Ensembles of biological and artificial microswimmers produce long-range velocity fields with strong nonequilibrium fluctuations, which result in dramatic increase diffusivity of embedded particles (tracers). While such enhanced diffusivity may point to enhanced mixing of the fluid, a rigorous quantification of the mixing efficiency requires analysis of pair dispersion of tracers, rather than simple one-particle diffusivity. Here, we calculate analytically the scale-dependent coefficient of relative diffusivity of passive tracers embedded in a dilute suspension of run-and-tumble microswimmers. Although each tracer is subject to strong fluctuations resulting in large absolute diffusivity, the small-scale relative dispersion is suppressed due to the correlations in fluid velocity which are relevant when the inter-tracers separation is below the persistence length of the swimmers motion. Our results suggest that the reorientation of swimming direction plays an important role in biological mixing and should be accounted in design of potential active matter devices capable of effective fluid mixing at microscale.
A dilute non-interacting suspension of micro-swimmers exhibits a finite velocity variance and short-ranged correlations that decay over a swimmer length. For a suspension of interacting straight swimmers, however, pair-interactions leads to a non-decaying velocity covariance, and a variance that diverges logarithmically with system size. The divergence is arrested on inclusion of orientation decorrelation mechanisms. Results for suspensions of run-and-tumble particles (RTPs) are presented, where the underlying straight-swimmer divergence leads to a broad cross-over between the ballistic and di…
We show that the Casimir effect can emerge in microswimmer suspensions. In principle, two effects conspire against the development of Casimir effects in swimmer suspensions. First, at low Reynolds number, the force on any closed volume vanishes, but here the relevant effect is the drag by the flow produced by the swimmers, which can be finite. Second, the fluid velocity and the pressure are linear on the swimmer force dipoles, and averaging over the swimmer orientations would lead to a vanishing effect. However, being the suspension a discrete system, the noise terms of the coarse grained equa…
Microorganisms can preferentially orient and move along gradients of a chemo-attractant (i.e., chemotax) while colonies of many microorganisms can collectively undergo complex dynamics in response to chemo-attractants that they themselves produce. For colonies or groups of micro-swimmers we investigate how an "auto-chemotactic" response that should lead to swimmer aggregation is affected by the non-trivial fluid flows that are generated by collective swimming. For this, we consider chemotaxis models based upon a hydrodynamic theory of motile suspensions that are fully coupled to chemo-attracta…
Ordered phases emerged in active suspensions of polar swimmers are under long-wavelength hydrodynamic mediated instabilities. In this letter, we show that chemical molecules dissolved in aqueous suspensions, as an unavoidable part of most wet active systems, can mediate long-range interactions and subsequently stabilize the ordered phases. Chemoattractant in living suspensions and dissolved molecules producing phoretic forces in synthesized Janus suspensions are reminiscent of such molecules. Communication between swimmers through the gradients of such chemicals generated by individual swimmer…
In addition to enabling movement towards environments with favourable living conditions, swimming by microorganisms has also been linked to enhanced mixing and improved nutrient uptake by their populations. Experimental studies have shown that Brownian tracer particles exhibit enhanced diffusion due to the swimmers, while theoretical models have linked this increase in diffusion to the flows generated by the swimming microorganisms, as well as collisions with the swimmers. In this study, we perform detailed simulations based on the force-coupling method and its recent extensions to the swimmin…
We study the effect of magnetic field on the rheology of magnetic micro-swimmers suspensions. We use a model of a dilute suspension under simple shear and subjected to a constant magnetic field. Particle shear stress is obtained for both pusher and puller types of micro-swimmers. In the limit of low shear rate, the rheology exhibits a constant shear stress, called actuated stress, which only depends on the swimming activity of the particles. This stress is induced by the magnetic field and can be positive (brake state) or negative (motor state). In the limit of low magnetic fields, a scaling r…
The agitation produced in a fluid by a suspension of micro-swimmers in the low Reynolds number limit is studied. In this limit, swimmers are modeled as force dipoles all with equal strength. The agitation is characterized by the active temperature defined, as in kinetic theory, as the mean square velocity, and by the equal-time spatial correlations. Considering the phase in which the swimmers are homogeneously and isotropically distributed in the fluid, it is shown that the active temperature and velocity correlations depend on a single scalar correlation function of the dipole-dipole correlat…
Several active organisms in nature tend to reside as a community in a viscous fluid medium. We analyze the variation of swimming characteristics of an active swimmer present in a dilute and disperse suspension, modeled as an effective Brinkman medium. This idealized representation of a collection of active swimmers allows one to distinguish the impact of the interior domain available to an individual swimmer as well as the contribution of its neighbors. Darcy's law along with the analytical solution enables the effective resistivity to be predicted as a function of the volume fraction which is…
The induced diffusion of tracers in a bacterial suspension is studied theoretically and experimentally at low bacterial concentrations. Considering the swimmer-tracer hydrodynamic interactions at low-Reynolds number and using a kinetic theory approach, it is shown that the induced diffusion coefficient is proportional to the swimmer concentration, their mean velocity and a coefficient $β$, as observed experimentally. The coefficient $β$ scales as the tracer-swimmer cross section times the mean square displacement produced by single scatterings. The displacements depend on the swimmer propulsio…
We use numerical simulations to probe the dynamics of concentrated suspensions of spherical microswimmers interacting hydrodynamically. Previous work in the dilute limit predicted orientational instabilities of aligned suspensions for both pusher and puller swimmers, which we confirm computationally. Unlike previous work, we show that isotropic suspensions of spherical swimmers are also always unstable. Both types of initial conditions develop long-time polar order, of a nature which depends on the hydrodynamic signature of the swimmer but very weakly on the volume fraction up to very high vol…
We present a comprehensive theory of the dynamics and fluctuations of a two-dimensional suspension of polar active particles in an incompressible fluid confined to a substrate. We show that, depending on the sign of a single parameter, a state with polar orientational order is anomalously stable (or anomalously unstable), with a nonzero relaxation (or growth) rate for angular fluctuations at zero wavenumber. This screening of the broken-symmetry mode in the stable state does lead to conventional rather than giant number fluctuations as argued by Bricard et al., Nature ${\bf 503}$, 95 (2013), b…
In this Letter, we study the collective behaviour of a large number of self-propelled microswimmers immersed in a fluid. Using unprecedently large-scale lattice Boltzmann simulations, we reproduce the transition to bacterial turbulence. We show that, even well below the transition, swimmers move in a correlated fashion that cannot be described by a mean-field approach. We develop a novel kinetic theory that captures these correlations and is non-perturbative in the swimmer density. To provide an experimentally accessible measure of correlations, we calculate the diffusivity of passive tracers …
Suspensions of rear- and front-actuated microswimmers immersed in a fluid, known respectively as ``pushers'' and ``pullers'', display qualitatively different collective behaviours: beyond a characteristic density, pusher suspensions exhibit a hydrodynamic instability leading to collective motion known as active turbulence, a phenomenon which is absent for pullers. In this Letter, we describe the collective dynamics of a binary pusher--puller mixture using kinetic theory and large-scale particle-resolved simulations. We derive and verify an instability criterion, showing that the critical densi…
We run numerical simulations of strongly confined suspensions of model spherical swimmers called "squirmers". Because of the confinement, the Stokeslet dipoles generated by the particles are quickly screened and the far-field flow is dominated by the source dipole for all the different kinds of squirmers. However, we show that the collective behavior of the suspension still depends on the self-propelling mechanism of the swimmers as polar states can only be observed for neutral squirmers. We demonstrate that the near-field hydrodynamic interactions play a crucial role in the alignment of the o…
Swimming microorganisms can influence the diffusion of passive particles. The effect of this swimmer-particle interaction depends on different properties, such as the hydrodynamic field of the swimmer and the relative sizes of microorganisms and particles. We investigated an enhancement of the diffusion of silica doublets in a suspension of microalgae Chlamydomonas reinhardtii in a flat capillary. Depending on the concentration of microswimmers, the translational and rotational diffusion constants increase by several orders of magnitude in the presence of the swimming algae. For low concentrat…
With a detail microscopic model for a self-propelled swimmer, we derive the rheological properties of a dilute suspension of such particles at small Peclet numbers. It is shown that, in addition to the Einstein's like contribution to the effective viscosity, that is proportional to the volume fraction of the swimmers, a contribution due to the activity of self-propelled particles influences the viscosity. As a result of the activity of swimmers, the effective viscosity would be a lower (higher) than the viscosity of the suspending medium when the particles are pushers (pullers). Such activity …
Micro-organisms usually can swim in their liquid environment by flagellar or ciliary beating. In this numerical work, we analyze the influence of flagellar beating on the orbits of a swimming cell in a shear flow. We also calculate the effect of the flagellar beating on the rheology of a dilute suspension of micro-swimmers. A three-dimensional model is proposed for Chlamydomonas Reinhardtii swimming with a breaststroke-like beating of two anterior flagella modeled by two counter-rotating fore beads. The active swimmer model reveals unusual angular orbits in a linear shear flow. Namely, the swi…
The majority of studies on self-propelled particles and microswimmers concentrates on objects that do not feature a deterministic bending of their trajectory. However, perfect axial symmetry is hardly found in reality, and shape-asymmetric active microswimmers tend to show a persistent curvature of their trajectories. Consequently, we here present a particle-scale statistical approach of circle-swimmer suspensions in terms of a dynamical density functional theory. It is based on a minimal microswimmer model and, particularly, includes hydrodynamic interactions between the swimmers. After deriv…
The presence of active forces in various biological and artificial systems may change how those systems behaves under forcing. We present a minimal model of a suspension of passive or active swimmers driven on the boundaries by time-dependent forcing. In particular, we consider a time-periodic drive from which we determine the linear response functions of the suspension. The meaning of these response functions are interpreted in terms of the storage and dissipation of energy through the particles within the system. We find that while a slowly driven active system responds in a way similar to a…
It is well known that suspensions of particles in a viscous fluid can affect the rheology significantly, producing a pronounced non-Newtonian response even in dilute suspension. However, it is unclear a priori which particle shapes lead to this behaviour. We present two simple symmetry conditions on the shape which are sufficient for a dilute suspension to be Newtonian for all strain sizes and one sufficient for Newtonian behavior for small strains. We also construct a class of shapes out of thin, rigid rods not found by the symmetry argument which share this property for small strains.
Using a fluid-particle dynamics approach, we numerically study the effects of hydrodynamic interactions on the collective dynamics of active suspensions within a simple model for bacterial motility: each microorganism is modeled as a stroke-averaged dumbbell swimmer with prescribed dipolar force pairs. Using both simulations and qualitative arguments, we show that, when the separation between swimmers is comparable to their size, the swimmers' motions are strongly affected by activity-induced hydrodynamic forces. To further understand these effects, we investigate semidilute suspensions of swi…
Sedimentation of a non-Brownian suspension of hard particles is studied. It is shown that in the low concentration limit a two-particle distribution function ensuring finite particle correlation length can be found and explicitly calculated. The sedimentation coefficient is computed. Results are compared with experiment.
Collective behaviour in suspensions of microswimmers is often dominated by the impact of long-ranged hydrodynamic interactions. These phenomena include active turbulence, where suspensions of pusher bacteria at sufficient densities exhibit large-scale, chaotic flows. To study this collective phenomenon, we use large-scale (up to $N=3\times 10^6$) particle-resolved lattice Boltzmann simulations of model microswimmers described by extended stresslets. Such system sizes enable us to obtain quantitative information about both the transition to active turbulence and characteristic features of the t…
Kinetic theory for dilute inertial suspension having soft-core potential is theoretically investigated. From the analysis of the scattering process, the expression of the scattering angle is analytically obtained. We derive the flow curve between the viscosity and the shear rate, which shows two-step discontinuous shear thickening when we change the softness of the particles. The molecular dynamics simulation shows that our theoretical results are consistent with the numerical ones.
Previous particle-based computer simulations have revealed a significantly more pronounced tendency of spontaneous global polar ordering in puller (contractile) microswimmer suspensions than in pusher (extensile) suspensions. We here evaluate a microscopic statistical theory to investigate the emergence of such order through a linear instability of the disordered state. For this purpose, input concerning the orientation-dependent pair-distribution function is needed, and we discuss corresponding approaches, particularly a heuristic variant of the Percus test-particle method applied to active s…
We study how hydrodynamic interactions affect the collective behaviour of active particles suspended in a fluid at high concentrations, with particular attention to lubrication forces which appear when the particles are very close to one another. We compute exactly the limiting behaviour of the hydrodynamic interactions between two spherical (circular) active swimmers in very close proximity to one another in the general setting in both three and (two) dimensions. Combining this with far-field interactions, we develop a novel numerical scheme which allows us to study the collective behaviour o…
A kinetic theory for a dilute gas-solid suspension under a simple shear is developed. With the aid of the corresponding Boltzmann equation, it is found that the flow curve (stress-strain rate relation) has a S-shape as a crossover from the Newtonian to the Bagnoldian for a granular suspension or from the Newtonian to a fluid having a viscosity proportional to the square of the shear rate for a suspension consisting of elastic particles. The existence of the S-shape in the flow curve directly leads to a discontinuous shear thickening (DST). This DST corresponds to the discontinuous transition o…
Swimming microorganisms rely on effective mixing strategies to achieve efficient nutrient influx. Recent experiments, probing the mixing capability of unicellular biflagellates, revealed that passive tracer particles exhibit anomalous non-Gaussian diffusion when immersed in a dilute suspension of self-motile Chlamydomonas reinhardtii algae. Qualitatively, this observation can be explained by the fact that the algae induce a fluid flow that may occasionally accelerate the colloidal tracers to relatively large velocities. A satisfactory quantitative theory of enhanced mixing in dilute active sus…
We present a new development of the force-coupling method (FCM) to address the accurate simulation of a large number of interacting micro-swimmers. Our approach is based on the squirmer model, which we adapt to the FCM framework, resulting in a method that is suitable for simulating semi-dilute squirmer suspensions. Other effects, such as steric interactions, are considered with our model. We test our method by comparing the velocity field around a single squirmer and the pairwise interactions between two squirmers with exact solutions to the Stokes equations and results given by other numeric…
Microfluidic technologies are commonly used for the manipulation of red blood cell (RBC) suspensions and analyses of flow-mediated biomechanics. To enhance the performance of microfluidic devices, understanding the dynamics of the suspensions processed within is crucial. We report novel aspects of the spatio-temporal dynamics of RBC suspensions flowing through a typical microchannel at low Reynolds number. Through experiments with dilute RBC suspensions, we find an off-centre two-peak (OCTP) profile of cells contrary to the centralised distribution commonly reported for low-inertia flows. This…
We present a mesoscopic hydrodynamic description of the dynamics of colloidal suspensions. We consider the system as a gas of Brownian particles suspended in a Newtonian heat bath subjected to stationary non-equilibrium conditions imposed by a velocity field. Using results already obtained in previous studies in the field by means of a generalized Fokker-Planck equation, we obtain a set of coupled differential equations for the local diffusion current and the evolution of the total stress tensor. We find that the dynamic shear viscosity of the system contains contributions arising from the fin…
Similarly to their biological counterparts, suspensions of chemically-active autophoretic swimmers exhibit nontrivial dynamics involving self-organization processes as a result of inter-particle interactions. Using a kinetic model for a dilute suspension of autochemotactic Janus particles, we analyse the effect of a confined pressure-driven flow on these collective behaviours and the impact of chemotactic aggregation on the effective viscosity of the active fluid. Four dynamic regimes are identified when increasing the strength of the imposed pressure-driven flow, each associated with a differ…
In this work, we investigate the dynamics of the number density fluctuations of a dilute suspension of active particles in a linear viscoelastic fluid. We propose a model for the frequency-dependent diffusion coefficient of the active particles, which captures the effect of rotational diffusion on the persistence of their self-propelled motion and the viscoelasticity of the medium. Using fluctuating hydrodynamics, the linearized equations for the active suspension are derived, from which we calculate its dynamic structure factor and the corresponding intermediate scattering function. For a Max…
Populations of swimming microorganisms produce fluid motions that lead to dramatically enhanced diffusion of tracer particles. Using simulations of suspensions of swimming particles in a periodic domain, we capture this effect and show that it depends qualitatively on the mode of swimming: swimmers ``pushed'' from behind by their flagella show greater enhancement than swimmers that are ``pulled'' from the front. The difference is manifested by an increase, that only occurs for pushers, of the diffusivity of passive tracers and the velocity correlation length with the size of the periodic domai…
We investigate previously unclarified effects of fluid elasticity on shear-thickening in dilute suspensions in an Oldroyd-B viscoelastic fluid using a novel direct numerical simulation based on the smoothed profile method. Fluid elasticity is determined by the Weissenberg number Wi and by viscosity ratio $1-β=η_p/(η_s+η_p)$ which measures the coupling between the polymer stress and flow: $η_p$ and $η_s$ are the polymer and solvent viscosity, respectively.As $1-β$ increases, while the stresslet does not change significantly compared to that in the $β\to 1$ limit, the growth rate of the normaliz…
We consider the tracer diffusion $D_{rr}$ that arises from the run-and-tumble motion of low Reynolds number swimmers, such as bacteria. In unbounded dilute suspensions, where the dipole swimmers move in uncorrelated runs of length $λ$, an exact solution showed that $D_{rr}$ is independent of $λ$. Here we verify this result in numerical simulations for a particular model swimmer, the spherical squirmer. We also note that in confined microenvironments, such as microscopic droplets, microfluidic devices and bacterial microzones in marine ecosystems, the size of the system can be comparable to $λ$…
We show that the shear-induced solidification of dilute charge-stabilized (DLVO) colloids is due to the interplay between the shear-induced formation and breakage of large non-Brownian clusters. While their size is limited by breakage, their number density increases with the shearing-time. Upon flow cessation, the dense packing of clusters interconnects into a rigid state by means of grainy bonds, each involving a large number of primary colloidal bonds. The emerging picture of shear-driven solidification in dilute colloidal suspensions combines the gelation of Brownian systems with the jammin…
We investigate the effect of particle volume fraction on the efficiency of deterministic lateral displacement (DLD) devices. DLD is a popular passive sorting technique for microfluidic applications. Yet, it has been designed for treating dilute suspensions, and its efficiency for denser samples is not well known. We perform 3D simulations based on the immersed-boundary, lattice-Boltzmann and finite-element methods to model the flow of red blood cells (RBCs) in different DLD devices. We quantify the DLD efficiency in terms of appropriate "failure" probabilities and RBC counts in designated devi…
We study the diffusion of a Brownian probe particle of size $R$ in a dilute dispersion of active Brownian particles (ABPs) of size $a$, characteristic swim speed $U_0$, reorientation time $τ_R$, and mechanical energy $k_s T_s = ζ_a U_0^2 τ_R /6$, where $ζ_a$ is the Stokes drag coefficient of a swimmer. The probe has a thermal diffusivity $D_P = k_B T/ζ_P$, where $k_B T$ is the thermal energy of the solvent and $ζ_P$ is the Stokes drag coefficient for the probe. When the swimmers are inactive, collisions between the probe and the swimmers sterically hinder the probe's diffusive motion. In compe…
We investigate the steady-state extensional rheology of a dilute suspension of spherical particles in a dilute polymer solution. For a particle-free polymeric fluid, in addition to the solvent viscosity, the extensional viscosity due to the polymers, $μ^\text{poly}$, contributes to the total non-dimensionalized extensional viscosity $1+μ^\text{poly}$. When a small volume fraction, $ϕ$, of spheres is added to a polymeric fluid, the stress is altered by the Einstein viscosity of 2.5$ϕ$ and two additional stress contributions: the interaction stresslet and the particle-induced polymer stress (PIP…
We present new data for the electrical conductivity of foams in which the liquid fraction ranges from two to eighty percent. We compare with a comprehensive collection of prior data, and we model all results with simple empirical formulæ. We achieve a unified description that applies equally to dry foams and emulsions, where the droplets are highly compressed, as well as to dilute suspensions of spherical particles, where the particle separation is large. In the former limit, Lemlich's result is recovered; in the latter limit, Maxwell's result is recovered.
We propose a model for the nonequilibrium enhancement of colloidal self-diffusion in an externally imposed shear flow in charged systems. The enhancement of diffusion is calculated in terms of the electrostatic, two--body interactions between the particles in the shear flow. In the high shear rate, low volume fraction limit in which our model is valid, we compare these calculations to the experiments of Qiu et al. [Phys. Rev. Lett. 61, 2554 (1988)].
The rheological response under simple shear of an active suspension of Escherichia coli is determined in a large range of shear rates and concentrations. The effective viscosity and the time scales characterizing the bacterial organization under shear are obtained. In the dilute regime, we bring evidences for a low shear Newtonian plateau characterized by a shear viscosity decreasing with concentration. In the semi-dilute regime, for particularly active bacteria, the suspension display a "super-fluid" like transition where the viscous resistance to shear vanishes, thus showing that macroscopic…
Microswimmers often exhibit surprising patterns due to the nonequilibrium nature of their dynamics. Collectively, suspensions of microswimmers appear as a liquid whose properties set it apart from its passive counterpart. To understand the impact of hydrodynamic interactions on the basic statistical features of a microswimmer's liquid, we investigate its structure by means of the pair distribution function. We perform particle-based simulations of microswimmers that include steric effects, shape anisotropy, and hydrodynamic interactions. We find that hydrodynamic interactions considerably alte…
We present a novel approach for characterizing the properties and performance of active matter in dilute suspension as well as in crowded environments. We use Super-Heterodyne Laser-Doppler-Velocimetry (SH-LDV) to study large ensembles of catalytically active Janus particles moving under UV-illumination. SH-LDV facilitates a model-free determination of the swimming speed and direction, with excellent ensemble averaging. In addition we obtain in-formation on the distribution of the catalytic activity. Moreover, SH-LDV operates away from walls and permits a facile correction for multiple scatter…
We experimentally study density fluctuations and energy spectra of bulk \textit{E. coli} suspensions of different concentrations. Our results verify the predicted scaling law of giant number fluctuations in three-dimensional (3D) wet active fluids. We find that such a scaling behavior persists at small scales even in low concentration suspensions well below the transition concentration to active turbulence. Our experiments also support the prediction on the energy spectra of dilute pusher swimmers and illustrate the spectral properties of the active turbulence of dense bacterial suspensions in…
