Microorganisms, such as spermatozoa, exhibit rich behaviours when in close proximity to each other. However, their locomotion is not fully understood when coupled mechanically and hydrodynamically. In this study, we develop hydrodynamic models to investigate the locomotion of paired spermatozoa, predicting the fine structure of their swimming. Experimentally, sperm pairs are observed to transition between different modes of flagellar synchronisation: in-phase, anti-phase and lagged synchronisation. Using our models, we assess their swimming performances in these synchronisation modes in terms of average swimming speed, average power consumption, and swimming efficiency. The swimming performances of paired spermatozoa are shown to depend on their flagellar phase lag, flagellar waveforms, and the mechanical coupling between their heads.

Cilia perform various functions, including sensing, locomotion, generation of fluid flows and mass transport, serving to underpin a vast range of biological and ecological processes. However, analysis of the mass transport typically fails to resolve the near-field dynamics around individual cilia, and therefore overlooks the intricate role of power/recovery strokes of ciliary motion. Selvan et al. (2023, Phys. Rev. Fluids 8, 123103) observed that the flow field due to a point torque (i.e. a rotlet) accurately resolves both the near- and far-field characteristics of a single cilium’s flow in a semi-infinite domain. In this paper, we calculate the mass transport between a no-slip boundary and an adjacent fluid, as a model system for nutrient exchange with ciliated tissues. We develop a Langevin model in the presence of a point torque (i.e. a single cilium) to examine the nutrient flux from a localised surface source. This microscopic transport model is validated using a macroscopic continuum model, which directly solves the advection–diffusion equation. Our findings reveal that the flow induced by a point torque can enhance the particles’ transport, depending on their diffusivity and the magnitude of the point torque. Additionally, the average mass transport affected by a single cilium can be enhanced or diminished by the presence of an externally imposed linear shear flow, with a strong dependence on the alignment of the cilium. Taken together, this framework serves as a useful minimal model for examining the average nutrient exchange between ciliated tissues and fluid environments.

The Liebau effect generates a net flow without the need for valves. For the Liebau effect pumping phenomenon to occur, the pump must have specific characteristics. It needs tubes with different elastic properties and an actuator to provide energy to the fluid. The actuator periodically compresses the more flexible element. Furthermore, asymmetry is a crucial factor that differentiates between two pumping mechanisms: impedance pumping and asymmetric pumping. In this work, a model based on the fluid dynamics of an asymmetric valveless pump under resonant conditions is proposed to determine which parameters influence the pumped flow rate. Experimental work is used to validate the model, after which each of the parameters involved in the pump performance is dimensionlessly analysed. This highlights the most significant parameters influencing the pump performance such as the actuator period, length tube ratio and tube diameters. The results point out ways to increase a valveless asymmetric pump’s net-propelled flow rate, which has exciting applications in fields such as biomedicine. The model also allows for predicting the resonance period, a fundamental operating parameter for asymmetric pumping.

Intermittent swimming behaviour is commonly observed in larval zebrafish, often attributed to energy-saving mechanisms. In this study, we utilize a hybrid approach combining deep reinforcement learning and the immersed boundary–lattice Boltzmann method to train a larval zebrafish-like swimmer to reach a target with minimized energy expenditure. We find that when the tail-beat period is fixed, continuous swimming emerges as the optimal strategy. However, when the tail-beat period is allowed to vary, intermittent swimming proves superior in energy performance, achieved through reductions in tail-beat amplitude and frequency. Our detailed analysis reveals that intermittent swimmers employ rapid backward tail flicks to attain high speeds, coupled with slower forward tail flicks and coasting phases to conserve energy. Furthermore, we derive scaling laws governing the swimming performance of trained fish. These results offer valuable insights into the intermittent swimming patterns of fish, with implications for understanding bio-inspired locomotion and informing the design of energy-efficient aquatic systems.

Although the interaction between microswimmers and walls during near-wall swimming has been extensively studied, the effect of microswimmer shapes and slip boundary conditions on the dynamic characteristics of near-wall microswimmers has received less attention. In this study, elliptical microswimmer models have been developed with various aspect ratios based on circular microswimmers. The lattice Boltzmann method has been used for the numerical simulation of the dynamic behaviour of microswimmers near walls. Under slip boundary conditions, the escape or capture of microswimmers by the walls is influenced by the swimming Reynolds number (Res), wall slip length (ls) and the aspect ratio (Cab) of a microswimmer. Changes in the Cab value of a microswimmer considerably affect its swimming state, especially for puller-type microswimmers. The tendency of pullers to be captured by the wall increases with increasing Cab. Moreover, changes in ls within the slip boundary condition of a puller can induce a transition in its movement state from a wall oscillation state to a stable sliding state and eventually to a wall lock-up state, a process influenced by the Cab value of the puller. Pusher-type microswimmers show a considerably increased tendency to escape from walls with increasing Cab and no wall lock-up state is observed, which is opposite to the case of pullers. Pushers and pullers show an increased tendency to be captured by the wall with increasing initial swimming angle of the microswimmer. The findings of this study enhance our understanding of the swimming patterns of natural microswimmers near walls and are of substantial importance for the design of artificial microswimmers and microfluidic devices.

Wave-assisted propulsion (WAP) systems directly convert wave energy into thrust using elastically mounted hydrofoils. The wave conditions as well as the design of the hydrofoil drive the fluid–structure interaction of the hydrofoil and, consequently, its performance. We employ simulations using a sharp-interface immersed boundary method to examine the effect of three key parameters on the flow physics, the fluid–structure interaction as well as thrust performance of these systems – the stiffness of the torsional spring, the location of the pitch axis and the Strouhal number. We demonstrate the utility of ‘maps’ of energy exchange between the flow and the hydrofoil system, as a way to understand and predict these characteristics. The force-partitioning method (FPM) is used to decompose the pressure forces into interpretable components and to quantify the mechanisms associated with thrust generation. Based on the results from FPM, a phenomenological model for the thrust generated by the WAP foil is presented. The parameters associated with this model are estimated based on data from over 450 distinct simulations. The predictions of the model are compared with the simulations and the use of this model for guiding WAP design is discussed.

Numerous flying and swimming creatures use the ground effect to boost their propulsive performance, with the ‘ground’ referring to either a solid boundary or a free surface. While our knowledge of how a solid boundary affects biolocomotion is relatively comprehensive, the ground effect of a free surface is not fully understood. To address this limitation, we conduct a numerical investigation on the propulsion performance of a flapping plate under a free surface, subject to a range of control parameters. When the Froude number ($Fr$) is very low (i.e. little surface deformation), the effects of a free surface are similar to those of a solid boundary, with enhanced thrust and input power but little change in efficiency. However, as $Fr$ increases (i.e. more surface deformation), our results reveal an optimal $Fr$ of approximately 0.6, where the free surface induces a more streamlined flow around the flapping plate, effectively reducing the added mass. This results in a significant decrease in input power and greatly enhanced efficiency.

Linear unsteady aerofoil theory, while successfully used for the prediction of unsteady aerofoil lift for many decades, has yet to be proven adequate for predicting the propulsive performance of oscillating aerofoils. In this paper we test the hypothesis that the central shortcoming of linear small-amplitude models, such as the Garrick function, is the failure to account for the flow acceleration caused by aerofoil thrust. A new analytical model is developed by coupling the Garrick function to a cycle-averaged actuator disc model, in a manner analogous to the blade-element momentum theory for wind turbines and propellers. This amounts to assuming the Garrick function to be locally valid and, in combination with a global control volume analysis, enables the prediction of flow acceleration at the aerofoil. The new model is demonstrated to substantially improve the agreement with large-eddy simulations of an aerofoil in combined heave and pitch motion.

Single-flagellated bacteria are ubiquitous in nature. They exhibit various swimming modes using their flagella to explore complex surroundings such as soil and porous polymer networks. Some single-flagellated bacteria swim with two distinct modes, one with the flagellum extended away from its body and another with the flagellum wrapped around it. The wrapped mode has been observed when bacteria swim under tight confinements or in highly viscous polymeric melts. In this study we investigate the hydrodynamics of these two modes inside a circular pipe. We find that the wrapped mode is slower than the extended mode in bulk but more efficient under strong confinement due to a hydrodynamic increase of its flagellum translation–rotation coupling and an Archimedes’ screw-like configuration that helps to move the fluid along the pipe.

Biological microorganisms and artificial micro-swimmers often locomote in heterogeneous viscous environments consisting of networks of obstacles embedded into viscous fluid media. In this work, we use the squirmer model and present a numerical investigation of the effects of shape on swimming in a heterogeneous medium. Specifically, we analyse the microorganism's propulsion speed as well as its energetic cost and swimming efficiency. The analysis allows us to probe the general characteristics of swimming in a heterogeneous viscous environment in comparison with the case of a purely viscous fluid. We found that a spheroidal microorganism always propels faster, expends less energy and is more efficient than a spherical microorganism in either a homogeneous fluid or a heterogeneous medium. Moreover, we determined that above a critical eccentricity, a spheroidal microorganism in a heterogeneous medium can swim faster than a spherical microorganism in a homogeneous fluid. Based on an analysis of the forces acting on the squirmer, we offer an explanation for the decrease in the squirmer's speed observed in heterogeneous media compared with homogeneous fluids.

Solid particles trapped in an acoustic standing wave have been observed to undergo propulsion. This phenomenon has been attributed to the generation of a steady streaming flow, with a reversal in the propulsion direction at a distinct frequency. We explain the mechanism underlying this reversal by considering the canonical problem of a sphere executing oscillatory rotation in an unbounded fluid that undergoes rectilinear oscillation; these two oscillations occur at identical frequency but with an arbitrary phase difference. Two distinct bifurcations in the flow field occur: (1) a stagnation point first forms with increasing frequency, which (2) splits into a saddle node and a vortex centre. Reversal in the propulsion direction is driven by reversal in the flow far from the sphere, which coincides with the second bifurcation. This flow is identified with that of a Stokeslet whose strength is the net force exerted on the particle, which has implications for studying the flow field around particles of non-spherical geometries and for modelling suspensions of particles in acoustic fields.

The squirmer is a popular model to analyse the fluid mechanics of a self-propelled object, such as a micro-organism. We demonstrate that some fore–aft symmetric squirmers can spontaneously self-propel above a critical Reynolds number. Specifically, we numerically study the effects of inertia on spherical squirmers characterised by an axially and fore–aft symmetric ‘quadrupolar’ distribution of surface-slip velocity; under creeping-flow conditions, such squirmers generate a pure stresslet flow, the stresslet sign classifying the squirmer as either a ‘pusher’ or ‘puller’. Assuming axial symmetry, and over the examined range of the Reynolds number $Re$ (defined based upon the magnitude of the quadrupolar squirming), we find that spontaneous symmetry breaking occurs in the puller case above $Re \approx 14.3$, with steady swimming emerging from that threshold consistently with a supercritical pitchfork bifurcation and with the swimming speed growing monotonically with $Re$.

We investigate numerically the propulsion characteristics of an oscillating foil undergoing coupled heave and pitch motion in a linearly density-stratified flow. A parameter space defined by the internal Froude number ($1 \le Fr \le 10$) and the maximum angle of attack ($5^\circ \le {\alpha _0} \le 30^\circ$) is considered in our study. The results demonstrate a significant enhancement in both thrust production and propulsive efficiency due to the stratification influence. Notably, the highest efficiency exceeding $80\,\%$ is achieved under moderate stratification conditions, surpassing the performance observed in a homogeneous fluid. We attribute this optimum performance to the proper match between the stratification effect and foil kinematics, which gives rise to intense vortex interactions and sufficient wave–mean flow interactions in the near wake of the oscillating foil. Consequently, the energy is transferred towards wake structures to form a high-intensity momentum jet in close proximity to the foil's trailing edge, indicating efficient propulsion. Furthermore, we find that the stratifications within the moderate-to-strong transitional regime display a reduced dependence of propulsive efficiency on the maximum angle of attack, primarily due to the delaying and alleviating effects on dynamic-stall events. Such a mechanism enables the oscillating foil to maintain a satisfactory performance by sufficiently high angles of attack without the penalty of stall events. Based on our findings, we propose that animals or artificial vehicles utilising oscillatory propulsion can benefit from the presence of density stratification in the surrounding fluid.

Various systems of mechanical devices and natural organisms use the repeated expansion and contraction of deformable structures to draw in and blow out fluid. Instead of such deformable continuous structures, a system that consists of multiple discrete bodies can also generate a directional flow through the cooperative movement of the individual bodies. In this study, we numerically investigate the collective effects of a multi-body system composed of eight circular cylinders, each of which oscillates separately in the radial direction to generate thrust. The cylinder array performs cooperative motion regulated by three motion parameters: phase difference, oscillation amplitude and frequency at a low Reynolds number ($Re = 10$). The phase difference between the cylinders is critical in determining the extent of the directional flow and the time-averaged thrust. The optimal phase difference that yields the maximum time-averaged thrust is consistent, regardless of the oscillation amplitude and frequency. However, the thrust generation performance becomes significantly weaker at a higher Reynolds number ($Re = 100$). This highlights that the hydrodynamic blockage in gaps between cylinders, which is induced by strong viscous diffusion at the low Reynolds number, is essential for the cooperative force generation of multiple closely spaced bodies. A new dimensionless geometric parameter based on the motion of the array is proposed to characterize the degree of bias in the generated flow and it successfully predicts the trend in the time-averaged thrust at the low Reynolds number with strong hydrodynamic blockage.

Natural flyers and swimmers employ flexible wings or fins to propel. While the complex interaction between the foil with deformation and the surrounding non-steady fluid environment defines the propulsion performance of the propellers, elucidating the interaction mechanism through theoretical models earns much challenge. Based on elastokinetics and linear potential flow theory, this study proposes a simplified analytical model to clarify the kinematics and the propulsion performance of a flexible thin foil pitching in flow. The dynamical forces, including the inertial force of the foil and the non-steady fluid pressure, are used to determine the averaged deformation angle of the foil. Combining the averaged deformation angle and the prescribed driving pitching motion, the kinematics of the foil is resolved analytically. Based on the analytical expressions for the corresponding pitching motion, analytical relations among the physical parameters of the stiffness and the mass of the foil and the driving frequency are given to these critical conditions, including resonance of the flow–structure system, equal pitching amplitude between the flexible foil and the rigid counterpart, phase angle transition from ${\rm \pi}/2$ to $- {\rm \pi}/2$. Subsequently, the performance of the foil, including the thrust, the power and the propulsive efficiency, as a function of the flexibility of the foil are derived, together with the introduction of a bluff body type offset drag to the thrust. The formulated analytical theory, which matches nicely with previous reports, will help to interpret the effect of the flexibility and regulate the propulsive performance of the flexible foil when pitching in fluid.

A classic lift decomposition (Von Kármán & Sears, J. Aeronaut. Sci., vol. 5, 1938, pp. 379–390) is conducted on potential flow simulations of a near-ground pitching hydrofoil. It is discovered that previously observed stable and unstable equilibrium altitudes are generated by a balance between positive wake-induced lift and negative quasi-steady lift while the added mass lift does not play a role. Using both simulations and experiments, detailed analyses of each lift component's near-ground behaviour provide further physical insights. When applied to three-dimensional pitching hydrofoils the lift decomposition reveals that the disappearance of equilibrium altitudes for ${A{\kern-4pt}R}\ {\rm (aspect\ ratio)} <1.5$ occurs due to the magnitude of the quasi-steady lift outweighing the magnitude of the wake-induced lift at all ground distances. Scaling laws for the quasi-steady lift, wake-induced lift and the stable equilibrium altitude are discovered. A simple scaling law for the lift of a steady foil in ground effect is derived. This scaling shows that both circulation enhancement and the velocity induced at a foil's leading edge by the bound vortex of its ground image foil are the essential physics to understand steady ground effect. The scaling laws for unsteady pitching foils can predict the equilibrium altitude to within $20\,\%$ of its value when $St\ {\rm (Strouhal\ number)} < 0.45$. For $St \ge 0.45$ there is a wake instability effect, not accounted for in the scaling relations, that significantly alters the wake-induced lift. These results not only provide key physical insights and scaling laws for steady and unsteady ground effects, but also for two schooling hydrofoils in a side-by-side formation with an out-of-phase synchronization.

In this study, the effects of antagonistic muscle actuation on the propulsion of a bilaminar-structure fish fin ray were investigated using a two-dimensional computational flow–structure interaction (FSI) model. The structure and material properties of the model were based on the realistic biological data of the sunfish fin. The effect of muscle actuation was modelled using root displacement offset between the two hemitrichs. Parametric FSI simulations were conducted by assuming a sinusoidal function of the offset over a cycle and varying the amplitude and phase difference between the actuations and pitching/plunging motions. The results show that the phase of muscle actuation is a critical factor affecting its effects. Three performance regions can be identified with different phase ranges, including a thrust-favour region, an efficiency-favour region and a thrust-efficiency-unfavour region. In each region, the relationships among the root actuations, fin-ray kinematics, vortex dynamics and resulting performance are studied and discussed. Furthermore, a strong positive correlation between the trailing–leading amplitude ratio and thrust coefficient as well as a negative relationship between the efficiency and angle of attack at the centre of mass of the fin ray are observed.

Metachronal rowing is a biological propulsion mechanism employed by many swimming invertebrates (e.g. copepods, ctenophores, krill and shrimp). Animals that swim using this mechanism feature rows of appendages that oscillate in a coordinated wave. In this study, we used observations of a swimming ctenophore (comb jelly) to examine the hydrodynamic performance and vortex dynamics associated with metachronal rowing. We first reconstructed the beating kinematics of ctenophore appendages based on a high-speed video of a metachronally coordinated row. Following the reconstruction, two numerical models were developed and simulated using an in-house immersed-boundary-method-based computational fluid dynamics solver. The two models included the original geometry (16 appendages in a row) and a sparse geometry (8 appendages, formed by removing every other appendage along the row). We found that appendage tip vortex interactions contribute to hydrodynamic performance via a vortex-weakening mechanism. Through this mechanism, appendage tip vortices are significantly weakened during the drag-producing recovery stroke. As a result, the swimming ctenophore produces less overall drag, and its thrust-to-power ratio is significantly improved (up to 55.0 % compared with the sparse model). Our parametric study indicated that such a propulsion enhancement mechanism is less effective at higher Reynolds numbers. Simulations were also used to investigate the effects of substrate curvature on the unsteady hydrodynamics. Our results illustrated that, compared with a flat substrate, arranging appendages on a curved substrate can boost the overall thrust generation by up to 29.5 %. These findings provide new insights into the fluid dynamic principles of propulsion enhancement underlying metachronal rowing.

Self-propulsion of chemically active droplets and phoretic disks has been studied widely; however, most research overlooks the influence of disk shape on swimming dynamics. Inspired by experimentally observed prolate composite droplets and elliptical camphor disks, we employ simulations to investigate the phoretic dynamics of an elliptical disk that emits solutes uniformly in the creeping flow regime. By varying the disk's eccentricity $e$ and the Péclet number $Pe$, we distinguish five disk behaviours: stationary, steady, orbiting, periodic and chaotic. We perform a linear stability analysis (LSA) to predict the onset of instability and the most unstable eigenmode when a stationary disk transitions spontaneously to steady self-propulsion. In addition to the LSA, we use an alternative approach to determine the perturbation growth rate, illustrating the competing roles of advection and diffusion. The steady motion features a transition from a puller-type to a neutral-type swimmer as $Pe$ increases, which occurs as a bimodal concentration profile at the disk surface shifts to a polarized solute distribution, driven by convective solute transport. An elliptical disk achieves an orbiting motion through a chiral symmetry-breaking instability, wherein it repeatedly follows a circular path while simultaneously rotating. The periodic swinging motion, emerging from a steady motion via a supercritical Hopf bifurcation, is characterized by a wave-like trajectory. We uncover a transition from normal diffusion to superdiffusion as eccentricity $e$ increases, corresponding to a random-walking circular disk and a ballistically swimming elliptical counterpart, respectively.

Three-dimensional schools of hydrodynamically axisymmetric swimmers self-propelling at a constant velocity are studied. We introduce a low-order model for the induced velocity based on the far-field approximation. We inquire if, by holding suitable relative positions in the three-dimensional space, the swimmers can reduce the overall energy consumption of the school in comparison with the same number of isolated individuals at the same velocity. We find a considerable (several per cent) energy saving achievable by chain formations. The benefit increases asymptotically with the number of individuals, towards a finite limit that is a function of the minimum allowed spacing between each pair of neighbours.