In this paper we systematically derive a fourth-order continuum theory capable of reproducing mesoscale turbulence in a three-dimensional suspension of microswimmers. We start from overdamped Langevin equations for a generic microscopic model (pushers or pullers), which include hydrodynamic interactions on both, small length scales (polar alignment of neighboring swimmers) and large length scales, where the solvent flow interacts with the order parameter field. The flow field is determined via the Stokes equation supplemented by an ansatz for the stress tensor. In addition to hydrodynamic interactions, we allow for nematic pair interactions stemming from excluded-volume effects. The results here substantially extend and generalize earlier findings [Phys. Rev. E 94, 020601(R) (2016)], in which we derived a two-dimensional hydrodynamic theory. From the corresponding mean-field Fokker-Planck equation combined with a self-consistent closure scheme, we derive nonlinear field equations for the polar and the nematic order parameter, involving gradient terms of up to fourth order. We find that the effective microswimmer dynamics depends on the coupling between solvent flow and orientational order. For very weak coupling corresponding to a high viscosity of the suspension, the dynamics of mesoscale turbulence can be described by a simplified model containing only an effective microswimmer velocity.
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…
Microswimmer suspensions self-organize into complex spatio-temporal flow patterns, including vortex lattices and mesoscale turbulence. Here we explore the consequences for the motion of passive tracers, based on a continuum model for the microswimmer velocity field. We observe two qualitatively different regimes distinguished via the dimensionless Kubo number $K$. At advection strengths right above the transition to turbulence, the flow field evolves very slowly ($K \gg 1$) and the spatial vortex structures lead to dominant trapping effects. In contrast, deep in the turbulent state, much faste…
Tracer particles immersed in suspensions of biological microswimmers such as E. coli or C. reinhardtii display phenomena unseen in conventional equilibrium systems, including strongly enhanced diffusivity relative to the Brownian value and non-Gaussian displacement statistics. In dilute, 3-dimensional suspensions, these phenomena have typically been explained by the hydrodynamic advection of point tracers by isolated microswimmers, while, at higher concentrations, correlations between pusher microswimmers such as E. coli can increase the effective diffusivity even further. Anisotropic tracers …
The measurement of a quantitative and macroscopic parameter to estimate the global motility of a large population of swimming biological cells is a challenge Experiments on the rheology of active suspensions have been performed. Effective viscosity of sheared suspensions of live unicellular motile micro-algae (\textit{Chlamydomonas Reinhardtii}) is far greater than for suspensions containing the same volume fraction of dead cells and suspensions show shear thinning behaviour. We relate these macroscopic measurements to the orientation of individual swimming cells under flow and discuss our res…
We investigate coarse-grained models of suspended self-thermophoretic microswimmers. Upon heating, the Janus spheres, with hemispheres made of different materials, induce a heterogeneous local solvent temperature that causes the self-phoretic particle propulsion. Starting from atomistic molecular dynamics simulations, we verify the coarse-grained description of the fluid in terms of a local molecular temperature field, and its role for the particle's thermophoretic self-propulsion and hot Brownian motion. The latter is governed by effective nonequilibrium temperatures, which are measured from …
This paper studies the influence of orienting external fields on pattern formation, particularly mesoscale turbulence, in microswimmer suspensions. To this end, we apply a hydrodynamic theory that can be derived from a microscopic microswimmer model [Phys. Rev. E 97, 022613 (2018)]. The theory combines a dynamic equation for the polar order parameter with a modified Stokes equation for the solvent flow. Here, we extend the model by including an external field that exerts an aligning torque on the swimmers (mimicking the situation in chemo-, photo-, magneto- or gravitaxis). Compared to the fiel…
A universal characteristic of mesoscale turbulence in active suspensions is the emergence of a typical vortex length scale, distinctly different from the scale-invariance of turbulent high-Reynolds number flows. Collective length-scale selection has been observed in bacterial fluids, endothelial tissue and active colloides, yet the physical origins of this phenomenon remain elusive. Here, we systematically derive an effective fourth-order field theory from a generic microscopic model that allows us to predict the typical vortex size in microswimmer suspensions. Building on a self-consistent cl…
We explore in this paper the phenomenon of photofocusing: a coupling between flow vorticity and biased swimming of microalgae toward a light source that produces a focusing of the microswimmer suspension. We combine experiments that investigate the stationary state of this phenomenon as well as the transition regime with analytical and numerical modeling. We show that the experimentally observed scalings on the width of the focalized region and the establishment length as a function of the flow velocity are well described by a simple theoretical model.
Hydrodynamical interactions of active micro-particles are pervasive in our planet's fluid environments. Hence, understanding the interactions of these self-propelled particles is essential for science and engineering. In this paper the suspensions of active swimmer-rotor particles have been mathematically modeled by extending a previously developed stochastic kinetic theory to analyze heterogeneous collections of microswimmers and microrotors with multiple spin velocity populations. The paper uses this modeling approach to derive insights on large scale properties such as suspension instabilit…
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…
Active microswimmers are known to affect the macroscopic viscosity of suspensions in a more complex manner than passive particles. For puller-like microswimmers an increase in the viscosity has been observed. It has been suggested that the persistence of the orientation of the microswimmers hinders the rotation that is normally caused by the vorticity. It was previously shown that some sorts of algaes are bottom-heavy swimmers, i.e. their centre of mass is not located in the centre of the body. In this way, the algae affects the vorticity of the flow when it is perpendicular oriented to the ax…
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…
Neural-Network-Assisted Boltzmann Approach for Dilute Microswimmer Suspensions
Institute of Industrial Science, The University of Tokyo（东京大学工业科学研究所）；Division of Chemical Engineering, Department of Materials Engineering Science, Graduate School of Engineering Science, The University of Osaka（大阪大学工程科学研究科材料工程科学系化学工程学部门）
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…
Some micro-algae are sensitive to light intensity gradients. This property is known as phototaxis: the algae swim toward a light source (positive phototaxis). We use this property to control the motion of micro-algae within a Poiseuille flow using light. The combination of flow vorticity and phototaxis results in a concentration of algae around the center of the flow. Intermittent light exposure allows analysis of the dynamics of this phenomenon and its reversibility. With this phenomenon, we hope to pave the way toward new algae concentration techniques (a bottleneck challenge in hydrogen alg…
Mesoscale simulations of diffusion and sedimentation in shape-anisotropic nanoparticle suspensions
Johannes Gutenberg University Mainz（美因茨约翰内斯·古腾堡大学）；Auburn University（奥本大学）；Leibniz-Institut für Polymerforschung Dresden e.V.（德累斯顿莱布尼茨聚合物研究所）；Technische Universität Dresden（德累斯顿工业大学）
We analyze the dynamics of a microswimmer in pressure-driven Poiseuille flow, where fluid inertia is small but non-negligible. Using perturbation theory and the reciprocal theorem, we show that in addition to the classical inertial lift of passive particles, the active nature generates a `swimming lift', which we evaluate for neutral and pusher/puller-type swimmers. Accounting for fluid inertia engenders a rich spectrum of novel complex dynamics including bistable states, where tumbling coexists with stable centerline swimming or swinging. The dynamics is sensitive to the swimmer's hydrodynami…
Dynamical density functional theory (DDFT) has been successfully derived and applied to describe on the one hand passive colloidal suspensions, including hydrodynamic interactions between individual particles. On the other hand, active "dry" crowds of self-propelled particles have been characterized using DDFT. Here we go one essential step further and combine these two approaches. We establish a DDFT for active microswimmer suspensions. For this purpose, simple minimal model microswimmers are introduced. These microswimmers self-propel by setting the surrounding fluid into motion. They hydrod…
We developed a mechanical model of spermatozoal swimming in bulk suspensions. We traced the spatiotemporal elastohydrodynamic interactions and found that spermatozoa engaged in self-organisation: flagellar undulatory motion generated a stable polar order. Conventional microswimmer theories use time-averaged stress fields to predict that the cellular structure is unstable. However, our present results reveal a novel self-organisation mechanism attributable to unsteady hydrodynamic interactions. We thus contribute to the theory of active soft matter dynamics.
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…
Microswimmers are active particles of microscopic size that self-propel by setting the surrounding fluid into motion. According to the kind of far-field fluid flow that they induce, they are classified into pushers and pullers. Many studies have explored similarities and differences between suspensions of either pushers or pullers, but the behavior of mixtures of the two is still to be investigated. Here, we rely on a minimal discrete microswimmer model, particle-resolved, including hydrodynamic interactions, to examine the orientational ordering in such binary pusher-puller mixtures. In agree…
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. …
We analyze a minimal model for a rigid spherical microswimmer and explore the consequences of its extended surface on the interplay between its self-propulsion and flow properties. The model is the first order representation of microswimmers, such as bacteria and algae, with rigid bodies and flexible propelling appendages. The flow field of such a microswimmer at finite distances significantly differs from that of a point-force (Stokeslet) dipole. For a suspension of microswimmers, we derive the grand mobility matrix that connects the motion of an individual swimmer to the active and passive f…
The nature of blood as a suspension of red blood cells makes computational hemodynamics a demanding task. Our coarse-grained blood model, which builds on a lattice Boltzmann method for soft particle suspensions, enables the study of the collective behavior of the order of 10^6 cells in suspension. After demonstrating the viscosity measurement in Kolmogorov flow, we focus on the statistical analysis of the cell orientation and rotation in Couette flow. We quantify the average inclination with respect to the flow and the nematic order as a function of shear rate and hematocrit. We further record…
We study synchronization in bulk suspensions of spherical microswimmers with chiral trajectories using large scale numerics. The model is generic. It corresponds to the lowest order solution of a general model for self-propulsion at low Reynolds numbers, consisting of a nonaxisymmetric rotating source dipole. We show that both purely circular and helical swimmers can spontaneously synchronize their rotation. The synchronized state corresponds to velocity alignment with high orientational order in both the polar and azimuthal directions. Finally, we consider a racemic mixture of helical swimmer…
Active matter exhibits various forms of non-equilibrium states in the absence of external forcing, including macroscopic steady-state currents. Such states are often too complex to be modelled from first principles and our understanding of their physics relies heavily on minimal models. These have mostly been studied in the case of "dry" active matter, where particle dynamics are dominated by friction with their surroundings. Significantly less is known about systems with long-range hydrodynamic interactions that belong to "wet" active matter. Dilute suspensions of motile bacteria, modelled as…
We study different types of microswimmers moving in channels with varying cross section and thereby interacting hydrodynamically with the channel walls. Starting from the Smoluchowski equation for a dilute suspension, for which interactions among swimmers can be neglected, we derive analytic expressions for the lateral probability distribution between plane channel walls. For weakly corrugated channels we extend the Fick--Jacobs approach to microswimmers and thereby derive an effective equation for the probability distribution along the channel axis. Two regimes arise dominated either by entro…
Geometric confinements are frequently encountered in the biological world and strongly affect the stability, topology, and transport properties of active suspensions in viscous flow. Based on a far-field analytical model, the low-Reynolds-number locomotion of a self-propelled microswimmer moving inside a clean viscous drop or a drop covered with a homogeneously distributed surfactant, is theoretically examined. The interfacial viscous stresses induced by the surfactant are described by the well-established Boussinesq-Scriven constitutive rheological model. Moreover, the active agent is represe…
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…
The collective motion of microswimmers in suspensions induce patterns of vortices on scales that are much larger than the characteristic size of a microswimmer, attaining a state called bacterial turbulence. Hydrodynamic turbulence acts on even larger scales and is dominated by inertial transport of energy. Using an established modification of the Navier-Stokes equation that accounts for the small scale forcing of hydrodynamic flow by microswimmers, we study the properties of a dense supensions of microswimmers in two dimensions, where the conservation of enstrophy can drive an inverse cascade…
Microswimmers such as E. Coli bacteria accumulate and exhibit an intriguing dynamics near walls, governed by hydrodynamic and steric interactions. Insight into the underlying mechanisms and predominant interactions demand a detailed characterization of the entrapment process. We employ a mesoscale hydrodynamics simulation approach to study entrapment of a E. coli-type cell at a no-slip wall. The cell is modeled by a spherocylindrical body with several explicit helical flagella. Three stages of the entrapment process can be distinguished: the approaching regime, where a cell swims toward the wa…
The dynamics and rheology of suspensions of fluid vesicles or red blood cells is investigated by a combination of molecular dynamics and mesoscale hydrodynamics simulations in two dimensions. The vesicle suspension is confined between two no-slip walls, which are driven externally to generate a shear flow with shear rate $\dotγ$. The flow behavior is studied as a function of $\dotγ$, the volume fraction of vesicles, and the viscosity contrast between inside and outside fluids. Results are obtained for the encounter and interactions of two vesicles, the intrinsic viscosity of the suspension, an…
Soft particles at fluid interfaces play an important role in many aspects of our daily life, such as the food industry, paints and coatings, and medical applications. Analytical methods are not capable of describing the emergent effects of the complex dynamics of suspensions of many soft particles, whereas experiments typically either only capture bulk properties or require invasive methods. Computational methods are therefore a great tool to complement experimental work. However, an efficient and versatile numerical method is needed to model dense suspensions of many soft particles. In this a…
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…
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…
Self-organised structures in mixed active-passive suspensions due to hydrodynamic interactions
Capital Fund Management S.A.（资本基金管理公司）；Institut Pasteur, Université Paris Cité, CNRS UMR3738（巴斯德研究所，巴黎西岱大学，法国国家科学研究中心）；Laboratoire de Physique de l’École normale supérieure, ENS, Université PSL, CNRS, Sorbonne Université, Université Paris Cité（巴黎高等师范学院物理实验室，巴黎文理研究大学，法国国家科学研究中心，索邦大学，巴黎西岱大学）；Department of Biomedical Engineering, Tohoku University（东北大学生物医学工程系）
The collective effects of microswimmers in active suspensions result in active turbulence, a spatiotemporally chaotic dynamics at mesoscale, which is characterized by the presence of vortices and jets at scales much larger than the characteristic size of the individual active constituents. To describe this dynamics, Navier-Stokes-based one-fluid models driven by small-scale forces have been proposed. Here, we provide a justification of such models for the case of dense suspensions in two dimensions (2d). We subsequently carry out an in-depth numerical study of the properties of one-fluid model…
Hydrodynamic interactions in systems comprised of self-propelled particles, such as swimming microorganisms, and passive tracers have a significant impact on the tracer dynamics compared to the equivalent "dry" sample. However, such interactions are often difficult to take into account in simulations due to their computational cost. Here, we perform a systematic investigation of swimmer-tracer interaction using an efficient force/counter-force based lattice-Boltzmann (LB) algorithm [J. de Graaf~\textit{et al.}, J. Chem. Phys.~\textbf{144}, 134106 (2016)] in order to validate its ability to cap…
We use boundary element simulations to study the interaction of model microswimmers with a neutrally buoyant spherical particle. The ratio of the size of the particle to that of the swimmer is varied from $R^\mathrm{P} / R^\mathrm{S} \ll 1$, corresponding to swimmer--tracer scattering, to $R^\mathrm{P} / R^\mathrm{S} \gg 1$, approximately equivalent to the swimmer interacting with a fixed, flat surface. We find that details of the swimmer and particle trajectories vary for different swimmers. However, the overall characteristics of the scattering event fall into two regimes, depending on the r…
