We propose a hybrid numerical model for the collective motion of fish groups, which integrates an agent-based model with a computational fluid dynamics (CFD)-based model. In the agent-based model, the fish group is represented by self-propelled particles (SPPs), incorporating social forces with local interactive rules. The CFD-based model treats the fish body with an undulated filament that responds to the hydrodynamic forces imposed by the surrounding fluid flow. These two models are coupled using a central pattern generator controller. We test this hybrid model with groups of 30–50 individuals. The results show that the group exhibits various collective behaviours, including tight schooling, sparse schooling and milling patterns, by adjusting the coefficients in the SPP model. Due to the hydrodynamic interactions, particularly with the obstacle avoidance model, both the individuals’ and the group’s mean speed fluctuate, differentiating it from traditional SPP models that typically consider volumeless particles. More interestingly, our findings indicate that fish benefit from collective motion in terms of energetic consumption in both schooling and milling patterns. It is important to note that the swimming fish are actuated using a very simple mechanism without any optimisation strategy. An additional study investigates the effects of the Reynolds number, demonstrating the capability of the current hybrid model to account for fish groups of varying body lengths or swimming speeds. Future applications of this model are promising, offering potential insights into the energetic advantages of collective motion in large-scale fish groups.

In a quiescent medium, chemically active particles propel themselves by emitting or absorbing solutes, creating concentration gradients that induce a slip at the particle surface. This self-propulsion occurs when solute advection overcomes diffusion. However, an imposed flow field can alter these dynamics. This study explores the propulsion characteristics and the related rheological consequences of chemically active particles in an imposed uniaxial extensional flow analytically and numerically. An asymptotic solution is obtained for weak imposed flow relative to self-induced diffusiophoretic slip. Meanwhile, finite element simulations are carried out over a wide range of imposed flow strength and Péclet number. The results reveal that the interplay between solute advection, imposed flow and diffusiophoretic slip significantly affects particle propulsion and suspension rheology. While solute advection and diffusiophoretic slip tend to create asymmetric solute distributions, promoting self-propulsion, imposed extensional flow promotes symmetric distributions, hindering self-propulsion. This not only delays the start of self-propulsion but also results in an early transition from a propulsion state to a stationary state characterised by an abrupt halt at relatively lower Péclet number compared to a quiescent medium. Post the abrupt halt, a stirring effect induced by particle activity and imposed extensional flow results in an increased magnitude of stresslet, thus a sudden change in the effective viscosity of the active suspension. The effect of imposed extensional flow on active particle dynamics and suspension rheology can be described succinctly by categorising the overall dynamics into three separate regimes, determined by the Péclet number and the intensity of the extensional flow.

The acoustofluidic method holds great promise for manipulating micro-organisms. When exposed to the steady vortex structures of acoustic streaming flow, these micro-organisms exhibit intriguing dynamic behaviours, such as hydrodynamic trapping and aggregation. To uncover the mechanisms behind these behaviours, we investigate the swimming dynamics of both passive and active particles within a two-dimensional acoustic streaming flow. By employing a theoretically calculated streaming flow field, we demonstrate the existence of stable bounded orbits for particles. Additionally, we introduce rotational diffusion and examine the distribution of particles under varying flow strengths. Our findings reveal that active particles can laterally migrate across streamlines and become trapped in stable bounded orbits closer to the vortex centre, whereas passive particles are confined to movement along the streamlines. We emphasise the influence of the flow field on the distribution and trapping of active particles, identifying a flow configuration that maximises their aggregation. These insights contribute to the manipulation of microswimmers and the development of innovative biological microfluidic chips.

Many particles, whether passive or active, possess elongated shapes. When these particles settle or swim in shear flows, they often form regions of accumulation and depletion. Additionally, the density contrast between the particles and the fluid can further alter the flow by increasing the local suspension density, resulting in a two-way buoyancy–flow coupling mechanism. This study investigates the buoyancy–flow coupled dispersion of active spheroids, examining the effects of elongation, orientation-dependent settling and gyrotaxis in a vertical pipe subjected to either downwards or upwards discharge. While the concentration and velocity profiles of passive settling spheroids and spherical gyrotactic swimmers can be analysed similarly to a recent study, notable differences in dispersion characteristics emerge due to different streamline-crossing mechanisms. For suspensions of elongated swimmers, the interplay between orientation-dependent settling, gyrotaxis-induced accumulation and shear-induced trapping results in distinct concentration and velocity distributions compared to those of neutrally buoyant particles and extremely dilute suspensions with negligible coupling effect. These differences further impact drift velocity, dispersivity, and the time elapsed to steady dispersion under varying flow rates. Interestingly, low-shear trapping of non-settling elongated swimmers around the centreline, commonly observed in planar Poiseuille flow, is absent in the vertical pipe due to the change of confinement from reflectional to rotational symmetry. However, elongated settling swimmers show a non-trivial concentration response to strong downwelling discharge. This phenomenon, linked to the centreline accumulation of passive settling spheroids, bears similarities to low-shear trapping observed in planar Poiseuille flow.

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.

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 both butterflies and dragonflies are four-winged insects, their wing geometries and kinematics differ significantly. Butterflies have a much narrower gap between their forewing and hindwing than dragonflies. While previous research has extensively investigated the forewing–hindwing interactions in dragonfly flight, this work focuses on their interactions in butterfly flight. The interactions are studied based on numerical simulations of the Navier–Stokes equations around a butterfly-inspired flapping wing with an adjustable slot, representing the narrow gap between the forewing and hindwing. The slot is controlled by a dihedral angle between the forewing and hindwing. The lift coefficients of wings with different slot sizes and locations are investigated in detail. The results show that the forewing–hindwing interactions can significantly enhance the lift if the slot is properly configured. When the slot is configured by elevating the forewing at a 10-degree dihedral angle relative to the hindwing during flapping flight, the wing generates over 20 % more lift than the model without a slot. The streamwise ram effect and tip-vortex capture are shown to be responsible for the lift enhancement by using a lift decomposition formula. The streamwise ram effect reduces the streamwise velocity beneath the forewing, decreasing the negative vortex lift associated with spanwise vorticity. The tip-vortex capture enhances the positive vortex lift associated with streamwise vorticity when the hindwing captures the tip vortex shedding from the forewing.

Aircraft with bio-inspired flapping wings that are operated in low-density atmospheric environments encounter unique challenges associated with the low density. The low density results in the requirement of high operating velocities of aircraft to generate sufficient lift resulting in significant compressibility effects. Here, we perform numerical simulations to investigate the compressibility effects on the lift generation of a bio-inspired wing during hovering flight using an immersed boundary method. The aim of this study is to develop a scaling law to understand how the lift is influenced by the Reynolds and Mach numbers, and the associated flow physics. Our simulations have identified a critical Mach number of approximately $0.6$ defined by the average wing-tip velocity. When the Mach number is lower than 0.6, compressibility does not have significant effects on the lift or flow fields, while when the Mach number is greater than $0.6$, the lift coefficient decreases linearly with increasing Mach number, due to the drastic change in the pressure on the wing surface caused by unsteady shock waves. Moreover, the decay rate is dependent on the Reynolds number and the angle of attack. Based on these observations, we propose a scaling law for the lift of a hovering flapping wing by considering compressible and viscous effects, with the scaled lift showing excellent collapse.

Hydrodynamic interactions between swimming or flying organisms can lead to complex flows on the scale of the group. These emergent fluid dynamics are often more complex than a linear superposition of individual organism flows, especially at intermediate Reynolds numbers. This paper presents an approach to estimate the flow induced by multiple swimmer wakes in proximity using a semianalytical model that conserves mass and momentum in the aggregation. The key equations are derived analytically, while the implementation and solution of these equations are carried out numerically. This model was informed by and compared with empirical measurements of induced vertical migrations of brine shrimp, Artemia salina. The response of individual swimmers to ambient background flow and light intensity was evaluated. In addition, the time-resolved three-dimensional spatial configuration of the swimmers was measured using a recently developed laser scanning system. Numerical results using the model found that the induced flow at the front of the aggregation was insensitive to the presence of downstream swimmers, with the induced flow tending towards asymptotic beyond a threshold aggregation length. Closer swimmer spacing led to higher induced flow speeds, in some cases leading to model predictions of induced flow exceeding swimmer speeds required to maintain a stable spatial configuration. This result was reconciled by comparing two different models for the near-wake of each swimmer. The results demonstrate that aggregation-scale flows result from a complex, yet predictable interplay between individual organism wake structure and aggregation configuration and size.

We analyse the motion of a flagellated bacterium in a two-fluid medium using slender body theory. The two-fluid model is useful for describing a body moving through a complex fluid with a microstructure whose length scale is comparable to the characteristic scale of the body. This is true for bacterial motion in biological fluids (entangled polymer solutions), where the entanglement results in a porous microstructure with typical pore diameters comparable to or larger than the flagellar bundle diameter, but smaller than the diameter of the bacterial head. Thus, the polymer and solvent satisfy different boundary conditions on the flagellar bundle and move with different velocities close to it. This gives rise to a screening length $L_B$ within which the fluids exchange momentum and the relative velocity between the two fluids decays. In this work, both the solvent and polymer of the two-fluid medium are modelled as Newtonian fluids with different viscosities $\mu _s$ and $\mu _p$ (viscosity ratio $\lambda = \mu _p/\mu _s$), thereby capturing the effects solely introduced by the microstructure of the complex fluid. From our calculations, we observe an increased drag anisotropy for a rigid, slender flagellar bundle moving through this two-fluid medium, resulting in an enhanced swimming velocity of the organism. The results are sensitive to the interaction between the bundle and the polymer, and we discuss two physical scenarios corresponding to two types of interaction. Our model provides an explanation for the experimentally observed enhancement of swimming velocity of bacteria in entangled polymer solutions and motivates further experimental investigations.

We present new constrained and free-swimming experiments and simulations in the inertial regime, with Reynolds number $\mbox{Re} = O(10^4)$, of a pair of two-dimensional and three-dimensional pitching hydrofoils interacting in a minimal school. The hydrofoils have an out-of-phase synchronisation, and they are varied through in-line, staggered and side-by-side formations within the two-dimensional interaction plane. It is discovered that there is a two-dimensionally stable equilibrium point for a side-by-side formation. This formation is super-stable, meaning that hydrodynamic forces will passively maintain this formation even under external perturbations, and the school as a whole has no net forces acting on it that cause it to drift to one side or the other. Previously discovered one-dimensionally stable equilibria driven by wake vortex interactions are shown to be, in fact, two-dimensionally unstable, at least for an out-of-phase synchronisation. Additionally, it is discovered that a trailing-edge vortex mechanism provides the restorative force to stabilise a side-by-side formation. The stable equilibrium is further verified by experiments and simulations for freely swimming foils where dynamic recoil motions are present. When constrained, swimmers in compact side-by-side formations experience collective efficiency and thrust increases up to 40 % and 100 %, respectively, whereas slightly staggered formations output an even higher efficiency improvement of 84 %, with an 87 % increase in thrust. Freely swimming foils in a stable side-by-side formation show efficiency and speed enhancements of up to 9 % and 15 %, respectively. These newfound schooling performance and stability characteristics suggest that fluid-mediated equilibria may play a role in the control strategies of schooling fish and fish-inspired robots.

Direct numerical simulation is performed for flow separation over a bump in a turbulent channel. Comparisons are made between a smooth bump and one where the lee side is covered with replicas of shark denticles – dermal scales that consist of a slender base (the neck) and a wide top (the crown). As flow over the bump is under an adverse pressure gradient (APG), a reverse pore flow is formed in the porous cavity region underneath the crowns of the denticle array. Remarkable thrust is generated by the reverse pore flow as denticle necks accelerate the fluid passing between them in the upstream direction. Several geometrical features of shark denticles, including some that had not previously been considered hydrodynamically functional, are identified to form the two-layer denticle structure that enables and sustains the reverse pore flow and thrust generation. The reverse pore flow is activated by the APG before massive flow detachment. The results indicate a proactive, on-demand drag reduction mechanism that leverages and transforms the APG into a favourable outcome.

The effects of the evolution of vortices on the aeroacoustics generated by a hovering wing are numerically investigated by using a hybrid method of an immersed boundary–finite difference method for the three-dimensional incompressible flows and a simplified model based on the Ffowcs Williams-Hawkings acoustic analogy. A low-aspect-ratio ($AR=1.5$) rectangular wing at low Reynolds ($Re=1000$) and Mach ($M=0.04$) numbers is investigated. Based on the simplified model, the far-field acoustics is shown to be dominated by the time derivative of the pressure on the wing surface. Results show that vortical structure evolution in the flow fields, which is described by the divergence of the convection term of the incompressible Navier–Stokes equations in a body-fixed reference frame, determines the time derivative of the surface pressure and effectively the far-field acoustics. It dominates over the centrifugal acceleration and Coriolis acceleration terms in determining the time derivative of the surface pressure. The position of the vortex is also found to affect the time derivative of the surface pressure. A scaling analysis reveals that the vortex acoustic source is scaled with the cube of the flapping frequency.

Spanwise vortex instability and the growth of secondary hairpin-like vortical structures in the wake of an oscillating foil are investigated numerically at Reynolds number 8000 in a range of chord-based Strouhal number ($0.32 \le St_c \le 0.56$). The phase-offset ($\phi$) between the heaving and pitching motion is $\phi = 90^\circ$. The wake at the lowest $St_c$ (0.32) is characterized by a single system of streamwise hairpin-like structures that evolve from the core vorticity outflux of the secondary leading edge vortex (LEV) over the foil boundary. The primary LEV features spanwise dislocations, but it does not reveal substantial changes advecting downstream. Increasing $St_c$ beyond 0.32 reveals that the transition in spanwise instability characterizes the deformation of primary LEV cores, which subsequently transforms to hairpin-like secondary structures. At higher $St_c$, stronger trailing edge vortices (TEVs) grow in close proximity to the primary LEVs, which contributes to an enhanced localized vortex compression and tilting near dislocations. This phenomenon amplifies the undulation amplitude of primary LEVs, eventually leading to vortex tearing. The larger circulation of TEVs with increasing $St_c$ provides an additional explanation for an accelerated vortex compression that coincides with a faster transition of spanwise LEV instability to secondary hairpin-like structures in the wake.

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.

Flow control of a low-aspect-ratio flat-plate heaving wing at an average angle of attack of $10^{\circ }$ by a steady-blowing jet is numerically studied by using a feedback immersed boundary–lattice Boltzmann method. Blowing jets at the leading edge, mid-chord and trailing edge are considered. The wing enjoys the highest lift production with the trailing-edge downstream blowing jet, which improves the average lift by 50.0 % at $Re = 1000$ and 22.9 % at $Re = 5000$ through the enhancement of the tip vortex circulation caused by the increase in the mass flux of the shear layer at the wing tips. This increase in mass flux decreases as $Re$ increases from 1000 to 5000 due to its self-limiting mechanism. A mid-chord vertical blowing jet induces a middle vortex which enhances the lift production but the enhancement is smaller than that of trailing-edge downstream blowing jet. Other jet arrangements do not significantly increase the lift coefficient, but the mid-chord upstream blowing jet experiences a significant reduction in the drag coefficient, leading to an increase of 50.6 % in the average lift-to-drag ratio. The effectiveness of the flow control is not significantly affected by the aspect ratio.

Flying insects exhibit remarkable capabilities in coordinating their olfactory sensory system and flapping wings during odour plume-tracking flights. While observations have indicated that their flapping wing motion can ‘sniff’ up the incoming plumes for better odour sampling range, how flapping motion impacts the odour concentration field around the antennae is unknown. Here, we reconstruct the body and wing kinematics of a forwards-flying butterfly based on high-speed images. Using an in-house computational fluid dynamics solver, we simulate the unsteady flow field and odourant transport process by solving the Navier–Stokes and odourant advection-diffusion equations. Our results show that, during flapping flight, the interaction between wing leading-edge vortices and antenna vortices strengthens the circulation of antenna vortices by over two-fold compared with cases without flapping motion, leading to a significant increase in odour intensity fluctuation along the antennae. Specifically, the interaction between the wings and antennae amplifies odour intensity fluctuations on the antennae by up to 8.4 fold. This enhancement is critical in preventing odour fatigue during odour-tracking flights. Further analysis reveals that this interaction is influenced by the inter-antennal angle. Adjusting this angle allows insects to balance between resistance to odour fatigue and the breadth of odour sampling. Narrower inter-antennal angles enhance fatigue resistance, while wider angles extend the sampling range but reduce resistance. Additionally, our findings suggest that while the flexibility of the wings and the thorax's pitching motion in butterflies do influence odour fluctuation, their impact is relatively secondary to that of the wing–antenna interaction.

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.

During stroke reversals, insect wings interact with their own wake flow from the preceding half-stroke, resulting in an unsteady aerodynamic mechanism known as ‘wing–wake interaction’ or ‘wake capture’. To better elucidate this mechanism, we numerically solved the incompressible Navier–Stokes equations at Reynolds numbers $10^2$ and $10^3$. Simulations were conducted for wing planforms defined using the beta function distribution with varying aspect ratios ($AR=2\unicode{x2013}6$) and radial centroid locations ($\hat {r}_1=0.4\unicode{x2013}0.6$), whilst employing representative normal hovering kinematics. The wake development from the considered flapping wing planforms was investigated, and the wake capture contribution to aerodynamic force production was quantified by comparing the force generation between the fifth and first stroke cycles at multiple sections along the wingspan. Our results revealed that on the inboard wing region experiencing an attached leading-edge vortex (LEV) structure, wing–wake interaction is dominated by an unsteady downwash effect, resulting in a reduction in local force production. However, in regions closer to the wingtip experiencing detachment of the LEV, wing–wake interaction is dominated by an unsteady upwash effect, leading to an increase in local force production. Consequently, the global wake capture force production is controlled by the extent of LEV detachment, which primarily increases with the increase of wing aspect ratio. This suggests that for normal hovering flapping wings, the typical loss in translational force production due to wingtip stall is partially mitigated by wake capture effects.

Understanding the propulsion of a swimmer in a large group of individuals holds the key to unravelling the intriguing dynamics of active matter collective motion. Here, we develop a two-dimensional (2-D) self-assembled rotor, powered by bacterial flagella. At a water–air interface, the average direction of rotation of a rotor is fixed. When the chiral rotor is put into a 2-D bacterial suspension, we examine the average and fluctuation of the angular velocity of the rotor. Remarkably, the average angular velocity of a rotor is found to increase up to 3 times when the density of surrounding bacterial suspension increases and the increase is nonlinear. In a dense suspension of bacteria, the existence of a rotor disrupts vortices in the surrounding active turbulence, and the acceleration of the rotor is independent of the activity level of the surrounding free bacteria. The nonlinear acceleration thus results from hydrodynamic interaction with surrounding crowdedness that can be quantitatively explained by hydrodynamic simulation. The simultaneity between the acceleration of rotor and free bacteria in active turbulence suggests that crowding-induced acceleration may promote the onset of instability. The result will inspire new active-matter-based microfluidic devices with improved transport properties.