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Organisms often swim through fluids that are spatially inhomogeneous. If the fluids are polymeric, gradients in polymer concentration may lead to gradients in both fluid viscosity and elasticity. In this letter, we present theoretical results for the dynamics of active particles, biological or otherwise, swimming through spatially inhomogeneous viscoelastic fluids. We model the active particles using the squirmer model, and show that spatial variations in fluid relaxation time lead to a novel mechanism for reorientation and taxis in viscoelastic fluids, which we refer to as a form of durotaxis in fluids., Comment: 5 pages, 1 figure

We study equilibrium states of a drop between flexible sheets clamped on both ends. Revisiting first the case of parallel sheets, we find multiple equilibria which we classify in a parameter space. In solution branching diagrams we identify hysteresis cycles and folds indicating abrupt transitions, yet not necessarily leading to channel collapse. Between nonparallel sheets, a drop can stay in equilibrium away from the ends even when the liquid is totally wetting, which is impossible between nonparallel rigid plates. We also show that nonparallel sheets can delay or prevent collapse in comparison to straight channels, suggesting thus a mechanism that protects slender structures inside micro-devices from damage by surface tension., Comment: 25 pages, 8 figures

We find the mixing induced by a small swimming organism in a density stratified fluid. We model the swimmer as a spherical squirmer and quantify mixing through a mixing efficiency that is the ratio of rate of change of potential energy of fluid to the total work done by the swimmer. Assessing mixing in the near field and far field of the swimmer separately, we find that the near-field mixing aligns with past work, but the overall mixing is much larger than that caused by a point-sized swimmer (like force-dipole), although still small in weak stratification. Equivalent results are also obtained for a homogeneous dilute suspension of non-interacting swimmers. Our study highlights the impact of swimmer size on mixing, revealing that small-sized swimmers cannot induce substantial mixing. However, we propose few (unexplored) pathways through which small swimmers could induce significant mixing, ultimately contributing towards oceanic mixing., Comment: 11 pages

A new gradient-based adaptive sampling method is proposed for design of experiments applications which balances space filling, local refinement, and error minimization objectives while reducing reliance on delicate tuning parameters. High order local maximum entropy approximants are used for metamodelling, which take advantage of boundary-corrected kernel density estimation to increase accuracy and robustness on highly clumped datasets, as well as conferring the resulting metamodel with some robustness against data noise in the common case of unreplicated experiments. Two-dimensional test cases are analyzed against full factorial and latin hypercube designs and compare favourably. The proposed method is then applied in a unique manner to the problem of adaptive spatial resolution in time-varying non-linear functions, opening up the possibility to adapt the method to solve partial differential equations., Comment: 33 pages, 14 figures

In this note, we study the effect of viscosity gradients on the energy dissipated by the motion of microswimmers and the associated efficiency of that motion. Using spheroidal squirmer model swimmers in weak linearly varying viscosity fields, we find that efficiency depends on whether they generate propulsion from the back (pushers) or the front (pullers). Pushers are faster and more efficient when moving down gradients but slower and less efficient moving up viscosity gradients, and the opposite is true for pullers. However, both pushers and pullers display negative viscotaxis, therefore pushers dynamically tend to the most efficient orientation while pullers the least. We also evaluate the effect of shape on power expenditure and efficiency when swimming in viscosity gradients and find that in general the change in both due to gradients monotonically decreases with increasing slenderness. This work shows how shape and gait play an important role in determining dynamics and efficiency in inhomogeneous environments, and demonstrating that both efficiency minimizing and maximizing stable dynamical states are possible., Comment: 12 pages, 1 figure

Active suspensions encompass a wide range of complex fluids containing microscale energy-injecting particles, such as cells, bacteria or artificially powered active colloids. Because they are intrinsically non-equilibrium, active suspensions can display a number of fascinating phenomena, including turbulent-like large-scale coherent motion and enhanced diffusion. Here, using a recently developed active Fast Stokesian Dynamics method, we present a detailed numerical study on the hydrodynamic diffusion in apolar active suspensions. Specifically, we simulate suspensions of active but non-self-propelling spherical squirmers, of either puller- or pusher-type, at volume fractions from 0.5% to 55%. Our results show little difference between pullers and pushers in their instantaneous and long-time dynamics, where the translational dynamics vary non-monotonically with the volume fraction, with a peak diffusivity at around 10% to 20%, in stark contrast to suspensions of self-propelling particles. On the other hand, the rotational dynamics tend to increase with the volume fraction as is the case for self-propelling particles. To explain these dynamics, we provide detailed scaling and statistical analyses based on the activity-induced hydrodynamic interactions and the observed microstructural correlations, which display a weak local order. Overall, these results elucidate and highlight the different effects of particle activity on the collective dynamics and transport phenomena in active fluids., Comment: This version replaces the previous ones that were unfortunately misguided by wrong data

In this paper, we explore the hydrodynamics of spheroidal active particles in viscosity gradients. This work provides a more accurate modeling approach, in comparison to spherical particles, for anisotropic organisms like Paramecium swimming through inhomogeneous environments, but more fundamentally examines the influence of particle shape on viscotaxis. We find that spheroidal squirmers generally exhibit dynamics consistent with their spherical analogs, irrespective of the classification of swimmers as pushers, pullers, or neutral swimmers. However, the slenderness of the spheroids tends to reduce the impact of viscosity gradients on their dynamics; when swimmers become more slender, the viscosity difference across their body is reduced, which leads to slower reorientation. We also derive the mobility tensor for passive spheroids in viscosity gradients generalizing previous results for spheres and slender bodies. This work enhances our understanding of how shape factors into the dynamics of passive and active particles in viscosity gradients, and offers new perspectives that could aid the control of both natural and synthetic swimmers in complex fluid environments., Comment: 19 pages, 5 figures

We study the imbibition of a wetting liquid between flexible sheets that are fixed on both ends. Assuming a narrow gap between the sheets, we solve the lubrication equation coupled with slender body deformation. When the sheets are parallel, we find that the deformation initially speeds up the flow, as shown in previous studies, but only up to the middle of the channel. As the channel contracts, the hydrodynamic resistance increases and ultimately slows down the filling process. Below a threshold stiffness, the channel collapses and imbibition stops. We propose a scaling of the filling duration near this threshold. Next we show that if the sheets are initially tilted with a minimal angle, the channel avoids collapse. The liquid front pulls the diverging sheets and spreads in a nearly parallel portion, which maintains the capillary propulsion and enhances the wicking. Therefore, while it is established that diverging rigid plates imbibe liquids slower than parallel ones, we show that elasticity reverses this principle: diverging flexible sheets imbibe liquids faster than parallel ones. We find an optimal tilt angle that gives the shortest filling time., Comment: 11 pages, 7 figures

Organisms often swim through density stratified fluids. In this Letter, we investigate the dynamics of small active particles swimming in density gradients and report theoretical evidence of taxis as a result of density stratification ($\textit{densitaxis}$). Specifically, we calculate the effect of density stratification on the dynamics of a force-free spherical squirmer and show that density stratification induces reorientation that tends to align swimming either parallel or normal to the gradient depending on the swimming gait. In particular, particles that propel by generating thrust in the front (pullers) rotate to swim parallel to gradients and hence display (positive or negative) densitaxis, while particles that propel by generating thrust in the back (pushers) rotate to swim normal to the gradients. This work could be useful to understand the motion of marine organisms in ocean, or be leveraged to sort or organize a suspension of active particles by modulating density gradients., Comment: 13 pages, 6 figures

Cytoskeletal gels are prototyped to reproduce the mechanical contraction of the cytoskeleton in-vitro. They are composed of a polymer network (backbone), swollen by the presence of a liquid solvent, and active molecules (molecular motors, MMs) that transduce chemical energy into the mechanical work of contraction. These motors attach to the polymer chains to shorten them and/or act as dynamic crosslinks, thereby constraining the thermal fluctuation of the chains. We describe both mechanisms thermodynamically as a microstructural reconfiguration, where the backbone stiffens to motivate solvent (out)flow and accommodate contraction. Via simple steady-state energetic analysis, under the simplest case of isotropic contraction, we quantify the mechanical energy required to achieve contraction as a function of polymer chain density and molecular motor density. We identify two limit cases, (fm) fast MM activation for which MMs provide all the available mechanical energy instantaneously and leave the polymer in a stiffened state, i.e. their activity occurs at a time scale that is much smaller than solvent diffusion, and (sm) slow MM activation for which the MM activation timescale is much longer. To achieve the same final contracted state, fm requires the largest amount of work per unit reference volume, while sm requires the least. For all intermediate cases where the timescale of MM activation is comparable with that of solvent flow, the required work ranges between the two cases. We provide all these quantities as a function of chain density and MM density. Finally, we compare our results with experiments and observe good agreement.

We present an experimental investigation of turbulent drag reduction with flexible and rigid polymer solutions. The flexible polymer is partially hydrolyzed polyacrylamide (HPAM) and the rigid polymer is xanthan gum (XG). The experiments are carried out at low drag reduction ($\%DR < 40$), high drag reduction ($\%DR > 40$) and maximum drag reduction (where the velocity profile $U^+$ roughly matches Virk's asymptote). We compare velocity profiles, streamwise and wall-normal Reynolds stresses and power spectra of streamwise velocity fluctuations measured by Laser Doppler Anemometry (LDA). Our results show that the effects of both XG and HPAM polymers on turbulence are similar, provided that the Reynolds numbers and $\%DR$ are also similar. At high levels of $\%DR$, the power spectral densities of streamwise velocity fluctuations of both XG and HPAM flows show a power-law decay near $-3$ instead of $-5/3$ in the inertial range. A slope of the power spectra of $-3$ was recently interpreted as evidence of elasto-inertial turbulence (EIT) in polymer jets. At relatively low concentrations, we observe that flexible polymer solutions are more effective at reducing drag, while XG only reaches MDR at very high concentrations. Thus, we hypothesize that the formation of polymer aggregates with higher concentrations contributes to increase viscoelasticity, and thus $\%DR$, with XG and other rigid polymer solutions.

We analyze a dilute suspension of active particles confined between walls and subjected to fields that can modulate particle speed as well as orientation. Generally, the particle distribution is different in the bulk compared to near the walls. In the bulk, particles tend to accumulate in the regions of low speed, but in the presence of an orienting field, particles rotate to align with the field and accumulate downstream in the field direction. At the walls, particles tend to accumulate pointing into the walls and thereby exert pressure on walls. But the presence of strong orienting fields can cause the particles to reorient away from the walls, and hence shows a possible mechanism for preventing contamination of surfaces. The pressure at the walls depends on the wall separation and the field strengths. This work demonstrates how multiple fields with different functionalities can be used to control active matter under confinement., Comment: 13 pages, 7 figures

Active particles (living or synthetic) often move through inhomogeneous environments, such as gradients in light, heat or nutrient concentration, that can lead to directed motion (or taxis). Recent research has explored inhomogeneity in the rheological properties of a suspending fluid, in particular viscosity, as a mechanical (rather than biological) mechanism for taxis. Theoretical and experimental studies have shown that gradients in viscosity can lead to reorientation due to asymmetric viscous forces. In particular, recent experiments with Chlamydomonas reinhardtii algae swimming across sharp viscosity gradients have observed that the microorganisms are redirected and scattered due to the viscosity change. Here we develop a simple theoretical model to explain these experiments. We model the swimmers as spherical squirmers and focus on small, but sharp, viscosity changes. We derive a law, analogous to Snell's law of refraction, that governs the orientation of active particles in the presence of a viscosity interface. Theoretical predictions show good agreement with experiments and provide a mechanistic understanding of the observed reorientation process., Comment: 12 pages, 5 figures

Turbulent flows of viscoplastic fluids at high Reynolds numbers have been investigated recently with direct numerical simulations (DNS) but experimental results have been limited. For this reason, we carry out an experimental study of fully turbulent flows of a yield stress fluid in a rectangular aspect ratio channel with a high-resolution laser doppler velocimetry (LDA) setup. We employ aqueous Carbopol solutions, often considered to be a simple yield stress fluid. We formulate different concentrations to address the effect of the rheology of the fluid on the turbulence statistics at an approximately constant Reynolds number. Additionally, we also perform experiments with a single Carbopol formulation at different Reynolds numbers to study its effect. The flow analysis is performed via rheology measurements, turbulence statistics and power spectral densities of velocity fluctuations. The addition of Carbopol to the flow increases turbulence anisotropy, with an enhancement of streamwise velocity fluctuations and a decrease in wall normal velocity fluctuations in comparison to water at the same mean velocity. This change is reflected on the power spectral densities of streamwise velocity fluctuations, where we observe a large increase in energy of large scale turbulent structures. Conversely, the energy of smaller scales is decreased in comparison to water, where the energy drops with a steeper scale than the Newtonian power law of $k_x^{-5/3}$. As we increase the Reynolds number with a Carbopol solution, the streamwise Reynolds stresses approach Newtonian values in the core, which suggests diminishing effects of shear-thinning. The power spectral densities reveal that the energy content at larger scales decreases slightly with the Reynolds number. However, the shear thinning effects do not disappear even as the Reynolds number approaches 50000.

The rheology of non-colloidal suspensions under cyclic shear is studied numerically. The main findings are a strain amplitude ($\gamma_0$) dependent response in the shear stress and second normal stress difference ($N_2$). Specifically, we find a reduced viscosity, an enhanced intracycle shear thinning, the onset of a finite $N_2$ and its frequency doubling, all near a critical strain amplitude $\gamma_c$ that scales with the volume fraction $\phi$ as $\gamma_c \sim \phi^{-2}$. These rheological changes also signify a reversible-irreversible transition (RIT), dividing stroboscopic particle dynamics into a reversible absorbing phase (for $\gamma_0<\gamma_c$) and a persistently diffusing phase (for $\gamma_0>\gamma_c$). We explain the results based on two flow-induced mechanisms and elucidate their connection in the context of RIT through the underlying microstructure, which tends towards hyperuniformity near $\gamma_0=\gamma_c$. Overall, we expect this correspondence between rheology and emergent dynamics to hold in a wide range of settings where structural organizations are dominated by volume exclusions., Comment: 9 pages, 7 figures

Long-chained, viscoelastic surfactant solutions (VES) have been widely employed in the oil and gas industry, particularly in hydraulic fracturing and gravel-packing operations, where turbulence is commonly reached due to high pumping rates. With this motivation, we experimentally investigate the turbulent duct flow of an under-studied class of wormlike micellar solutions that forms a gel at room temperature. The fluid is characterized via rotational rheometry, and the turbulent velocity and Reynolds stress profiles are measured via Laser Doppler Anemometry (LDA). Three surfactant concentrations are investigated at increasing Reynolds numbers. The turbulent flow fields of water, and semi-dilute solutions of partially hydrolyzed polyacrylamide (HPAM) and xanthan gum (XG) are used as comparisons. Our study reveals that the gel-like structure of the wormlike micellar gel is mostly broken down during turbulent flow, especially in the near-wall region where the results indicate the presence of a water layer. Turbulent flow at low concentrations of surfactant show a Newtonian-like flow field throughout most of the duct, where the energy spectra shows a -5/3 power law scale with wavenumber, whereas higher concentrations lead to drag reduction and lower power spectral densities at large wavenumbers. A comparison of the flow of polymeric fluids and the wormlike micellar solutions at maximum drag reduction (MDR) shows comparable drag-reduction effects, with a large decrease in Reynolds shear stresses, and increased turbulence anisotropy in the buffer layer due to the large streamwise fluctuations and near-zero wall-normal fluctuations. Additionally, the MDR regime was bounded by Virk's asymptote for both polymers and the micellar gel, which implies a similar mechanism of drag reduction at MDR.

Since its development, Stokesian Dynamics has been a leading approach for the dynamic simulation of suspensions of particles at arbitrary concentrations with full hydrodynamic interactions. Although originally developed for the simulation of passive particle suspensions, the Stokesian Dynamics framework is equally well suited to the analysis and dynamic simulation of suspensions of active particles, as we elucidate here. We show how the reciprocal theorem can be used to formulate the exact dynamics for a suspension of arbitrary active particles and then show how the Stokesian Dynamics method provides a rigorous way to approximate and compute the dynamics of dense active suspensions where many-body hydrodynamic interactions are important., Comment: 16 pages, 3 figures

In this work, we analyze the motion of an active particle, modeled as a spherical squirmer, in linearly varying viscosity fields. In general, the presence of a particle will disturb a background viscosity field and the disturbance generated depends on the boundary conditions imposed by the particle on the viscosity field. We find that, irrespective of the details of the disturbance, active squirmer-type particle tend to align down viscosity gradients (negative viscotaxis). However, the rate of rotation and the swimming speed along the gradient do depend on the details of the interaction of the particle and the background viscosity field. In addition, we explore the relative importance on the dynamics of the local viscosity changes on the surface of active particles versus the (nonlocal) changes in the flow field due to spatially varying viscosity (from that of a homogeneous fluid). We show that the relative importance of local versus nonlocal effects depends crucially on the boundary conditions imposed by the particle on the field. This work demonstrates the dangers in neglecting the disturbance of the background viscosity caused by the particle as well as in using the local effects alone to capture the particle motion., Comment: 12 pages, 3 figures

Long-chained surfactant solutions have found widespread use in the oil and gas industry due to a host of attractive properties. In this paper, we characterize one such commercially used viscoelastic surfactant that forms a wormlike micellar gel at room temperature and a viscoelastic solution at higher temperatures. We probe both states by conducting linear and nonlinear rheological tests and analyze their behaviour under the framework of micellar rheology. Our study outlines departure from behaviour exhibited by more conventional micellar systems and uncovers interesting dynamics like shear-induced fracture and possible shear-banding in these materials. In doing so we provide a detailed understanding of a novel class of wormlike micellar solutions, Comment: Corrected micellar length estimates and energetic calculations using formulae in Ref 62. Added explanatory text, Added References, Corrected typos

We derive a general formula for the inertialess dynamics of active particles in linear viscoelastic fluids by means of a modified reciprocal theorem. We then demonstrate that force-free active particles in Maxwell-like linear viscoelastic fluids with no retardation have exactly the same dynamics as in Newtonian fluids. In contrast, active particles in Jeffreys-like fluids with retardation can display markedly different dynamics, including net motion under reciprocal actuation thereby breaking the scallop theorem., Comment: Error in equation (17). Missing term in time derivative

We investigate the dynamics of a prolate spheroid in a shear flow of a shear-thinning Carreau fluid. The motion of a prolate particle is developed analytically for asymptotically weak shear thinning and then integrated numerically. We find that shear-thinning rheology does not lift the degeneracy of Jeffery orbits observed in Newtonian fluids but the instantaneous rate of rotation and trajectories of the orbits are modified. Qualitatively, shear thinning has a similar effect to elongating the particle in a Newtonian fluid. The period of rotation increases as the particle slows down more when aligned with the flow due to a reduction of shear stresses. Unlike Jeffery orbits in Newtonian fluids, in shear-thinning fluids the period of orbits depends on the specific trajectory (or initial orientation of the particle)., Comment: 16 pages, 6 figures

Microswimmers in nature often experience spatial gradients of viscosity. In this work we develop theoretical results for the dynamics of active particles, biological or otherwise, swimming through viscosity gradients. We model the active particles using the squirmer model, and show how viscosity gradients lead to viscotaxis for squirmers, and how the effects of viscosity gradients depend on the swimming gait of the microswimmers. We also show how such gradients in viscosity can be used to control active particles and suggest a mechanism to sort them based on their swimming style., Comment: 5 pages, 2 figures

Many microorganisms swim in fluids with complex rheological properties. Although much is now understood about motion of these swimmers in Newtonian fluids, the understanding is still developing in non-Newtonian fluids --- this understanding is crucial for various biomimetic and biomedical applications. Here we study a common model for microswimmers, the squirmer model, in two common viscoelastic fluid models, the Giesekus fluid model and fluids of differential type (grade three), at zero Reynolds number. Through this article we address a recent commentary that discussed suitable values of parameters in these model and pointed at higher order viscoelastic effects on the squirming motion.

We examine swimmers comprising of two rigid spheres which oscillate periodically along their axis of symmetry, considering both when the oscillation is in phase and anti-phase, and study the effects of fluid viscoelasticity on their net motion. These swimmers both display reciprocal motion in a Newtonian fluid and hence no net swimming is achieved over one cycle. Conversely, we find that when the two spheres are of different sizes, the effect of viscoelasticity acts to propel the swimmers forward in the direction of the smaller sphere. Finally, we compare the motion of rigid spheres oscillating in viscoelastic fluids with elastic spheres in Newtonian fluids where we find similar results., Comment: 3 figures

A wetting liquid is driven through a thin gap due to surface tension and when the gap boundaries are elastic, the liquid deforms the gap as it rises. But when the fluid boundaries are also permeable (or poroelastic), the liquid can permeate the boundaries as the fluid rises and change their properties, for example by swelling and softening, thereby altering the dynamics of the rise. In this paper, we study the dynamics of capillary rise between two poroelastic sheets to understand the effects of boundary permeability and softening. We find that if the bending rigidity of sheets is reduced, due to liquid permeation, the sheets coalesce faster compared to the case of impermeable sheets. We show that as a direct consequence of this faster coalescence, the volume of fluid captured between the sheets can be significantly lower., Comment: 10 pages, 5 figures

Active particles moving through fluids generate disturbance flows due to their activity. For simplicity, the induced flow field is often modeled by the leading terms in a far-field approximation of the Stokes equations, whose coefficients are the force, torque and stresslet (zeroth and first-order force moments) of the active particle. This level of approximation is quite useful, but may also fail to predict more complex behaviors that are observed experimentally. In this study, to provide a better approximation, we evaluate the contribution of the second-order force moments to the flow field and, by reciprocal theorem, present explicit formulas for the stresslet dipole, rotlet dipole and potential dipole for an arbitrarily-shaped active particle. As examples of this method, we derive modified Fax\'en laws for active spherical particles and resolve higher-order moments for active rod-like particles., Comment: 15 pages

Particle motion in non-Newtonian fluids can be markedly different than in Newtonian fluids. Here we look at the change in dynamics for a few problems involving rigid spherical particles in shear-thinning fluids in the absence of inertia. We give analytical formulas for sedimenting spheres, obtained by means of the reciprocal theorem, and demonstrate quantitively differences in comparison to a Newtonian fluid. We also calculate the first correction to the suspension viscosity, the Einstein viscosity, for a dilute suspension of spheres in a weakly shear-thinning fluid., Comment: 12 pages, 1 figure

In this work, we numerically investigate the dynamics of a self-propelling autophoretic Janus particle in a weakly viscoelastic fluid. The self-propulsion is achieved by an asymmetry in the properties of the surface of the Janus particle that drives a surface slip velocity and bulk flow. Here we investigate the effect of viscoelasticity on this advection-diffusion problem over a range of P\'eclet and Damk\"ohler numbers. Particles are found to swim faster, or slower, in viscoelastic fluids, and we show how reaction and diffusion rates affect the viscoelastic stresses that lead to changes in propulsion., Comment: 10 pages, 9 figures

A generalized reciprocal theorem is formulated for the motion and hydrodynamic force moments of an active particle in an arbitrary background flow of a (weakly nonlinear) complex fluid. This formalism includes as special cases a number of previous calculations of the motion of both passive and active particles in Newtonian and non-Newtonian fluids., Comment: 6 pages

Biological locomotion in nature is often achieved by the interaction between a flexible body and its surrounding medium. The interaction of a flexible body with granular media is less understood compared with viscous fluids partially due to its complex rheological properties. In this work we explore the effect of flexibility on granular propulsion by considering a simple mechanical model where a rigid rod is connected to a torsional spring that is under a displacement actuation using a granular resistive force theory. Through a combined numerical and asymptotic investigation, we characterize the propulsive dynamics of such a flexible flapper in relation to the actuation amplitude and spring stiffness and compare these dynamics with those observed in a viscous fluid. In addition, we demonstrate that the maximum possible propulsive force can be obtained in the steady propulsion limit with a finite spring stiffness and large actuation amplitude. These results may apply to the development of synthetic locomotive systems that exploit flexibility to move through complex terrestrial media., Comment: 9 pages, 5 figures

At low Reynolds numbers the locomotive capability of a body can be dramatically hindered by the absence of inertia. In this work, we show how propulsive performance in this regime can be significantly enhanced by employing spatially varying flexibility. As a prototypical example, we consider the propulsive thrust generated by a filament periodically driven at one end. The rigid case leads to zero propulsion, as so constrained by Purcell's scallop theorem, while for uniform filaments there exists a bending stiffness maximizing the propulsive force at a given frequency; here we demonstrate explicitly how considerable further improvement (greater than 37% enhancement compared with the optimal uniform stiffness) can be achieved by simply varying the stiffness along the filament. The optimal flexibility distribution is strongly configuration-dependent: while increasing the flexibility towards the tail-end enhances the propulsion of a clamped filament, for a hinged filament decreasing the flexibility towards the tail-end is instead favorable. The results reveal new design principles for maximizing propulsion at low Reynolds numbers, potentially useful for developing synthetic micro-swimmers requiring large propulsive force for various biomedical applications., Comment: 5 figures

Swimming at low Reynolds number in Newtonian fluids is only possible through non-reciprocal body deformations due to the kinematic reversibility of the Stokes equations. We consider here a model swimmer consisting of two linked spheres, wherein one sphere is rigid and the other an incompressible neo-Hookean solid. The two spheres are connected by a rod which changes its length periodically. We show that the deformations of the body are non-reciprocal despite the reversible actuation and hence, the elastic two-sphere swimmer propels forward. Our results indicate that even weak elastic deformations of a body can affect locomotion and may be exploited in designing artificial microswimmers., Comment: 9 pages, 2 figures

In this work, we study active particles with prescribed surface velocities in non-Newtonian fluids. We employ the reciprocal theorem to obtain the velocity of an active spherical particle with an arbitrary axisymmetric slip-velocity in an otherwise quiescent second-order fluid. We then determine how the motion of a diffusiophoretic Janus particle is affected by complex fluid rheology, namely viscoelasticity and shear-thinning viscosity, compared to a Newtonian fluid, assuming a fixed slip-velocity. We find that a Janus particle may go faster or slower in a viscoelastic fluid, but is always slower in a shear-thinning fluid as compared to a Newtonian fluid., Comment: 9 pages, 3 figures

In this paper, we give formulas for the swimming of simplified two-dimensional bodies in complex fluids using the reciprocal theorem. By way of these formulas we calculate the swimming velocity due to small-amplitude deformations on the simplest of these bodies, a two-dimensional sheet, to explore general conditions on the swimming gait under which the sheet may move faster, or slower, in a viscoelastic fluid compared to a Newtonian fluid. We show that in general, for small amplitude deformations, a speed increase can only be realized by multiple deformation modes in contrast to slip flows. Additionally, we show that a change in swimming speed is directly due to a change in thrust generated by the swimmer.

Microorganisms develop coordinated beating patterns on surfaces lined with cilia known as metachronal waves. For a chain of cilia attached to a flat ciliate, it has been shown that hydrodynamic interactions alone can lead the system to synchronize. However, several microorganisms possess a curve shaped ciliate body and so to understand the effect of this geometry on the formation of metachronal waves, we evaluate the hydrodynamic interactions of cilia near a large spherical body. Using a minimal model, we show that for a chain of cilia around the sphere, the natural periodicity in the geometry leads the system to synchronize. We also report an emergent wave-like behavior when an asymmetry is introduced to the system.

Undulatory locomotion is ubiquitous in nature and observed in different media, from the swimming of flagellated microorganisms in biological fluids, to the slithering of snakes on land, or the locomotion of sandfish lizards in sand. Despite the similarity in the undulating pattern, the swimming characteristics depend on the rheological properties of different media. Analysis of locomotion in granular materials is relatively less developed compared with fluids partially due to a lack of validated force models but recently a resistive force theory in granular media has been proposed and shown useful in studying the locomotion of a sand-swimming lizard. Here we employ the proposed model to investigate the swimming characteristics of a slender filament, of both finite and infinite length, undulating in a granular medium and compare the results with swimming in viscous fluids. In particular, we characterize the effects of drifting and pitching in terms of propulsion speed and efficiency for a finite sinusoidal swimmer. We also find that, similar to Lighthill's results using resistive force theory in viscous fluids, the sawtooth swimmer is the optimal waveform for propulsion speed at a given power consumption in granular media. The results complement our understanding of undulatory locomotion and provide insights into the effective design of locomotive systems in granular media.

Many microorganisms find themselves immersed in fluids displaying non-Newtonian rheological properties such as viscoelasticity and shear-thinning viscosity. The effects of viscoelasticity on swimming at low Reynolds numbers have already received considerable attention, but much less is known about swimming in shear-thinning fluids. A general understanding of the fundamental question of how shear-thinning rheology influences swimming still remains elusive. To probe this question further, we study a spherical squirmer in a shear-thinning fluid using a combination of asymptotic analysis and numerical simulations. Shear-thinning rheology is found to affect a squirming swimmer in nontrivial and surprising ways; we predict and show instances of both faster and slower swimming depending on the surface actuation of the squirmer. We also illustrate that while a drag and thrust decomposition can provide insights into swimming in Newtonian fluids, extending this intuition to problems in complex media can prove problematic.

The use of the reciprocal theorem has been shown to be a powerful tool to obtain the swimming velocity of bodies at low Reynolds number. The use of this method for lower-dimensional swimmers, such as cylinders and sheets, is more problematic because of the undefined or ill-posed resistance problems that arise in the rigid-body translation of these shapes. Here we show that this issue can be simply circumvented and give concise formulas obtained via the reciprocal theorem for the self-propelled motion of deforming two-dimensional bodies. We also discuss the connection between these formulae and Fax\'en's laws.

In G.I. Taylor's historic paper on swimming microorganisms, a two dimensional sheet was proposed as a model for flagellated cells passing traveling waves as a means of locomotion. Using a perturbation series, Taylor computed swimming speeds up to fourth order in amplitude. Here we systematize that expansion so that it can be carried out formally to arbitrarily high order. The resultant series diverges for an order one value of the wave amplitude, but may be transformed into series with much improved convergence properties and which yield results comparing favorably to those obtained numerically via a boundary integral method for moderate and large values of the wave amplitudes.

Motivated by recent experiments, we consider theoretically the compression of droplets pinned at the bottom on a surface of finite area. We show that if the droplet is sufficiently compressed at the top by a surface, it will always develop a shape instability at a critical compression. When the top surface is flat, the shape instability occurs precisely when the apparent contact angle of the droplet at the pinned surface is pi, regardless of the contact angle of the upper surface, reminiscent of past work on liquid bridges and sessile droplets as first observed by Plateau. After the critical compression, the droplet transitions from a symmetric to an asymmetric shape. The force required to deform the droplet peaks at the critical point then progressively decreases indicative of catastrophic buckling. We characterize the transition in droplet shape using illustrative examples in two dimensions followed by perturbative analysis as well as numerical simulation in three dimensions. When the upper surface is not flat, the simple apparent contact angle criterion no longer holds, and a detailed stability analysis is carried out to predict the critical compression., Comment: 11 pages, 8 figures

When swimming in close proximity, some microorganisms such as spermatozoa synchronize their flagella. Previous work on swimming sheets showed that such synchronization requires a geometrical asymmetry in the flagellar waveforms. Here we inquire about a physical mechanism responsible for such symmetry-breaking in nature. Using a two-dimensional model, we demonstrate that flexible sheets with symmetric internal forcing, deform when interacting with each other via a thin fluid layer in such a way as to systematically break the overall waveform symmetry, thereby always evolving to an in-phase conformation where energy dissipation is minimized. This dynamics is shown to be mathematically equivalent to that obtained for prescribed waveforms in viscoelastic fluids, emphasizing the crucial role of elasticity in symmetry-breaking and synchronization., Comment: 8 pages, 4 figures

Spermatozoa flagella are known to synchronize when swimming in close proximity. We use a model consisting of two-dimensional sheets propagating transverse waves of displacement to demonstrate that fluid forces lead to such synchronization passively. Using two distinct asymptotic descriptions (small amplitude and long wavelength), we derive the synchronizing dynamics analytically for ar- bitrarily shaped waveforms in Newtonian fluids, and show that phase locking will always occur for sufficiently asymmetric shapes. We characterize the effect of the geometry of the waveforms and the separation between the swimmers on the synchronizing dynamics, the final stable conformations, and the energy dissipated by the cells. For two closely-swimming cells, synchronization always oc- curs at the in-phase or opposite-phase conformation, depending solely on the geometry of the cells. In contrast, the work done by the swimmers is always minimized at the in-phase conformation. As the swimmers get further apart, additional fixed points arise at intermediate values of the relative phase. In addition, computations for large-amplitude waves using the boundary integral method reveal that the two asymptotic limits capture all the relevant physics of the problem. Our results provide a theoretical framework to address other hydrodynamic interactions phenomena relevant to populations of self-propelled organisms., Comment: 29 pages, 12 figures

Experimental studies have demonstrated that spermatozoa synchronize their flagella when swimming in close proximity. In a Newtonian fluid, it was shown theoretically that such synchronization arises passively due to hydrodynamic forces between the two swimmers if their waveforms exhibit a front-back geometrical asymmetry. Motivated by the fact that most biological fluids possess a polymeric microstructure, we address here synchronization in a viscoelastic fluid analytically. Using a two-dimensional infinite sheet model we show that the presence of polymeric stresses removes the geometrical asymmetry constraint, and therefore even symmetric swimmers synchronize. Such synchronization occurs on asymptotically faster time scales than in a Newtonian fluid, and the swimmers are seen to be driven into a stable in-phase conformation minimizing the energy dissipated in the surrounding fluid.

Some microorganisms, such as spermatozoa, synchronize their flagella when swimming in close proximity. Using a simplified model (two infinite, parallel, two-dimensional waving sheets), we show that phase-locking arises from hydrodynamics forces alone, and has its origin in the front-back asymmetry of the geometry of their flagellar waveform. The time-evolution of the phase difference between co-swimming cells depends only on the nature of this geometrical asymmetry, and microorganisms can phase-lock into conformations which minimize or maximize energy dissipation.