Measuring the 3D wake of swimming snakes (Natrix tessellata) using volumetric particle image velocimetry.

We describe a method for measuring the 3D vortical structures produced by an anguilliform swimmer using volumetric velocimetry. The wake of freely swimming dice snakes (Natrix tessellata) was quantified, revealing the creation of multiple vortices along the body of the snake due to its undulation. The 3D structure of the vortices generally consisted of paired vortex tubes, some of which were linked together to form a hairpin structure. The observations match predictions from computational fluid dynamic studies of other anguilliform swimmers. Quantitative measurements allowed us to study vortex circulation and size, and global kinetic energy of the flow, which varied with swimming speed, vortex topology and individual characteristics. Our findings provide a baseline for comparing wake structures of snakes with different morphologies and ecologies and investigating the energetic efficiency of anguilliform swimming.

Illuminance-tuned collective motion in fish.

We experimentally investigate the role of illumination on the collective dynamics of a large school (ca. 50 individuals) of Hemigrammus rhodostomus. The structure of the group, defined using two order parameters, is quantified while progressively altering the visual range of the fish through controlled cycles of ambient light intensity. We show that, at low light levels, the individuals within the group are unable to form a cohesive group, while at higher illuminance the degree of alignment of the school correlates with the light intensity. When increasing the illuminance, the school structure is successively characterized by a polarized state followed by a highly regular and stable rotational configuration (milling). Our study shows that vision is necessary to achieve cohesive collective motion for free swimming fish schools, while the short-range lateral line sensing is insufficient in this situation. The present experiment therefore provides new insights into the interaction mechanisms that govern the emergence and intensity of collective motion in biological systems.

Special issue: bioinspired fluid-structure interaction.

Fluid-structure interaction (FSI) studies the interaction between fluid and solid objects. It helps understand how fluid motion affects solid objects and vice versa. FSI research is important in engineering applications such as aerodynamics, hydrodynamics, and structural analysis. It has been used to design efficient systems such as ships, aircraft, and buildings. FSI in biological systems has gained interest in recent years for understanding how organisms interact with their fluidic environment. Our special issue features papers on various biological and bio-inspired FSI problems. Papers in this special issue cover topics ranging from flow physics to optimization and diagonistics. These papers offer new insights into natural systems and inspire the development of new technologies based on natural principles.

Divergence of climbing escape flight performance in Morpho butterflies living in different microhabitats.

Habitat specialization can influence the evolution of animal movement in promoting divergent locomotor abilities adapted to contrasting environmental conditions, differences in vegetation clutter or predatory communities. While the effect of habitat on the evolution of locomotion and particularly escape performance has been well investigated in terrestrial animals, it remains understudied in flying animals. Here, we investigated whether specialization of Morpho butterfly species into different vertical strata of the Amazonian forest affects the performance of upward escape flight manoeuvres. Using stereoscopic high-speed videography, we compared the climbing flight kinematics of seven Morpho species living either in the forest canopy or in the understory. We show that butterflies from canopy species display strikingly higher climbing speed and steeper ascent angle compared with understory species. Although climbing speed increased with wing speed and angle of attack, the higher climb angle observed in canopy species was best explained by their higher body pitch angle, resulting in more upward-directed aerodynamic thrust forces. Climb angle also scales positively with weight-normalized wing area, and this weight-normalized wing area was higher in canopy species. This shows that a combined divergence in flight behaviour and morphology contributes to the evolution of increased climbing flight abilities in canopy species.

Wake and aeroelasticity of a flexible pitching foil.

A flexible foil undergoing pitching oscillations is studied experimentally in a wind tunnel with different imposed free stream velocities. The chord-based Reynolds number is in the range 1600-4000, such that the dynamics of the system is governed by inertial forces and the wake behind the foil exhibits the reverse Bénard-von Kármán vortex street characteristic of flapping-based propulsion. Particle image velocimetry (PIV) measurements are performed to examine the flow around the foil, whilst the deformation of the foil is also tracked. The first natural frequency of vibration of the foil is within the range of flapping frequencies explored, determining a strongly-coupled dynamics between the elastic foil deformation and the vortex shedding. Cluster-based reduced order modelling is applied on the PIV data in order to identify the coherent flow structures. Analysing the foil kinematics and using a control-volume calculation of the average drag forces from the corresponding velocity fields, we determine the optimal flapping configurations for thrust generation. We show that propulsive force peaks occur at dimensionless frequencies shifted with respect to the elastic resonances that are marked by maximum trailing edge oscillation amplitudes. The thrust peaks are better explained by a wake resonance, which we examine using the tools of classic hydrodynamic stability on the mean propulsive jet profiles.

Hydrodynamical Fingerprint of a Neighbour in a Fish Lateral Line.

For fish, swimming in group may be favorable to individuals. Several works reported that in a fish school, individuals sense and adjust their relative position to prevent collisions and maintain the group formation. Also, from a hydrodynamic perspective, relative-position and kinematic synchronisation between adjacent fish may considerably influence their swimming performance. Fish may sense the relative-position and tail-beat phase difference with their neighbors using both vision and the lateral-line system, however, when swimming in dark or turbid environments, visual information may become unavailable. To understand how lateral-line sensing can enable fish to judge the relative-position and phase-difference with their neighbors, in this study, based on a verified three-dimensional computational fluid dynamics approach, we simulated two fish swimming adjacently with various configurations. The lateral-line signal was obtained by sampling the surface hydrodynamic stress. The sensed signal was processed by Fast Fourier Transform (FFT), which is robust to turbulence and environmental flow. By examining the lateral-line pressure and shear-stress signals in the frequency domain, various states of the neighboring fish were parametrically identified. Our results reveal that the FFT-processed lateral-line signals in one fish may potentially reflect the relative-position, phase-differences, and the tail-beat frequency of its neighbor. Our results shed light on the fluid dynamical aspects of the lateral-line sensing mechanism used by fish. Furthermore, the presented approach based on FFT is especially suitable for applications in bioinspired swimming robotics. We provide suggestions for the design of artificial systems consisting of multiple stress sensors for robotic fish to improve their performance in collective operation.

Burst-and-coast swimmers optimize gait by adapting unique intrinsic cycle.

This paper addresses the physical mechanism of intermittent swimming by considering the burst-and-coast regime of fish swimming at different speeds. The burst-and-coast regime consists of a cycle with two successive phases, i.e., a phase of active undulation powered by the fish muscles followed by a passive gliding phase. Observations of real fish whose swimming gait is forced in a water flume from low to high speed regimes are performed, using a full description of the fish kinematics and mechanics. We first show that fish modulate a unique intrinsic cycle to sustain the demanded speed by modifying the bursting to coasting ratio while maintaining the duration of the cycle nearly constant. Secondly, we show using numerical simulations that the chosen kinematics by correspond to optimized gaits for swimming speeds larger than 1 body length per second.

Exploring the functional meaning of head shape disparity in aquatic snakes.

Phenotypic diversity, or disparity, can be explained by simple genetic drift or, if functional constraints are strong, by selection for ecologically relevant phenotypes. We here studied phenotypic disparity in head shape in aquatic snakes. We investigated whether conflicting selective pressures related to different functions have driven shape diversity and explore whether similar phenotypes may give rise to the same functional output (i.e., many-to-one mapping of form to function). We focused on the head shape of aquatically foraging snakes as they fulfill several fitness-relevant functions and show a large amount of morphological variability. We used 3D surface scanning and 3D geometric morphometrics to compare the head shape of 62 species in a phylogenetic context. We first tested whether diet specialization and size are drivers of head shape diversification. Next, we tested for many-to-one mapping by comparing the hydrodynamic efficiency of head shape characteristic of the main axes of variation in the dataset. We 3D printed these shapes and measured the forces at play during a frontal strike. Our results show that diet and size explain only a small amount of shape variation. Shapes did not fully functionally converge as more specialized aquatic species evolved a more efficient head shape than others. The shape disparity observed could thus reflect a process of niche specialization.

On the Fluid Dynamical Effects of Synchronization in Side-by-Side Swimmers.

In-phase and anti-phase synchronization of neighboring swimmers is examined experimentally using two self-propelled independent flexible foils swimming side-by-side in a water tank. The foils are actuated by pitching oscillations at one extremity-the head of the swimmers-and the flow engendered by their undulations is analyzed using two-dimensional particle image velocimetry in their frontal symmetry plane. Following recent observations on the behavior of real fish, we focus on the comparison between in-phase and anti-phase actuation by fixing all other geometric and kinematic parameters. We show that swimming with a neighbor is beneficial for both synchronizations tested, as compared to swimming alone, with an advantage for the anti-phase synchronization. We show that the advantage of anti-phase synchronization in terms of swimming performance for the two-foil "school" results from the emergence of a periodic coherent jet between the two swimmers.

On the energetics and stability of a minimal fish school.

The physical basis for fish schooling is examined using three-dimensional numerical simulations of a pair of swimming fish, with kinematics and geometry obtained from experimental data. Energy expenditure and efficiency are evaluated using a cost of transport function, while the effect of schooling on the stability of each swimmer is examined by probing the lateral force and the lateral and longitudinal force fluctuations. We construct full maps of the aforementioned quantities as functions of the spatial pattern of the swimming fish pair and show that both energy expenditure and stability can be invoked as possible reasons for the swimming patterns and tail-beat synchronization observed in real fish. Our results suggest that high cost of transport zones should be avoided by the fish. Wake capture may be energetically unfavorable in the absence of kinematic adjustment. We hereby hypothesize that fish may restrain from wake capturing and, instead, adopt side-to-side configuration as a conservative strategy, when the conditions of wake energy harvesting are not satisfied. To maintain a stable school configuration, compromise between propulsive efficiency and stability, as well as between school members, ought to be considered.

Hydrodynamics of frontal striking in aquatic snakes: drag, added mass, and the possible consequences for prey capture success.

Transient locomotion under water is highly constrained by drag and added mass, yet some aquatic snakes catch their prey using a fast forward acceleration, with the mouth opened. These aquatic snakes show a convergence of their head shape in comparison with closely related species that do not forage under water. As both drag and added mass are related to some extent to the shape of the moving object, we explored how shape impacts the hydrodynamic forces applied to the head of a snake during a prey capture event. We compared two 3D-printed heads representing typical shapes of aquatically-foraging and non-aquatically-foraging snakes, and frontal strike kinematics based on in vivo observations. By using direct force measurements, we calculated the drag and added mass coefficient of the two models. Our results show that both drag and added mass are reduced in aquatic snakes. The drag coefficient of the aquatic model is 0.24, which is almost two times smaller than the non-aquatic model. The added mass coefficient of the aquatic model is 0.15 versus 0.24 for the non-aquatic model, showing that the convergence of head shape in aquatically foraging snakes is associated with a hydrodynamic advantage during frontal striking. The vorticity field measurements with particle image velocimetry show that a less intense recirculation bubble behind the jaw of the aquatic model, compared to the non-aquatic model, might be the basis of this advantage.

Insect and insect-inspired aerodynamics: unsteadiness, structural mechanics and flight control.

Flying insects impress by their versatility and have been a recurrent source of inspiration for engineering devices. A large body of literature has focused on various aspects of insect flight, with an essential part dedicated to the dynamics of flapping wings and their intrinsically unsteady aerodynamic mechanisms. Insect wings flex during flight and a better understanding of structural mechanics and aeroelasticity is emerging. Most recently, insights from solid and fluid mechanics have been integrated with physiological measurements from visual and mechanosensors in the context of flight control in steady airs and through turbulent conditions. We review the key recent advances concerning flight in unsteady environments and how the multi-body mechanics of the insect structure-wings and body-are at the core of the flight control question. The issues herein should be considered when applying bio-informed design principles to robotic flapping wings.

Simple phalanx pattern leads to energy saving in cohesive fish schooling.

The question of how individuals in a population organize when living in groups arises for systems as different as a swarm of microorganisms or a flock of seagulls. The different patterns for moving collectively involve a wide spectrum of reasons, such as evading predators or optimizing food prospection. Also, the schooling pattern has often been associated with an advantage in terms of energy consumption. In this study, we use a popular aquarium fish, the red nose tetra fish, Hemigrammus bleheri, which is known to swim in highly cohesive groups, to analyze the schooling dynamics. In our experiments, fish swim in a shallow-water tunnel with controlled velocity, and stereoscopic video recordings are used to track the 3D positions of each individual in a school, as well as their tail-beating kinematics. Challenging the widespread idea of fish favoring a diamond pattern to swim more efficiently [Weihs D (1973) Nature 241:290-291], we observe that when fish are forced to swim fast-well above their free-swimming typical velocity, and hence in a situation where efficient swimming would be favored-the most frequent configuration is the "phalanx" or "soldier" formation, with all individuals swimming side by side. We explain this observation by considering the advantages of tail-beating synchronization between neighbors, which we have also characterized. Most importantly, we show that schooling is advantageous as compared with swimming alone from an energy-efficiency perspective.

Does aquatic foraging impact head shape evolution in snakes?

Evolutionary trajectories are often biased by developmental and historical factors. However, environmental factors can also impose constraints on the evolutionary trajectories of organisms leading to convergence of morphology in similar ecological contexts. The physical properties of water impose strong constraints on aquatic feeding animals by generating pressure waves that can alert prey and potentially push them away from the mouth. These hydrodynamic constraints have resulted in the independent evolution of suction feeding in most groups of secondarily aquatic tetrapods. Despite the fact that snakes cannot use suction, they have invaded the aquatic milieu many times independently. Here, we test whether the aquatic environment has constrained head shape evolution in snakes and whether shape converges on that predicted by biomechanical models. To do so, we used three-dimensional geometric morphometrics and comparative, phylogenetically informed analyses on a large sample of aquatic snake species. Our results show that aquatic snakes partially conform to our predictions and have a narrower anterior part of the head and dorsally positioned eyes and nostrils. This morphology is observed, irrespective of the phylogenetic relationships among species, suggesting that the aquatic environment does indeed drive the evolution of head shape in snakes, thus biasing the evolutionary trajectory of this group of animals.

Large-amplitude undulatory swimming near a wall.

The propulsive dynamics of a flexible undulating foil in a self-propelled swimming configuration near a wall is studied experimentally. Measurements of the swimming speed and the propulsive force are presented, together with image acquisition of the kinematics of the foil and particle image velocimetry (PIV) in its wake. The presence of the wall enhances the cruising velocity in some cases up to 25% and the thrust by a 45% , for swept angles of 160 and 240°. The physical mechanisms underlying this effect are discussed by studying the vorticity dynamics in the wake of the foil. Proper orthogonal decomposition is applied to the PIV measurements in order to analyse the kinetic energy modal distribution in the flow and to relate it to the propulsion generated by the foil.

Passive elastic mechanism to mimic fish-muscle action in anguilliform swimming.

Swimmers in nature use body undulations to generate propulsive and manoeuvring forces. The anguilliform kinematics is driven by muscular actions all along the body, involving a complex temporal and spatial coordination of all the local actuations. Such swimming kinematics can be reproduced artificially, in a simpler way, by using the elasticity of the body passively. Here, we present experiments on self-propelled elastic swimmers at a free surface in the inertial regime. By addressing the fluid-structure interaction problem of anguilliform swimming, we show that our artificial swimmers are well described by coupling a beam theory with the potential flow model of Lighthill. In particular, we show that the propagative nature of the elastic wave producing the propulsive force is strongly dependent on the dissipation of energy along the body of the swimmer.

Force balance in the take-off of a pierid butterfly: relative importance and timing of leg impulsion and aerodynamic forces.

Up to now, the take-off stage has remained an elusive phase of insect flight that was relatively poorly explored compared with other maneuvers. An overall assessment of the different mechanisms involved in force production during take-off has never been explored. Focusing on the first downstroke, we have addressed this problem from a force balance perspective in butterflies taking off from the ground. In order to determine whether the sole aerodynamic wing force could explain the observed motion of the insect, we have firstly compared a simple analytical model of the wing force with the acceleration of the insect's center of mass estimated from video tracking of the wing and body motions. Secondly, wing kinematics were also used for numerical simulations of the aerodynamic flow field. Similar wing aerodynamic forces were obtained by the two methods. However, neither are sufficient, nor is the inclusion of the ground effect, to predict faithfully the body acceleration. We have to resort to the leg forces to obtain a model that best fits the data. We show that the median and hind legs display an active extension responsible for the initiation of the upward motion of the insect's body, occurring before the onset of the wing downstroke. We estimate that legs generate, at various times, an upward force that can be much larger than all other forces applied to the insect's body. The relative timing of leg and wing forces explains the large variability of trajectories observed during the maneuvers.

Spatiotemporal spectral analysis of a forced cylinder wake.

The wake of a circular cylinder performing rotary oscillations is studied using hydrodynamic tunnel experiments at Re=100. Two-dimensional particle image velocimetry on the midplane perpendicular to the axis of a cylinder is used to characterize the spatial development of the flow and its stability properties. The lock-in phenomenon that determines the boundaries between regions of the forcing parameter space where the wake is globally unstable or convectively unstable [see Thiria and Wesfreid, J. Fluids Struct. 25, 654 (2009) for a review] is scrutinized using the experimental data. A method based on the analysis of power density spectra of the flow allows us to give a detailed description of the forced wake, shedding light on the energy distribution in the different frequency components and in particular on a cascade-like mechanism evidenced for a high amplitude of the forcing oscillation. In addition, a calculation of the drag from the velocity field is performed, allowing us to relate the resulting force on the body to the wake properties.

Rather than resonance, flapping wing flyers may play on aerodynamics to improve performance.

Saving energy and enhancing performance are secular preoccupations shared by both nature and human beings. In animal locomotion, flapping flyers or swimmers rely on the flexibility of their wings or body to passively increase their efficiency using an appropriate cycle of storing and releasing elastic energy. Despite the convergence of many observations pointing out this feature, the underlying mechanisms explaining how the elastic nature of the wings is related to propulsive efficiency remain unclear. Here we use an experiment with a self-propelled simplified insect model allowing to show how wing compliance governs the performance of flapping flyers. Reducing the description of the flapping wing to a forced oscillator model, we pinpoint different nonlinear effects that can account for the observed behavior--in particular a set of cubic nonlinearities coming from the clamped-free beam equation used to model the wing and a quadratic damping term representing the fluid drag associated to the fast flapping motion. In contrast to what has been repeatedly suggested in the literature, we show that flapping flyers optimize their performance not by especially looking for resonance to achieve larger flapping amplitudes with less effort, but by tuning the temporal evolution of the wing shape (i.e., the phase dynamics in the oscillator model) to optimize the aerodynamics.

How wing compliance drives the efficiency of self-propelled flapping flyers.

Wing flexibility governs the flying performance of flapping-wing flyers. Here, we use a self-propelled flapping-wing model mounted on a "merry go round" to investigate the effect of wing compliance on the propulsive efficiency of the system. Our measurements show that the elastic nature of the wings can lead not only to a substantial reduction in the consumed power, but also to an increment of the propulsive force. A scaling analysis using a flexible plate model for the wings points out that, for flapping flyers in air, the time-dependent shape of the elastic bending wing is governed by the wing inertia. Based on this prediction, we define the ratio of the inertial forces deforming the wing to the elastic restoring force that limits the deformation as the elastoinertial number N(ei). Our measurements with the self-propelled model confirm that it is the appropriate structural parameter to describe flapping flyers with flexible wings.