To explore the impact of surface viscosity on coexisting fluid domains in biomembranes we consider two-phase fluid deformable surfaces as model systems for biomembranes. Such surfaces are modelled by incompressible surface Navier–Stokes–Cahn–Hilliard-like equations with bending forces. We derive this model using the Lagrange–d’Alembert principle considering various dissipation mechanisms. The highly nonlinear model is solved numerically to explore the tight interplay between surface evolution, surface phase composition, surface curvature and surface hydrodynamics. It is demonstrated that hydrodynamics can enhance bulging and furrow formation, which both can further develop to pinch-offs. The numerical approach builds on a Taylor–Hood element for the surface Navier–Stokes part, a semi-implicit approach for the Cahn–Hilliard part, higher-order surface parametrizations, appropriate approximations of the geometric quantities, and mesh redistribution. We demonstrate convergence properties that are known to be optimal for simplified subproblems.

‘Storm oil’ – nearly water-insoluble oil poured into the ocean and acting as a surfactant – has been used since ancient times to smooth the waves on the ocean. It was first scientifically described by Benjamin Franklin (Phil. Trans. R. Soc. Lond., vol. 64, 1774, pp. 445–460). In a recent paper, by combining highly controlled experiments in a wave tank and direct numerical simulations, Erinin et al. (J. Fluid Mech., vol. 972, 2023, R5) have now beautifully revealed the strong effect of soluble surfactants on the dynamics of plunging breakers. Remarkably, it is not the change in surface tension which mainly matters, but the surface tension gradient which emerges through compression and dilation of the plunging breaker surface.

Fish often swim in crystallized group formations (schooling) and orient themselves against the incoming flow (rheotaxis). At the intersection of these two phenomena, we investigate the emergence of unique schooling patterns through passive hydrodynamic mechanisms in a fish pair, the simplest subsystem of a school. First, we develop a fluid dynamics-based mathematical model for the positions and orientations of two fish swimming against a flow in an infinite channel, modelling the effect of the self-propelling motion of each fish as a point-dipole. The resulting system of equations is studied to gain an understanding of the properties of the dynamical system, its equilibria and their stability. The system is found to have five types of equilibria, out of which only upstream swimming in in-line and staggered formations can be stable. A stable near-wall configuration is observed only in limiting cases. It is shown that the stability of these equilibria depends on the flow curvature and streamwise interfish distance, below critical values of which, the system may not have a stable equilibrium. The study reveals that simply through passive fluid dynamics, in the absence of any other feedback/sensing, we can justify rheotaxis and the existence of stable in-line and staggered schooling configurations.

The monitoring of intracranial pressure (ICP) fluctuations, which is needed in the context of a number of neurological diseases, requires the insertion of pressure sensors, an invasive procedure with considerable risk factors. Intracranial pressure fluctuations drive the wave-like pulsatile motion of cerebrospinal fluid (CSF) along the compliant spinal canal. Systematically derived simplified models relating the ICP fluctuations with the resulting CSF flow rate can be useful in enabling indirect evaluations of the former from non-invasive magnetic resonance imaging (MRI) measurements of the latter. As a preliminary step in enabling these predictive efforts, a model is developed here for the pulsating viscous motion of CSF in the spinal canal, assumed to be a linearly elastic compliant tube of slowly varying section, with a Darcy pressure-loss term included to model the fluid resistance introduced by the trabeculae, which are thin collagen-reinforced columns that form a web-like structure stretching across the spinal canal. Use of Fourier-series expansions enables predictions of CSF flow rate for realistic anharmonic ICP fluctuations. The flow rate predicted using a representative ICP waveform together with a realistic canal anatomy is seen to compare favourably with in vivo phase-contrast MRI measurements at multiple sections along the spinal canal. The results indicate that the proposed model, involving a limited number of parameters, can serve as a basis for future quantitative analyses targeting predictions of ICP temporal fluctuations based on MRI measurements of spinal-canal anatomy and CSF flow rate.

To generate aerodynamic forces required for flight, two-winged insects (Diptera) move their wings back and forth at high wing-beat frequencies. This results in exceptionally high wing-stroke accelerations, and consequently relatively high acceleration-dependent fluid forces. Quasi-steady fluid force models have reasonable success in relating the generated aerodynamic forces to the instantaneous wing motion kinematics. However, existing approaches model the stroke-rate and stroke-acceleration effects independently from each other, which might be too simplified for capturing the complex unsteady aerodynamics of accelerating wings. Here, we use computational-fluid-dynamics simulations to systematically explore how aerodynamic forces and flow dynamics depend on wing-stroke rate, wing-stroke acceleration and wing-planform geometry. Based on this, we developed and calibrated a novel unsteady aerodynamic force model for insect wings with stroke accelerations. This includes improved versions of the translational-force model and the added-mass force model, and we identify a third novel component generated by the interaction of the two. This term reflects the delay in bound-circulation build-up as the wing accelerates. The physical interpretation of this effect is analogous to the Wagner effect experienced by a wing starting from rest. Here, we show that this effect can be modelled in the context of flapping wings as a stroke-acceleration-dependent correction on the translational-force model. Our revised added-mass model includes a viscous force component, which is relatively small but not negligible. We subsequently applied our new model to realistic wing-beat kinematics of hovering Dipteran insects, in a quasi-steady approach. This revealed that stroke-acceleration-related aerodynamic forces contribute substantially to lift and drag production, particularly for high-frequency flapping mosquito wings.

Sucrose is among the main products of photosynthesis that are deemed necessary for plant growth and survival. It is produced in the mesophyll cells of leaves and translocated to different parts of the plant through the phloem. Progress in understanding this transport process remains fraught with experimental difficulties, thereby prompting interest in theoretical approaches and laboratory studies. The Münch pressure and mass flow model is one of the accepted hypotheses describing the physics of sucrose transport in the phloem. It is based on osmosis creating an energy potential difference between the source and the sink. The flow responding to this energy potential is assumed laminar and described by the Hagen–Poiseuille equation. This study revisits such osmotically driven flows in tubes with membrane walls by including the effects of Taylor dispersion on mass transport. This effect has been overlooked in phloem flow studies. Taylor dispersion can increase the effective transport of solutes by increasing the apparent diffusion coefficient. It is shown that, in addition to the conventional diffusive correction derived for impermeable tube walls, a new adjustment to the mean advective terms arises because of osmotic effects. Because the molecular Schmidt number is very large for sucrose in water, the sucrose front speed and travel times have a direct dependence on the Péclet number for different ranges of the Münch number. This study establishes upper limits on expected Taylor dispersion enhancement of sucrose transport.

We investigate aspects of the spherical squirmer model employing both large-scale numerical simulations and asymptotic methods when the squirmer is placed in weakly elastic fluids. The fluids are modelled by differential equations, including the upper-convected Maxwell (UCM)/Oldroyd-B, finite-extensibility nonlinear elastic model with Peterlin approximation (FENE-P) and Giesekus models. The squirmer model we examine is characterized by two dimensionless parameters related to the fluid velocity at the surface of the micro-swimmer: the slip parameter $\xi $ and the swirl parameter $\zeta $. We show that, for swimming in UCM/Oldroyd-B fluids, the elastic stress becomes singular at a critical Weissenberg number, Wi, that depends only on $\xi$. This singularity for the UCM/Oldroyd-B models is independent of the domain exterior to the swimmer, or any other forces considered in the momentum balance for the fluid – we believe that this is the first time such a singularity has been explicitly demonstrated. Moreover, we show that the behaviour of the solution at the poles is purely extensional in character and is the primary reason for the singularity in the Oldroyd-B model. When the Giesekus or the FENE-P models are utilized, the singularity is removed. We also investigate the mechanism behind the speed and rotation rate enhancement associated with the addition of swirl in the swimmer's gait. We demonstrate that, for all models, the speed is enhanced by swirl, but the mechanism of enhancement depends intrinsically on the rheological model employed. Special attention is paid to the propulsive role of the pressure and elucidated upon. We also study the region of convergence of the perturbation solutions in terms of Wi. When techniques that accelerate the convergence of series are applied, transformed solutions are derived that are in very good agreement with the results obtained by simulations. Finally, both the analytical and numerical results clearly indicate that the low-Wi region is more important than one would expect and demonstrate all the major phenomena observed when swimming with swirl in a viscoelastic fluid.

This work analyses the viscous flow and elastic deformation created by the forced axial motion of a rigid cylinder within an elastic liquid-filled tube. The examined configuration is relevant to various minimally invasive medical procedures in which slender devices are inserted into fluid-filled biological vessels, such as vascular interventions, interventional radiology, endoscopies and laparoscopies. By applying the lubrication approximation, thin shell elastic model, as well as scaling analysis and regular and singular asymptotic schemes, the problem is examined for small and large deformation limits (relative to the gap between the cylinder and the tube). At the limit of large deformations, forced insertion of the cylinder is shown to involve three distinct regimes and time scales: (i) initial shear dominant regime, (ii) intermediate regime of dominant fluidic pressure and a propagating viscous-peeling front, (iii) late-time quasi-steady flow regime of the fully peeled tube. A uniform solution for all regimes is presented for a suddenly applied constant force, showing initial deceleration and then acceleration of the inserted cylinder. For the case of forced extraction of the cylinder from the tube, the negative gauge pressure reduces the gap between the cylinder and the tube, increasing viscous resistance or creating friction due to contact of the tube and cylinder. Matched asymptotic schemes are used to calculate the dynamics of the near-contact and contact limits. We find that the cylinder exits the tube in a finite time for sufficiently small or large forces. However, for an intermediate range of forces, the radial contact creates a steady locking of the cylinder inside the tube.

We study the flow induced by eye rotations in the anterior chamber (AC) of the eye, the region between the cornea and the iris. We model the geometry of the AC as a thin domain sitting on the surface of a sphere, and study both the simpler case of a constant-height domain as well as a more realistic AC shape. We model eye rotations as harmonic in time with prescribed frequency $\unicode[STIX]{x1D714}_{f}$ and amplitude $\unicode[STIX]{x1D6FD}$, and use lubrication theory to simplify the governing equations. We write the equations in a reference frame moving with the domain and show that fluid motion is governed by three dimensionless parameters: the aspect ratio $\unicode[STIX]{x1D716}$ of the AC, the angular amplitude $\unicode[STIX]{x1D6FD}$ and the Womersley number $\unicode[STIX]{x1D6FC}$. We simplify the equations under the physiologically realistic assumptions that $\unicode[STIX]{x1D716}$ is small and $\unicode[STIX]{x1D6FC}$ large, leading to a linear system that can be decomposed into three harmonics: a dominant frequency component, with frequency $\unicode[STIX]{x1D714}_{f}$, and a steady streaming component and a third component with frequency $2\unicode[STIX]{x1D714}_{f}$. We solve the problem analytically for the constant-height domain and numerically as the solution of ordinary differential equations in the more realistic geometry. Both the primary flow and the steady streaming are shown to have a highly three-dimensional structure, which has not been highlighted in previous numerical works. We show that the steady streaming is particularly relevant from the clinical point of view, as it induces fluid mixing in the AC. Furthermore, the steady flow component is the dominant mixing mechanism during the night, when the thermal flow induced by temperature variations across the AC is suppressed.

This paper investigates the transport of a solute carried by the cerebrospinal fluid (CSF) in the spinal canal. The analysis is motivated by the need for a better understanding of drug dispersion in connection with intrathecal drug delivery (ITDD), a medical procedure used for treatment of some cancers, infections and pain, involving the delivery of the drug to the central nervous system by direct injection into the CSF via the lumbar route. The description accounts for the CSF motion in the spinal canal, described in our recent publication (Sánchez et al., J. Fluid Mech., vol. 841, 2018, pp. 203–227). The Eulerian velocity field includes an oscillatory component with angular frequency $\unicode[STIX]{x1D714}$, equal to that of the cardiac cycle, and associated tidal volumes that are a factor $\unicode[STIX]{x1D700}\ll 1$ smaller than the total CSF volume in the spinal canal, with the small velocity corrections resulting from convective acceleration providing a steady-streaming component with characteristic residence times of order $\unicode[STIX]{x1D700}^{-2}\unicode[STIX]{x1D714}^{-1}\gg \unicode[STIX]{x1D714}^{-1}$. An asymptotic analysis for $\unicode[STIX]{x1D700}\ll 1$ accounting for the two time scales $\unicode[STIX]{x1D714}^{-1}$ and $\unicode[STIX]{x1D700}^{-2}\unicode[STIX]{x1D714}^{-1}$ is used to investigate the prevailing drug-dispersion mechanisms and their dependence on the solute diffusivity, measured by the Schmidt number $S$. Convective transport driven by the time-averaged Lagrangian velocity, obtained as the sum of the Eulerian steady-streaming velocity and the Stokes-drift velocity associated with the non-uniform pulsating flow, is found to be important for all values of $S$. By way of contrast, shear-enhanced Taylor dispersion, which is important for values of $S$ of order unity, is shown to be negligibly small for the large values $S\sim \unicode[STIX]{x1D700}^{-2}\gg 1$ corresponding to the molecular diffusivities of all ITDD drugs. Results for a model geometry indicate that a simplified equation derived in the intermediate limit $1\ll S\ll \unicode[STIX]{x1D700}^{-2}$ provides sufficient accuracy under most conditions, and therefore could constitute an attractive reduced model for future quantitative analyses of drug dispersion in the spinal canal. The results can be used to quantify dependences of the drug-dispersion rate on the frequency and amplitude of the pulsation of the intracranial pressure, the compliance and specific geometry of the spinal canal and the molecular diffusivity of the drug.

The present study is aimed at assessing the existing results concerning the stability of canonical shear flows in channels and tubes with deformable walls, in light of consistent formulations of the nonlinear solid constitutive model and linearised interface conditions at the fluid–solid interface. We show that a class of unstable shear-wave modes at low Reynolds number, predicted by previous studies for pressure-driven flows through neo-Hookean tubes and channels, is absent upon use of consistent interfacial conditions. Furthermore, we analyse the consequences of the change in solid model on the stability of the canonical shear flows by using both neo-Hookean and Mooney–Rivlin models. We show that the salient features of the stability of the system are adequately captured by a consistent formulation of the neo-Hookean solid model, thus precluding the need to employ more detailed solid models. The stability analysis of planar flows past a neo-Hookean solid subjected to three-dimensional disturbances showed that two-dimensional disturbances are more unstable than the corresponding three-dimensional disturbances within the consistent formulations. We show that prior inconsistent formulations of the solid constitutive equation predict a physically spurious spanwise instability in disagreement with experiments thereby demonstrating their inapplicability to predict instabilities in flow past deformable solid surfaces. Using the consistent formulation, the present work provides an accurate picture, over a range of Reynolds numbers, of the stability of canonical shear flows through deformable channels and tubes. Importantly, it is shown how inconsistencies in either the bulk constitutive relation or in the linearisation of the interface conditions can separately lead to physically spurious instabilities. The predictions of this work are relevant to experimental studies in flow through deformable tubes and channels in the low and moderate Reynolds number regime.

Leaves falling in air and marine larvae settling in water are examples of unsteady descents due to complex interactions between gravitational and aerodynamic forces. Understanding passive flight is relevant to many branches of engineering and science, ranging from estimating the behaviour of re-entry space vehicles to analysing the biomechanics of seed dispersion. The motion of regularly shaped objects falling freely in homogenous fluids is relatively well understood. However, less is known about how density stratification of the fluid medium affects passive flight. In this paper, we experimentally investigate the descent of heavy discs in stably stratified fluids for Froude numbers of order 1 and Reynolds numbers of order 1000. We specifically consider fluttering descents, where the disc oscillates as it falls. In comparison with pure water and homogeneous saltwater fluid, we find that density stratification significantly enhances the radial dispersion of the disc, while simultaneously decreasing the vertical descent speed, fluttering amplitude and inclination angle of the disc during descent. We explain the physical mechanisms underlying these observations in the context of a quasi-steady force and torque model. These findings could have significant impact on the design of unpowered vehicles and on the understanding of geological and biological transport where density and temperature variations may occur.

The individual and combined influences of aspect ratio ($A$), Reynolds number ($Re$) and Rossby number ($Ro$) on the leading-edge vortex (LEV) of a rotating wing of insect-like planform are investigated numerically. A previous study from our group has determined the wingspan to be an appropriate length scale governing the large-scale LEV structure. In this study, the $A$ range considered is further extended, to show that this scaling works well as $A$ is varied by a factor of 4 ($1.8\leqslant A\leqslant 7.28$) and over a $Re$ range of two orders of magnitude. The present study also extends this scaling for wings with an offset from the rotation axis, which is typically the case for actual insects and often for experiments. Remarkably, the optimum range of $A$ based on the lift coefficients at different $Re$ coincides with that observed in nature. The scaling based on the wingspan is extended to the acceleration terms of the Navier–Stokes equations, suggesting a modified scaling of $Ro$, which decouples the effects of $A$. A detailed investigation of the flow structures, by increasing $Ro$ in a wide range, reveals the weakening of the LEV due to the reduced spanwise flow, resulting in a reduced lift. Overall, the use of span-based scaling of $Re$ and $Ro$, together with $A$, may help reconcile apparent conflicting trends between observed variations in aerodynamic performance in different sets of experiments and simulations.

Micro-organisms encounter heterogeneous viscous environments consisting of networks of obstacles embedded in a viscous fluid medium. In this paper we analyse the characteristics of swimming in a porous medium modelled by the Brinkman equation via a spherical squirmer model. The idealized geometry allows an analytical and exact solution of the flow surrounding a squirmer. The propulsion speed obtained agrees with previous results using the Lorentz reciprocal theorem. Our analysis extends these results to calculate the power dissipation and hence the swimming efficiency of the squirmer in a Brinkman medium. The analytical solution enables a systematic analysis of the structure of the flow surrounding the squirmer, which can be represented in terms of singularities in Brinkman flows. We also discuss the spatial decay of flows due to squirming motion in a Brinkman medium in comparison with the decay in a purely viscous fluid. The results lay the foundation for subsequent studies on hydrodynamic interactions, nutrient transport and uptake by micro-organisms in heterogeneous viscous environments.

The effect of air viscosity on the flow around an insect wing increases as insect size decreases. For the smallest insects (wing length $R$ below 1 mm), the viscous effect is so large that lift-generation mechanisms used by their larger counterparts become ineffective. How the weight-supporting vertical force is generated is unknown. To elucidate the aerodynamic mechanisms responsible, we measure the wing kinematics of the tiny wasp Encarsia formosa (0.6 mm $R$) in hovering or very slow ascending flight and compute and analyse the aerodynamic forces. We find that the insects perform two unusual wing motions. One is ‘rowing’: the wings move fast downward and backward, like stroking oars. The other is the previously discovered Weis-Fogh ‘fling’. The rowing produces 70 % of the required vertical force and the Weis-Fogh ‘fling’ the other 30 %. The oaring wing mainly produces an approximately up-pointing drag, resulting in the vertical force. Because each oaring produces a starting flow, the drag is unsteady in nature and much greater than that in steady motion at the same velocities and angles of attack. Furthermore, our computation shows that if the tiny wasps employed the usual wing kinematics of the larger insects (flapping back and forth in a horizontal plane), the vertical force produced would be only $1/3$ of that by the real wing kinematics; i.e. they must use the special wing movements to overcome the problem of large viscous effects encountered by the commonly used flapping kinematics. We have observed for the first time very small insects using drag to support their weight and we explain how a net vertical force is generated when the drag principle is applied.

While hydrodynamic interactions for aggregates of swimmers have received significant attention in the low Reynolds number realm ($Re\ll 1$), there has been far less work at higher Reynolds numbers, in which fluid and body inertia are involved. Here we study the collective behaviour of multiple self-propelled plates in tandem configurations, which are driven by harmonic flapping motions of identical frequency and amplitude. Both fast modes with compact configurations and slow modes with sparse configurations were observed. The Lighthill conjecture that orderly configurations may emerge passively from hydrodynamic interactions was verified on a larger scale with up to eight plates. The whole group may consist of subgroups and individuals with regular separations. Hydrodynamic forces experienced by the plates near their multiple equilibrium locations are all springlike restoring forces, which stabilize the orderly formation and maintain group cohesion. For the cruising speed of the whole group, the leading subgroup or individual plays the role of ‘leading goose’.

The propagation of periodically generated vortex rings (period $T$) is numerically investigated by imposing pulsed jets of velocity $U_{jet}$ and duration $T_{s}$ (no flow between pulses) at the inlet of a cylinder of diameter $D$ exiting into a tank. Because of the step-like nature of pulsed jet waveforms, the average jet velocity during a cycle is $U_{ave}=U_{jet}T_{s}/T$. By using $U_{ave}$ in the definition of the Reynolds number ($Re=U_{ave}D/\unicode[STIX]{x1D708}$, $\unicode[STIX]{x1D708}$: kinematic viscosity of fluid) and non-dimensional period ($T^{\ast }=TU_{ave}/D=T_{s}U_{jet}/D$, i.e. equivalent to formation time), then based on the results, the vortex ring velocity $U_{v}/U_{jet}$ becomes approximately independent of the stroke ratio $T_{s}/T$. The results also show that $U_{v}/U_{jet}$ increases by reducing $Re$ or increasing $T^{\ast }$ (more sensitive to $T^{\ast }$) according to a power law of the form $U_{v}/U_{jet}=0.27T^{\ast 1.31Re^{-0.2}}$. An empirical relation, therefore, for the location of vortex ring core centres ($S$) over time ($t$) is proposed ($S/D=0.27T^{\ast 1+1.31Re^{-0.2}}t/T_{s}$), which collapses (scales) not only our results but also the results of experiments for non-periodic rings. This might be due to the fact that the quasi-steady vortex ring velocity was found to have a maximum of 15 % difference with the initial (isolated) one. Visualizing the rings during the periodic state shows that at low $T^{\ast }\leqslant 2$ and high $Re\geqslant 1400$ here, the stopping vortices become unstable and form hairpin vortices around the leading ones. However, by increasing $T^{\ast }$ or decreasing $Re$ the stopping vortices remain circular. Furthermore, rings with short $T^{\ast }=1$ show vortex pairing after approximately one period in the downstream, but higher $T^{\ast }\geqslant 2$ generates a train of vortices in the quasi-steady state.

Observations in experiments and simulations show that the kinematic behaviour of an elastic capsule, suspended and rotating in shear flow, depends upon the flow strength, the capsule membrane material properties and its at-rest shape. We develop a linear stability description of the periodically rotating base state of this coupled system, as represented by a boundary integral flow formulation with spherical harmonic basis functions describing the elastic capsule geometry. This yields Floquet multipliers that classify the stability of the capsule motion for elastic capillary numbers $Ca$ ranging from $Ca=0.01$ to 5. Viscous dissipation rapidly damps most perturbations. However, for all cases, a single component grows or decays slowly, depending upon $Ca$, over many periods of the rotation. The transitions in this stability behaviour correspond to the different classes of rotating motion observed in previous studies.

As a first step towards understanding the distribution of swimming micro-organisms in flowing shallow water containing vegetation, we formulate a continuum model for dilute suspensions in horizontal shear flow, with a maximum Reynolds number of 100, past a single, rigid, vertical, circular cylinder that extends from a flat horizontal bed and penetrates the free water surface. A numerical platform was developed to solve this problem, in four stages: first, a scheme for computation of the flow field; second, a solver for the Fokker–Planck equation governing the probability distribution of the swimming direction of gyrotactic cells under the combined action of gravity, ambient vorticity and rotational diffusion; third, the construction of a database for the mean swimming velocity and the translational diffusivity tensor as functions of the three vorticity components, using parameters appropriate for the swimming alga, Chlamydomonas nivalis; fourth, a solver for the three-dimensional concentration distribution of the gyrotactic micro-organisms. Upstream of the cylinder, the cells are confined to a vertical strip of width equal to the cylinder diameter, which enables us to visualise mixing in the wake. The flow downstream of the cylinder is divided into three zones: parallel vortex shedding in the top zone near the water surface, oblique vortex shedding in the middle zone and quasi-steady flow in the bottom zone. Secondary (vertical) flow occurs just upstream and downstream of the cylinder. Frequency spectra of the velocity components in the wake of the cylinder show two dominant frequencies of vortex shedding, in the parallel- and oblique-shedding zones respectively, together with a low frequency, equal to the difference between those two frequencies, that corresponds to a beating modulation. The concentration distribution is calculated for both active particles and passive, non-swimming, particles for comparison. The concentration distribution is very similar for both active and passive particles, except near the top surface, where upswimming causes the concentration of active particles to reach values greater than in the upstream strip, and in a thin boundary layer on the downstream surface of the cylinder, where a high concentration of active particles occurs as a result of radial swimming.