The angle of attack of leading edge differs with the overall angle of attack of the wing. The kinematics of the bat's wing is helpful to interpret the secret of the bat's flight.

Wearable Vibration Sensor for Measuring the Wing Flapping of Insects.

In this study, we fabricated a novel wearable vibration sensor for insects and measured their wing flapping. An analysis of insect wing deformation in relation to changes in the environment plays an important role in understanding the underlying mechanism enabling insects to dynamically interact with their surrounding environment. It is common to use a high-speed camera to measure the wing flapping; however, it is difficult to analyze the feedback mechanism caused by the environmental changes caused by the flapping because this method applies an indirect measurement. Therefore, we propose the fabrication of a novel film sensor that is capable of measuring the changes in the wingbeat frequency of an insect. This novel sensor is composed of flat silver particles admixed with a silicone polymer, which changes the value of the resistor when a bending deformation occurs. As a result of attaching this sensor to the wings of a moth and a dragonfly and measuring the flapping of the wings, we were able to measure the frequency of the flapping with high accuracy. In addition, as a result of simultaneously measuring the relationship between the behavior of a moth during its search for an odor source and its wing flapping, it became clear that the frequency of the flapping changed depending on the frequency of the odor reception. From this result, a wearable film sensor for an insect that can measure the displacement of the body during a particular behavior was fabricated.

Data-driven CFD Scaling of Bioinspired Mars Flight Vehicles for Hover.

One way to improve our model of Mars is through aerial sampling and surveillance, which could provide information to augment the observations made by ground-based exploration and satellite imagery. Flight in the challenging ultra-low-density Martian environment can be achieved with properly scaled bioinspired flapping wing vehicle configurations that utilize the same high lift producing mechanisms that are employed by insects on Earth. Through dynamic scaling of wings and kinematics, we investigate the ability to generate solutions for a broad range of flapping wing flight vehicles masses ranging from insects O(10-3) kg to the Mars helicopter Ingenuity O(100) kg. A scaling method based on a neural-network trained on 3D Navier-Stokes solutions is proposed to determine approximate wing size and kinematic values that generate bioinspired hover solutions. We demonstrate that a family of solutions exists for designs that range from 1 to 1000 grams, which are verified and examined using a 3D Navier-Stokes solver. Our results reveal that unsteady lift enhancement mechanisms, such as delayed stall and rotational lift, are present in the bioinspired solutions for the scaled vehicles hovering in Martian conditions. These hovering vehicles exhibit payloads of up to 1 kg and flight times on the order of 100 minutes when considering the respective limiting cases of the vehicle mass being comprised entirely of payload or entirely of a battery and neglecting any transmission inefficiencies. This method can help to develop a range of Martian flying vehicle designs with mission viable payloads, range, and endurance.

Elastic wing deformations mitigate flapping asymmetry during manoeuvres in rose chafers (Protaetia cuprea).

To manoeuvre in air, flying animals produce asymmetric flapping between contralateral wings. Unlike the adjustable vertebrate wings, insect wings lack intrinsic musculature, preventing active control over wing shape during flight. However, the wings elastically deform as a result of aerodynamic and inertial forces generated by the flapping motions. How these elastic deformations vary with flapping kinematics and flight performance in free-flying insects is poorly understood. Using high-speed videography, we measured how contralateral wings elastically deform during free-flight manoeuvring in rose chafer beetles (Protaetia cuprea). We found that asymmetric flapping during aerial turns was associated with contralateral differences in chord-wise wing deformations. The highest instantaneous difference in deformation occurred during stroke reversals, resulting from differences in wing rotation timing. Elastic deformation asymmetry was also evident during mid-strokes, where wing compliance increased the angle of attack of both wings, but reduced the asymmetry in the angle of attack between contralateral wings. A biomechanical model revealed that wing compliance can increase the torques generated by each wing, providing higher potential for manoeuvrability, while concomitantly contributing to flight stability by attenuating steering asymmetry. Such stability may be adaptive for insects such as flower chafers that need to perform delicate low-speed landing manoeuvres among vegetation.

Wind-associated detours promote seasonal migratory connectivity in a flapping flying long-distance avian migrant.

It is essential to gain knowledge about the causes and extent of migratory connectivity between stationary periods of migrants to further the understanding of processes affecting populations, and to allow efficient implementation of conservation efforts throughout the annual cycle. Avian migrants likely use optimal routes with respect to mode of locomotion, orientation and migration strategy, influenced by external factors such as wind and topography. In self-powered flapping flying birds, any increases in fuel loads are associated with added flight costs. Energy-minimizing migrants are therefore predicted to trade-off extended detours against reduced travel across ecological barriers with no or limited foraging opportunities. Here, we quantify the extent of detours taken by different populations of European nightjars Caprimulgus europaeus, to test our predictions that they used routes beneficial according to energetic principles and evaluate the effect of route shape on seasonal migratory connectivity. We combined data on birds tracked from breeding sites along a longitudinal gradient from England to Sweden. We analysed the migratory connectivity between breeding and main non-breeding sites, and en route stopover sites just south of the Sahara desert. We quantified each track's route extension relative to the direct route between breeding and wintering sites, respectively, and contrasted it to the potential detour derived from the barrier reduction along the track while accounting for potential wind effects. Nightjars extended their tracks from the direct route between breeding and main non-breeding sites as they crossed the Mediterranean Sea-Sahara desert, the major ecological barrier in the Palaearctic-African migration system. These clockwise detours were small for birds from eastern sites but increased from east to west breeding longitude. Routes of the tracked birds were associated with partial reduction in the barrier crossing resulting in a trade-off between route extension and barrier reduction, as expected in an energy-minimizing migrant. This study demonstrates how the costs of barrier crossings in prevailing winds can disrupt migratory routes towards slightly different goals, and thereby promote migratory connectivity. This is an important link between individual migration strategies in association with an ecological barrier, and both spatially and demographic population patterns.

Speed control and force-vectoring of bluebottle flies in a magnetically levitated flight mill.

Flies fly at a broad range of speeds and produce sophisticated aerial maneuvers with precisely controlled wing movements. Remarkably, only subtle changes in wing motion are used by flies to produce aerial maneuvers, resulting in little directional tilt of the aerodynamic force vector relative to the body. Therefore, it is often considered that flies fly according to a helicopter model and control speed mainly via force vectoring by body pitch change. Here, we examined the speed control of bluebottle flies using a magnetically levitated (MAGLEV) flight mill, as they fly at different body pitch angles and with different augmented aerodynamic damping. We identified wing kinematic contributors to the changes of estimated aerodynamic force through testing and comparing two force-vectoring models - a constant force-vectoring model and a variable force-vectoring model - while using Akaike's information criterion for selection of the best-approximating model. The results show that the best-approximating variable force-vectoring model, which includes the effects of wing kinematic changes, yields a considerably more accurate prediction of flight speed, particularly in higher velocity range, as compared with those of the constant force-vectoring model. Examination of the variable force-vectoring model reveals that, in the flight mill tethered flight, flies use a collection of wing kinematic variables to control primarily the force magnitude, while the force direction is also modulated, albeit to a smaller extent compared with those due to the changes in body pitch. The roles of these wing kinematic variables are analogous to those of throttle, and collective and cyclic pitch of helicopters.

Wing morphology, flight type and migration distance predict accumulated fuel load in birds.

Birds often accumulate large fat and protein reserves to fuel long-distance flights. While it is well known that species that fly the longest accumulate the largest amounts of fuel, considerable cross-species variation in fuel load is seen after controlling for overall migration distance. It remains unclear whether this variation can be explained by aerodynamic attributes of different species, despite obvious ecological and conservation implications. Here, we collected data on wing morphology, flight type, migration distance and fuel load from 213 European bird species and explored three questions: (1) does maximum fuel load relate to migration distance across species?; (2) does wing morphology, as described by wing aspect ratio and wing loading, influence maximum fuel load?; and (3) does flight type influence maximum fuel load? Our results indicate that maximum fuel load increases with migration across species, but residual variance is high. The latter variance is explained by aspect ratio and flight type, while wing loading and body mass explain little variance. Birds with slender wings accumulate less fuel than species with low wing aspect ratio when covering a similar migration distance. Continuously flapping species accumulate the largest amounts of fuel, followed by flapping and soaring species and flapping and gliding species, while the smallest fuel loads were observed in birds with passerine-type flight. These results highlight complex eco-evolutionary adaptations to migratory behaviour, pointing toward the importance of energy minimisation.

Enhancing Output Power of a Cantilever-Based Flapping Airflow Energy Harvester Using External Mechanical Interventions.

This paper presents a flapping airflow energy harvester based on oscillations of a horizontal cantilever beam facing the direction of airflow. A wing is attached to the free end of a cantilever beam and a bluff body is placed in front of the wing from where vortex falls off, producing vortices under the wing and driving it to oscillate. An electromagnetic transducer is integrated to convert the flow induced vibration into electrical energy. This flapping energy harvester, however, may stop oscillating or vibrate in the second mode under high electrical damping, and thus may be unable to achieve its optimum performance. Simple yet effective mechanical interventions can be applied to the harvester to enhance its power output, i.e., to increase flow velocity and to apply external magnetic interaction. The effect of airflow velocities on output power was investigated experimentally and the results show that the energy harvester scavenges more power in airflow at higher Reynolds numbers (higher flow velocity at R e < 24,000). The external magnetic excitation is achieved though placing one magnet to the wing and another one above the wing to induce a repelling force, aiding the beam to oscillate in high electrical damping. Experimental results show that the power output can be enhanced by 30% when the magnet interaction is properly integrated.

Research on Key Techniques of Insect Flapping Onset Control Based on Electrical Stimulation.

In this paper, an insect flapping onset control method based on electrical stimulation is proposed. The beetle (Allomyrina dithotomus, Coleoptera) is employed for the research carrier, and it's left and right longitudinal muscles are electrically stimulated to control the flapping onset behavior. The control principle of insect flapping onset utilizing electrical stimulation is analyzed firstly followed by the movement function of the dorsal longitudinal muscle. Subsequently, a micro-control system, which is composed of a PC controller, coordinator and electronic backpack, is designed to realize the wireless control of beetle movements. Finally, the verification experiment is implemented to verify the effectiveness of dorsal longitudinal muscle stimulation with respect to the beetle flapping onset, whereas the comparative experiment emphasizes on determining optimal simulating parameters. The experimental results demonstrate that when the period, duty ratio, number of and amplitude of pulses stimulation signal are assigned to 5 ms, 20%, 90 and 3.3 V respectively, the beetle flapping onset can be controlled with an average response time of 1.69 s. Simultaneously, the optimization of duty ratio from 20% to 40%, and the number of pulses from 90 to 100, is proved to the best parameter configuration.

Aerodynamic characteristics along the wing span of a dragonfly Pantala flavescens.

We investigated the characteristics of interwing aerodynamic interactions across the span of the high aspect ratio, flexible wings of dragonflies under tethered and free-flying conditions. This revealed that the effects of the interactions on the hindwings vary across four spanwise regions. (i) Close to the wing root, a trailing-edge vortex (TEV) is formed by each stroke, while the formation of a leading-edge vortex (LEV) is limited by the short translational distance of the hindwing and suppressed by the forewing-induced flow. (ii) In the region away from the wing root but not quite up to midspan, the formation of the hindwing LEV is influenced by that of the forewing LEV. This vortex synergy can increase the circulation of the hindwing LEV in the corresponding cross-section by 22% versus that of the hindwing in isolation. (iii) In the region about half-way between the wing root and wing tip there is a transition dominated by downwash from the forewing resulting in flow attached to the hindwing. (iv) A LEV is developed in the remaining, outer region of the wing at the end of a stroke when the hindwing captures the vortex shed by the forewing. The interaction effects depend not only on the wing phasing but also on the flapping offset and flight direction. The aerodynamics of the hindwings vary substantially from the wing root to the wing tip. For a given phasing, this spanwise variation in the aerodynamics can be exploited in the design of artificial wings to achieve greater agility and higher efficiency.

Wing inertia as a cause of aerodynamically uneconomical flight with high angles of attack in hovering insects.

Flying insects can maintain maneuverability in the air by flapping their wings, and, to save energy, the wings should operate following optimal kinematics. However, unlike conventional rotary wings, insects operate their wings at aerodynamically uneconomical and high angles of attack (AoA). Although insects have continuously received attention from biologists and aerodynamicists, the high AoA operation in insect flight has not been clearly explained. Here, we used a theoretical blade-element model to examine the impact of wing inertia on the power requirement and flapping AoA, based on 3D free-hovering flight wing kinematics of a horned beetle, Allomyrinadichotoma The relative simplicity of the model allowed us to search for the best AoA distributed along the wingspan, which generate the highest vertical force per unit power. We show that, although elastic elements may be involved in flight muscles to store and save energy, the insect still has to use substantial power to accelerate its wings, because inertial energy stores should be used to overcome aerodynamic drag before being stored elastically. At the same flapping speed, a wing operating at a higher AoA requires lower inertial torque, and therefore lower inertial power output, at stroke reversals than a wing operating at an aerodynamically optimal low AoA. An interactive aerodynamic-inertial effect thereby enables the wing to flap at sufficiently high AoA, which causes an aerodynamically uneconomical flight in an effort to minimize the net flight energy.

New approaches for assessing squid fin motions: coupling proper orthogonal decomposition with volumetric particle tracking velocimetry.

Squid, which swim using a coupled fin/jet system powered by muscular hydrostats, pose unique challenges for the study of locomotion. The high flexibility of the fins and complex flow fields generated by distinct propulsion systems require innovative techniques for locomotive assessment. For this study, we used proper orthogonal decomposition (POD) to decouple components of the fin motions and defocusing digital particle tracking velocimetry (DDPTV) to quantify the resultant 3D flow fields. Kinematic footage and DDPTV data were collected from brief squid, Lolliguncula brevis [3.1-6.5 cm dorsal mantle length (DML)], swimming freely in a water tunnel at speeds of 0.39-7.20 DML s-1 Both flap and wave components were present in all fin motions, but the relative importance of the wave components was higher for arms-first swimming than for tail-first swimming and for slower versus higher speed swimming. When prominent wave components were present, more complex interconnected vortex ring wakes were observed, while fin movements dominated by flapping resulted in more spatially separated vortex ring patterns. Although the jet often produced the majority of the thrust for steady rectilinear swimming, our results demonstrated that the fins can contribute more thrust than the jet at times, consistently produce comparable levels of lift to the jet during arms-first swimming, and can boost overall propulsive efficiency. By producing significant drag signatures, the fins can also aid in stabilization and maneuvering. Clearly, fins play multiple roles in squid locomotion, and when coupled with the jet, allow squid to perform a range of swimming behaviors integral to their ecological success.

The four-flipper swimming method of plesiosaurs enabled efficient and effective locomotion.

The extinct ocean-going plesiosaurs were unique within vertebrates because they used two flipper pairs identical in morphology for propulsion. Although fossils of these Mesozoic marine reptiles have been known for more than two centuries, the function and dynamics of their tandem-flipper propulsion system has always been unclear and controversial. We address this question quantitatively for the first time in this study, reporting a series of precisely controlled water tank experiments that use reconstructed plesiosaur flippers scaled from well-preserved fossils. Our aim was to determine which limb movements would have resulted in the most efficient and effective propulsion. We show that plesiosaur hind flippers generated up to 60% more thrust and 40% higher efficiency when operating in harmony with their forward counterparts, when compared with operating alone, and the spacing and relative motion between the flippers was critical in governing these increases. The results of our analyses show that this phenomenon was probably present across the whole range of plesiosaur flipper motion and resolves the centuries-old debate about the propulsion style of these marine reptiles, as well as indicating why they retained two pairs of flippers for more than 100 million years.