Fish and robots swimming together: attraction towards the robot demands biomimetic locomotion

The integration of biomimetic robots in a fish school may enable a better understanding of collective behaviour, offering a new experimental method to test group feedback in response to behavioural modulations of its ‘engineered’ member. Here, we analyse a robotic fish and individual golden shiners (Notemigonus crysoleucas) swimming together in a water tunnel at different flow velocities. We determine the positional preference of fish with respect to the robot, and we study the flow structure using a digital particle image velocimetry system. We find that biomimetic locomotion is a determinant of fish preference as fish are more attracted towards the robot when its tail is beating rather than when it is statically immersed in the water as a ‘dummy’. At specific conditions, the fish hold station behind the robot, which may be due to the hydrodynamic advantage obtained by swimming in the robot''s wake. This work makes a compelling case for the need of biomimetic locomotion in promoting robot–animal interactions and it strengthens the hypothesis that biomimetic robots can be used to study and modulate collective animal behaviour.     

Oceanographic and behavioural assumptions in models of the fate of coral and coral reef fish larvae

A predictive model of the fate of coral reef fish larvae in a reef system is proposed that combines the oceanographic processes of advection and turbulent diffusion with the biological process of horizontal swimming controlled by olfactory and auditory cues within the timescales of larval development. In the model, auditory cues resulted in swimming towards the reefs when within hearing distance of the reef, whereas olfactory cues resulted in the larvae swimming towards the natal reef in open waters by swimming against the concentration gradients in the smell plume emanating from the natal reef. The model suggested that the self-seeding rate may be quite large, at least 20% for the larvae of rapidly developing reef fish species, which contrasted with a self-seeding rate less than 2% for non-swimming coral larvae. The predicted self-recruitment rate of reefs was sensitive to a number of parameters, such as the time at which the fish larvae reach post-flexion, the pelagic larval duration of the larvae, the horizontal turbulent diffusion coefficient in reefal waters and the horizontal swimming behaviour of the fish larvae in response to auditory and olfactory cues, for which better field data are needed. Thus, the model suggested that high self-seeding rates for reef fish are possible, even in areas where the ‘sticky water’ effect is minimal and in the absence of long-term trapping in oceanic fronts and/or large-scale oceanic eddies or filaments that are often argued to facilitate the return of the larvae after long periods of drifting at sea.     

Hydrodynamic constraints on prey-capture performance in forward-striking snakes

Some specialized aquatic snakes such as Natrix tessellata strike at fish by rapidly accelerating their head towards the prey with their mouth opened widely. This strategy is believed to be suboptimal as relatively high drag forces act on the open jaws and, therefore, probably limit strike speed. Moreover, the bow wave in front of the snake''s jaws could push prey away from the mouth, thus potentially explaining the relatively low capture success observed in these animals (<20%). Here, we used laser-scan based computational fluid dynamics to test these potential constraints on prey-capture performance for N. tessellata. Our simulations showed that drag force indeed increases drastically for striking at a high gape angle. However, we estimated the overall cost in slowing down strike speed to be less pronounced due to the instationary dynamics of the system. In contrast to the expectations, forward displacement of prey was relatively limited (<13% of head length), and forceful collisions between prey and the leading edge of the jaw regularly occurred. However, our models showed that precise aiming by the snake was needed to reduce the chance of deviating the prey to a path bypassing the mouth. Our study also indicated several hydrodynamic advantages for snakes to strike at relatively large prey.     

Analytical insights into optimality and resonance in fish swimming

This paper provides analytical insights into the hypothesis that fish exploit resonance to reduce the mechanical cost of swimming. A simple body–fluid fish model, representing carangiform locomotion, is developed. Steady swimming at various speeds is analysed using optimal gait theory by minimizing bending moment over tail movements and stiffness, and the results are shown to match with data from observed swimming. Our analysis indicates the following: thrust–drag balance leads to the Strouhal number being predetermined based on the drag coefficient and the ratio of wetted body area to cross-sectional area of accelerated fluid. Muscle tension is reduced when undulation frequency matches resonance frequency, which maximizes the ratio of tail-tip velocity to bending moment. Finally, hydrodynamic resonance determines tail-beat frequency, whereas muscle stiffness is actively adjusted, so that overall body–fluid resonance is exploited.     

Numerical model of self-propulsion in a fluid.

We provide initial evidence that a structure formed from an articulated series of linked elements, where each element has a given stiffness, damping and driving term with respect to its neighbours, may 'swim' through a fluid under certain conditions. We derive a Lagrangian for this system and, in particular, we note that we allow the leading edge to move along the x-axis. We assume that no lateral displacement of the leading edge of the structure is possible, although head 'yaw' is allowed. The fluid is simulated using a computational fluid dynamics technique, and we are able to determine and solve Euler-Lagrange equations for the structure. These two calculations are solved simultaneously by using a weakly coupled solver. We illustrate our method by showing that we are able to induce both forward and backward swimming. A discussion of the relevance of these simulations to a slowly swimming body, such as a mechanical device or a fish, is given.     

Unsteady motion: escape jumps in planktonic copepods,their kinematics and energetics

We describe the kinematics of escape jumps in three species of 0.3–3.0 mm-sized planktonic copepods. We find similar kinematics between species with periodically alternating power strokes and passive coasting and a resulting highly fluctuating escape velocity. By direct numerical simulations, we estimate the force and power output needed to accelerate and overcome drag. Both are very high compared with those of other organisms, as are the escape velocities in comparison to startle velocities of other aquatic animals. Thus, the maximum weight-specific force, which for muscle motors of other animals has been found to be near constant at 57 N (kg muscle)⁻¹, is more than an order of magnitude higher for the escaping copepods. We argue that this is feasible because most copepods have different systems for steady propulsion (feeding appendages) and intensive escapes (swimming legs), with the muscular arrangement of the latter probably adapted for high force production during short-lasting bursts. The resulting escape velocities scale with body length to power 0.65, different from the size-scaling of both similar sized and larger animals moving at constant velocity, but similar to that found for startle velocities in other aquatic organisms. The relative duration of the pauses between power strokes was observed to increase with organism size. We demonstrate that this is an inherent property of swimming by alternating power strokes and pauses. We finally show that the Strouhal number is in the range of peak propulsion efficiency, again suggesting that copepods are optimally designed for rapid escape jumps.     

Shear-induced orientational dynamics and spatial heterogeneity in suspensions of motile phytoplankton

Fluid flow, ubiquitous in natural and man-made environments, has the potential to profoundly impact the transport of microorganisms, including phytoplankton in aquatic habitats and bioreactors. Yet, the effect of ambient flow on the swimming behaviour of phytoplankton has remained poorly understood, largely owing to the difficulty of observing cell–flow interactions at the microscale. Here, we present microfluidic experiments where we tracked individual cells for four species of motile phytoplankton exposed to a spatially non-uniform fluid shear rate, characteristic of many flows in natural and artificial environments. We observed that medium-to-high mean shear rates (1–25 s⁻¹) produce heterogeneous cell concentrations in the form of regions of accumulation and regions of depletion. The location of these regions relative to the flow depends on the cells'' propulsion mechanism, body shape and flagellar arrangement, as captured by an effective aspect ratio. Species having a large effective aspect ratio accumulated in the high-shear regions, owing to shear-induced alignment of the swimming orientation with the fluid streamlines. Species having an effective aspect ratio close to unity exhibited little preferential accumulation at low-to-moderate flow rates, but strongly accumulated in the low-shear regions under high flow conditions, potentially owing to an active, behavioural response of cells to shear. These observations demonstrate that ambient fluid flow can strongly affect the motility and spatial distribution of phytoplankton and highlight the rich dynamics emerging from the interaction between motility, morphology and flow.     

Distribution of properties from sampling inhomogeneous materials by line transects

A theory related to the statistics of the structure and properties of inhomogeneous materials has been proposed, dealing with the distribution of properties from sampling of inhomogeneous structures by line transects. It is demonstrated that the empirical cumulative distribution functions of the intercepts from the microstructural constituents and their variances are important fingerprints of the inhomogeneous structures.Simulated sampling of the images of the microstructure by line transects has been proposed as a powerful method for characterization the distribution of properties of inhomogeneous structures. One of the advantages of the method based on the expected values of the order statistics of the intercepts is that the probability bounds of the property are determined without prior knowledge of the functional relationship between the property and the intercepts from the structural constituents.The concept intercept variance has an important application in determining the minimum transect length that stabilizes the variation of the intercept at a low value. Additionally, the concept ‘intercept variance’ can be used for topological optimization of the microstructure regarding the risk of intercepting a large amount of the weaker constituent. Equations regarding the variance of the intercepted fraction characterizing transect lengths with a specified distribution have also been derived.     

Homeostatic swimming of zooplankton upon crowding: the case of the copepod Centropages typicus

Crowding has a major impact on the dynamics of many material and biological systems, inducing effects as diverse as glassy dynamics and swarming. While this issue has been deeply investigated for a variety of living organisms, more research remains to be done on the effect of crowding on the behaviour of copepods, the most abundant metazoans on Earth. To this aim, we experimentally investigate the swimming behaviour, used as a dynamic proxy of animal adaptations, of males and females of the calanoid copepod Centropages typicus at different densities of individuals (10, 50 and 100 ind. l⁻¹) by performing three-dimensional single-organism tracking. We find that the C. typicus motion is surprisingly unaffected by crowding over the investigated density range. Indeed, the mean square displacements as a function of time always show a crossover from ballistic to Fickian regime, with poor variations of the diffusion constant on increasing the density. Close to the crossover, the displacement distributions display exponential tails with a nearly density-independent decay length. The trajectory fractal dimension, D_3D ≅ 1.5, and the recently proposed ‘ecological temperature’ also remain stable on increasing the individual density. This suggests that, at least over the range of animal densities used, crowding does not impact on the characteristics of C. typicus swimming motion, and that a homeostatic mechanism preserves the stability of its swimming performance.     

Vortex wake,downwash distribution,aerodynamic performance and wingbeat kinematics in slow-flying pied flycatchers

Many small passerines regularly fly slowly when catching prey, flying in cluttered environments or landing on a perch or nest. While flying slowly, passerines generate most of the flight forces during the downstroke, and have a ‘feathered upstroke’ during which they make their wing inactive by retracting it close to the body and by spreading the primary wing feathers. How this flight mode relates aerodynamically to the cruising flight and so-called ‘normal hovering’ as used in hummingbirds is not yet known. Here, we present time-resolved fluid dynamics data in combination with wingbeat kinematics data for three pied flycatchers flying across a range of speeds from near hovering to their calculated minimum power speed. Flycatchers are adapted to low speed flight, which they habitually use when catching insects on the wing. From the wake dynamics data, we constructed average wingbeat wakes and determined the time-resolved flight forces, the time-resolved downwash distributions and the resulting lift-to-drag ratios, span efficiencies and flap efficiencies. During the downstroke, slow-flying flycatchers generate a single-vortex loop wake, which is much more similar to that generated by birds at cruising flight speeds than it is to the double loop vortex wake in hovering hummingbirds. This wake structure results in a relatively high downwash behind the body, which can be explained by the relatively active tail in flycatchers. As a result of this, slow-flying flycatchers have a span efficiency which is similar to that of the birds in cruising flight and which can be assumed to be higher than in hovering hummingbirds. During the upstroke, the wings of slowly flying flycatchers generated no significant forces, but the body–tail configuration added 23 per cent to weight support. This is strikingly similar to the 25 per cent weight support generated by the wing upstroke in hovering hummingbirds. Thus, for slow-flying passerines, the upstroke cannot be regarded as inactive, and the tail may be of importance for flight efficiency and possibly manoeuvrability.     

Nonlinear instability in flagellar dynamics: a novel modulation mechanism in sperm migration?

Throughout biology, cells and organisms use flagella and cilia to propel fluid and achieve motility. The beating of these organelles, and the corresponding ability to sense, respond to and modulate this beat is central to many processes in health and disease. While the mechanics of flagellum–fluid interaction has been the subject of extensive mathematical studies, these models have been restricted to being geometrically linear or weakly nonlinear, despite the high curvatures observed physiologically. We study the effect of geometrical nonlinearity, focusing on the spermatozoon flagellum. For a wide range of physiologically relevant parameters, the nonlinear model predicts that flagellar compression by the internal forces initiates an effective buckling behaviour, leading to a symmetry-breaking bifurcation that causes profound and complicated changes in the waveform and swimming trajectory, as well as the breakdown of the linear theory. The emergent waveform also induces curved swimming in an otherwise symmetric system, with the swimming trajectory being sensitive to head shape—no signalling or asymmetric forces are required. We conclude that nonlinear models are essential in understanding the flagellar waveform in migratory human sperm; these models will also be invaluable in understanding motile flagella and cilia in other systems.     

Hydrodynamic drag constrains head enlargement for mouthbrooding in cichlids

Presumably as an adaptation for mouthbrooding, many cichlid fish species have evolved a prominent sexual dimorphism in the adult head. Since the head of fishes serves as a bow during locomotion, an evolutionary increase in head volume to brood more eggs can trade-off with the hydrodynamic efficiency of swimming. Here, the differences between males and females in three-dimensional shape and size of the external head surfaces and the effect thereof on drag force during locomotion was analysed for the Nile tilapia (Oreochromis niloticus), a maternal mouthbrooder. To do so, three-dimensional body surface reconstructions from laser scans and computational fluid dynamics simulations were performed. After scaling the scanned specimens to post-cranial body volume, in order to theoretically equalize propulsive power, the external volume of the head of females was 27% larger than that of males (head length + 14%; head width + 9%). These differences resulted in an approximate 15% increase in drag force. Yet, hydrodynamics imposed important constraints on the adaptation for mouthbrooding as a much more drastic drop in swimming efficiency seems avoided by mainly enlarging the head along the swimming direction.     

Biomechanical properties of single chondrocytes and chondrons determined by micromanipulation and finite-element modelling

A chondrocyte and its surrounding pericellular matrix (PCM) are defined as a chondron. Single chondrocytes and chondrons isolated from bovine articular cartilage were compressed by micromanipulation between two parallel surfaces in order to investigate their biomechanical properties and to discover the mechanical significance of the PCM. The force imposed on the cells was measured directly during compression to various deformations and then holding. When the nominal strain at the end of compression was 50 per cent, force relaxation showed that the cells were viscoelastic, but this viscoelasticity was generally insignificant when the nominal strain was 30 per cent or lower. The viscoelastic behaviour might be due to the mechanical response of the cell cytoskeleton and/or nucleus at higher deformations. A finite-element analysis was applied to simulate the experimental force-displacement/time data and to obtain mechanical property parameters of the chondrocytes and chondrons. Because of the large strains in the cells, a nonlinear elastic model was used for simulations of compression to 30 per cent nominal strain and a nonlinear viscoelastic model for 50 per cent. The elastic model yielded a Young''s modulus of 14 ± 1 kPa (mean ± s.e.) for chondrocytes and 19 ± 2 kPa for chondrons, respectively. The viscoelastic model generated an instantaneous elastic modulus of 21 ± 3 and 27 ± 4 kPa, a long-term modulus of 9.3 ± 0.8 and 12 ± 1 kPa and an apparent viscosity of 2.8 ± 0.5 and 3.4 ± 0.6 kPa s for chondrocytes and chondrons, respectively. It was concluded that chondrons were generally stiffer and showed less viscoelastic behaviour than chondrocytes, and that the PCM significantly influenced the mechanical properties of the cells.     

Airflow elicits a spider's jump towards airborne prey. I. Airflow around a flying blowfly

The hunting spider Cupiennius salei uses airflow generated by flying insects for the guidance of its prey-capture jump. We investigated the velocity field of the airflow generated by a freely flying blowfly close to the flow sensors on the spider''s legs. It shows three characteristic phases (I–III). (I) When approaching, the blowfly induces an airflow signal near the spider with only little fluctuation (0.013 ± 0.006 m s⁻¹) and a strength that increases nearly exponentially with time (maximum: 0.164 ± 0.051 m s⁻¹ s.d.). The spider detects this flow while the fly is still 38.4 ± 5.6 mm away. The fluctuation of the airflow above the sensors increases linearly up to 0.037 m s⁻¹ with the fly''s altitude. Differences in the time of arrival and intensity of the fly signal at different legs probably inform the spider about the direction to the prey. (II) Phase II abruptly follows phase I with a much higher degree of fluctuation (fluctuation amplitudes: 0.114 ± 0.050 m s⁻¹). It starts when the fly is directly above the sensor and corresponds to the time-dependent flow in the wake below and behind the fly. Its onset indicates to the spider that its prey is now within reach and triggers its jump. The spider derives information on the fly''s position from the airflow characteristics, enabling it to properly time its jump. The horizontal velocity of the approaching fly is reflected by the time of arrival differences (ranging from 0.038 to 0.108 s) of the flow at different legs and the exponential velocity growth rate (16–79 s⁻¹) during phase I. (III) The air flow velocity decays again after the fly has passed the spider.     

Magnetically Propelled Fish‐Like Nanoswimmers

The swimming locomotion of fish involves a complex interplay between a deformable body and induced flow in the surrounding fluid. While innovative robotic devices, inspired by physicomechanical designs evolved in fish, have been created for underwater propulsion of large swimmers, scaling such powerful locomotion into micro‐/nanoscale propulsion remains challenging. Here, a magnetically propelled fish‐like artificial nanoswimmer is demonstrated that emulates the body and caudal fin propulsion swimming mechanism displayed by fish. To mimic the deformable fish body for periodic shape changes, template‐electrosynthesized multisegment nanowire swimmers are used to construct the artificial nanofishes (diameter 200 nm; length 4.8 μm). The resulting nanofish consists a gold segment as the head, two nickel segments as the body, and one gold segment as the caudal fin, with three flexible porous silver hinges linking each segment. Under an oscillating magnetic field, the propulsive nickel elements bend the body and caudal fin periodically to generate travelling‐wave motions with speeds exceeding 30 μm s^?1. The propulsion dynamics is studied theoretically using the immersed boundary method. Such body‐deformable nanofishes exhibit a high swimming efficiency and can serve as promising biomimetic nanorobotic devices for nanoscale biomedical applications.     

Boxfish swimming paradox resolved: forces by the flow of water around the body promote manoeuvrability

The shape of the carapace protecting the body of boxfishes has been attributed an important hydrodynamic role in drag reduction and in providing automatic, flow-direction realignment and is therefore used in bioinspired design of cars. However, tight swimming-course stabilization is paradoxical given the frequent, high-performance manoeuvring that boxfishes display in their spatially complex, coral reef territories. Here, by performing flow-tank measurements of hydrodynamic drag and yaw moments together with computational fluid dynamics simulations, we reverse several assumptions about the hydrodynamic role of the boxfish carapace. Firstly, despite serving as a model system in aerodynamic design, drag-reduction performance was relatively low compared with more generalized fish morphologies. Secondly, the current theory of course stabilization owing to flow over the boxfish carapace was rejected, as destabilizing moments were found consistently. This solves the boxfish swimming paradox: destabilizing moments enhance manoeuvrability, which is in accordance with the ecological demands for efficient turning and tilting.     

流固耦合S-型自主游动柔性鱼运动特性分析

以S-型游动鱼作为对象,采用不可压缩Neo-Hookean材料描述鱼游动过程中的柔性变形及其与流体相互作用产生推力的自主游动特性。基于浸入边界法(IBM)的有限单元(FEM)离散模型,模拟了考虑鱼体肌肉作用力效应的柔性鱼在不可压缩粘性流体中自主游动的动力学过程,分析了流固耦合柔性鱼的游动机制及柔性变形的水弹性动力学特性。研究发现:随着鱼尾部不断呈S-型往复摆动,鱼体运动速度也呈波动形状,且受流固耦合的水动力学特性控制,并与鱼体的刚度密切相关。游动过程中,在鱼头的部位首先有涡旋形成,随着鱼向前游动,这些涡旋附着在鱼体表面向后滑移,并在鱼尾末端脱落形成S-型状的尾流。S-型状尾流直接影响鱼游动的速度及稳 定性。     

Larvicidal activity of green synthesized silver nanoparticles using Excoecaria agallocha L. (Euphorbiaceae) leaf extract against Aedes aegypti

Green synthesis of silver nanoparticles (AgNPs) using plant extracts has been achieved by eco‐friendly reducing and capping agents. The present study was conducted to evaluate the larvicidal efficacies of AgNPs synthesized using aqueous leaf extracts of Excoecaria agallocha against dengue vector, Aedes aegypti. The 3^rd and 4^th instar larvae of A. aegypti were exposed to various concentrations of aqueous extracts of E. agallocha, synthesized AgNPs and also crude solvent extracts (methanol and chloroform) for 24 h. The formation of AgNPs using aqueous leaf extracts was observed after 30 min with a characteristic colour change. The results recorded from UV‐Vis spectrum, XRD, FTIR, EDX, SEM and HR‐TEM were used to characterize and confirm the biosynthesis of AgNPs. The highest larvicidal efficacy of synthesized AgNPs was observed against 3^rd instar larvae at LC₅₀ 4.65 mg/L, LC₉₀ 14.17 mg/L and 4^th instar larvae with a concentration of LC₅₀ 6.10 mg/L, LC₉₀ 15.64 mg/L. A significant larvicidal activity was also observed with crude methanolic extracts against 3^rd instar larvae at a concentration LC₅₀ 41.74 mg/L, LC₉₀ 123.61 mg/L and 4^th instar larvae at a concentration of LC₅₀ 52.06 mg/L, LC₉₀ 166.40 mg/L as compared to the chloroform extract.Inspec keywords: silver, nanoparticles, nanofabrication, microorganisms, cellular biophysics, organic compounds, ultraviolet spectra, visible spectra, X‐ray diffraction, Fourier transform infrared spectra, X‐ray chemical analysis, scanning electron microscopy, transmission electron microscopyOther keywords: larvicidal activity, green synthesised silver nanoparticles, Excoecaria agallocha L. leaf extract, Aedes aegypti, plant extracts, capping agents, larvicidal efficacies, aqueous leaf extracts, excoecaria agallocha, dengue vector, Aedes aegypti, aegypti, aqueous extraction, E. agallocha, crude solvent extracts, methanol, chloroform, characteristic colour change, ultraviolet‐visible spectrum, X‐ray diffraction, Fourier‐transform infrared spectroscopy, EDX, scanning electron microscopy, high‐resolution transmission electron microscopy, AgNP biosynthesis, larvicidal efficacy, third instar larvae, instar larvae, crude methanolic extracts, chloroform extraction, time 24 h