Self-propelled swimmers such as bacteria agglomerate into clusters as a result of their persistent motion. In 1D, those clusters do not coalesce macroscopically and the stationary cluster size distribution (CSD) takes an exponential form. We develop a minimal lattice model for active particles in narrow channels to study how clustering is affected by the interplay between self-propulsion speed diversity and confinement. A mixture of run-and-tumble particles with a distribution of self-propulsion speeds is simulated in 1D. Particles can swap positions at rates proportional to their relative self-propulsion speed. Without swapping, we find that the average cluster size $L_\text{c}$ decreases with diversity and follows a non-arithmetic power mean of the single-component $L_\text{c}$'s, unlike the case of tumbling-rate diversity previously studied. Effectively, the mixture is thus equivalent to a system of identical particles whose self-propulsion speed is the harmonic mean self-propulsion speed of the mixture. With swapping, particles escape more quickly from clusters. As a consequence, $L_\text{c}$ decreases with swapping rates and depends less strongly on diversity. We derive a dynamical equilibrium theory for the CSDs of binary and fully polydisperse systems. Similarly to the clustering behaviour of one-component models, our qualitative results for mixtures are expected to be universal across active matter. Using literature experimental values for the self-propulsion speed diversity of unicellular swimmers known as choanoflagellates, which naturally differentiate into slower and faster cells, we predict that the error in estimating their $L_\text{c}$ via one-component models which use the conventional arithmetic mean self-propulsion speed is around $30\%$.
Phase transitions and collective dynamics of active colloidal suspensions are fascinating topics in soft matter physics, particularly for out-of-equilibrium systems, which can lead to rich rheological behaviours in the presence of steady shear flow. In this article, the role of self-propulsion in the rheological response of a dense colloidal suspension is investigated by using particle-resolved simulations. First, the interplay between activity and shear in the solid to the liquid transition of the suspension is analysed. While both self-propulsion and shear destroy order and melt the system b…
In the last years self-motile droplets attracted the attention of scientists from different fields ranging from applied biology to theoretical physics, because of their promising technological applications and important biological implications. In this Chapter we review the state of the art of the research on active droplets with a particular focus on theoretical and numerical studies. In particular, we reviewed the active gel theory, namely a generalization of the standard Landau-de Gennes theory for liquid crystals adapted to take into account internal active injection due to the presence of…
We study the behaviour of interacting self-propelled particles, whose self-propulsion speed decreases with their local density. By combining direct simulations of the microscopic model with an analysis of the hydrodynamic equations obtained by explicitly coarse graining the model, we show that interactions lead generically to the formation of a host of patterns, including moving clumps, active lanes and asters. This general mechanism could explain many of the patterns seen in recent experiments and simulations.
The ability of cells to move through small spaces depends on the mechanical properties of the cellular cytoskeleton and on nuclear deformability. In mammalian cells, the cytoskeleton is comprised of three interacting, semi-flexible polymer networks: actin, microtubules, and intermediate filaments (IF). Recent experiments of mouse embryonic fibroblasts with and without vimentin have shown that the IF vimentin plays a role in confined cell motility. We, therefore, develop a minimal model of cells moving through confined geometries that effectively includes all three types of cytoskeletal filamen…
We investigate how the competing presence of a nonuniform motility landscape and an external confining field affects the properties of active particles. We employ the active Ornstein-Uhlenbeck particle (AOUP) model with a periodic swim velocity profile to derive analytical approximations for the steady-state probability distribution of position and velocity, encompassing both the Unified Colored Noise Approximation and the theory of potential-free active particles with spatially dependent swim velocity recently developed. We test the theory by confining an active particle in a harmonic trap, w…
Motility-induced phase separation, MIPS, is arguably the most well-studied collective phenomenon occurring in active matter without alignment interactions (scalar active matter). Its basic origin is simple: since self-propelled particles accumulate where they move slowly, having a propulsion speed that decreases steeply enough with density, due to collisions or chemical interactions, leads to a feedback loop that induces the formation of a dense phase. In these notes, I will discuss some of the main theoretical and computational efforts that have been made over the last decade in understanding…
Drops of active liquid crystal have recently shown the ability to self-propel, which was associated with topological defects in the orientation of active filaments [Sanchez {\em et al.}, Nature {\bf 491}, 431 (2013)]. Here, we study the onset and different aspects of motility of a three-dimensional drop of active fluid on a planar surface. We analyse theoretically how motility is affected by orientation profiles with defects of various types and locations, by the shape of the drop, and by surface friction at the substrate. In the scope of a thin drop approximation, we derive exact expressions …
Designing microscopic and nanoscopic self-propelled particles and characterising their motion has become a major scientific challenge over the past decades. To this purpose, phoretic effects, namely propulsion mechanisms relying on local field gradients, have been the focus of many theoretical and experimental studies. In this review, we adopt a tutorial approach to present the basic physical mechanisms at stake in phoretic motion, and describe the different experimental works that lead to the fabrication of active particles based on this principle. We also present the collective effects obser…
The collective behaviour of confluent cell sheets is strongly influenced both by polar forces, arising through cytoskeletal propulsion and by active inter-cellular forces, which are mediated by interactions across cell-cell junctions. We use a phase-field model to explore the interplay between these two contributions and compare the dynamics of a cell sheet when the polarity of the cells aligns to (i) their main axis of elongation, (ii) their velocity, and (iii) when the polarity direction executes a persistent random walk.In all three cases, we observe a sharp transition from a jammed state (…
Growing, Buckling, and Swirling: motility from polymerization
International Centre for Theoretical Sciences, Tata Institute of Fundamental Research（国际理论科学中心，塔塔基础研究所）；Center for Computational Biology, Flatiron Institute（计算生物学中心，平顿研究所）；Courant Institute of Mathematical Sciences, New York University（纽约大学库朗数学科学研究所）
Active agents - like phoretic particles, bacteria, sperm, and cytoskeletal filaments in motility assays - show a large variety of motility-induced collective behaviors, such as aggregation, clustering and phase separation. The behavior of dense suspensions of phoretic particles and of bacteria during biofilm formation is determined by two principle physical mechanisms: (i) volume exclusion (short-range steric repulsion) and (ii) quorum sensing (longer-range reduced propulsion due to alteration of the local chemical environment). To systematically characterize such systems, we study semi-penetr…
We propose a novel mechanism of cell motility, which relies on the coupling of actin polymerization at the cell membrane to geometric confinement. We consider a polymerizing viscoelastic cytoskeletal gel confined in a narrow channel, and show analytically that spontaneous motion occurs. Interestingly, this does not require specific adhesion with the channel walls, and yields velocities potentially larger than the polymerization velocity. The contractile activity of myosin motors is not necessary to trigger motility in this mechanism, but is shown quantitatively to increase the velocity. Our mo…
Microorganisms are able to overcome the thermal randomness of their surroundings by harvesting energy to navigate in viscous fluid environments. In a similar manner, synthetic colloidal microswimmers are capable of mimicking complex biolocomotion by means of simple self-propulsion mechanisms. Although experimentally the speed of active particles can be controlled by e.g. self-generated chemical and thermal gradients, an in-situ change of swimming direction remains a challenge. In this work, we study self-propulsion of half-coated spherical colloids in critical binary mixtures and show that the…
Synthetic microswimmers offer models for cell motility and their tunability makes them promising candidates for biomedical applications. Here we measure the acoustic propulsion of bimetallic micro-rods that, when trapped at the nodal plane of a MHz acoustic resonator, swim with speeds of up to 300 microns per second. While past acoustic streaming models predict speeds that are more than one order of magnitude smaller than our measurements, we demonstrate that the acoustic locomotion of the rods is driven by a viscous, non-reciprocal mechanism relying on shape anisotropy akin to that used by sw…
We perform Brownian dynamics simulations in two dimensions to study the collective behavior of circle swimmers, which are driven by both, an (effective) translational and rotational self-propulsion, and interact via steric repulsion. We find that active rotation generally opposes motility-induced clustering and phase separation, as demonstrated by a narrowing of the coexistence region upon increase of the propulsion angular velocity. Moreover, although the particles are intrinsically assigned to rotate counterclockwise, a novel state of clockwise vortices emerges at an optimal value of the eff…
Bacterial migration through confined spaces is critical for several phenomena like: biofilm formation, bacterial transport in soils, and bacterial therapy against cancer . In the present work, \textit{E. coli} (strain K12-MG1655 WT) motility was characterized by recording and analyzing individual bacterium trajectories in a simulated quasi-2-dimensional porous medium. The porous medium was simulated by enclosing, between slide and cover slip, a bacterial-culture sample mixed with uniform 2.98 $μm$ spherical latex particles. The porosity of the medium was controlled by changing the latex partic…
We consider an active Brownian particle moving in a disordered two-dimensional energy or motility landscape. The averaged mean-square-displacement (MSD) of the particle is calculated analytically within a systematic short-time expansion. As a result, for overdamped particles, both an external random force field and disorder in the self-propulsion speed induce ballistic behaviour adding to the ballistic regime of an active particle with sharp self-propulsion speed. Spatial correlations in the force and motility landscape contribute only to the cubic and higher order powers in time for the MSD. …
Combining experiments on active colloids, whose propulsion velocity can be controlled via a feedback loop, and theory of active Brownian motion, we explore the dynamics of an overdamped active particle with a motility that depends explicitly on the particle orientation. In this case, the active particle moves faster when oriented along one direction and slower when oriented along another, leading to an anisotropic translational dynamics which is coupled to the particle's rotational diffusion. We propose a basic model of active Brownian motion for orientation-dependent motility. Based on this m…
In equilibrium, colloidal suspensions governed by short-range attractive and long-range repulsive interactions form thermodynamically stable clusters. Using Brownian dynamics computer simulations, we investigate how this equilibrium clustering is affected when such particles are self-propelled. We find that the clustering process is stable under self-propulsion. For the range of interaction parameters studied and at low particle density, the cluster size increases with the speed of self-propulsion (activity) and for higher activity the cluster size decreases, showing a non-monotonic variation …
The co-localization of density modulations and particle polarization is a characteristic emergent feature of motile active matter in activity gradients. It can therefore play the role of a smoking gun for the mesoscale detection of intrinsic microscopic activity. We employ the active-Brownian-particle (ABP) model to derive precise analytical expressions for the density and polarization profiles of a single Janus-type swimmer in the vicinity of an abrupt activity step, including situations with an orientation-dependent propulsion speed. Our results agree well with measurement data for a hot mic…
We study the dynamical steady-states of a monolayer of chemically active self-phoretic colloids as a function of packing fraction and self-propulsion speed by means of Brownian dynamics simulations. We focus on the case that a chemical field induces competing attractive positional and repulsive orientational interactions. Analyzing the distribution of cluster size and local density as well as the hexatic order parameter, we distinguish four distinct dynamical states which include collapsed, active gas, dynamical clustering, and motility-induced phase-separated states. The long-range chemical f…
Combining experiments and computer simulations, we use a spatially periodic and flashing light-field to direct the motion of phototactic active colloids. Here, the colloids self-organize into a density spike pattern, which resembles a shock wave and propagates over long distances, almost without dispersing. The underlying mechanism involves a synchronization of the colloids with the light-field, so that particles see the same intensity gradient each time the light-pattern is switched on, but no gradient in between (for example). This creates a pulsating transport whose strength and direction c…
Numerous physical models have been proposed to explain how cell motility emerges from internal activity, mostly focused on how crawling motion arises from internal processes. Here we offer a classification of self-propulsion mechanisms based on general physical principles, showing that crawling is not the only way for cells to move on a substrate. We consider a thin drop of active matter on a planar substrate and fully characterize its autonomous motion for all three possible sources of driving: (i) the stresses induced in the bulk by active components, which allow in particular tractionless m…
We develop a model of amoeboid cell motility based on active gel theory. Modeling the motile apparatus of a eukaryotic cell as a confined layer of finite length of poroelastic active gel permeated by a solvent, we first show that, due to active stress and gel turnover, an initially static and homogeneous layer can undergo a contractile-type instability to a polarized moving state in which the rear is enriched in gel polymer. This agrees qualitatively with motile cells containing an actomyosin-rich uropod at their rear. We find that the gel layer settles into a steadily moving, inhomogeneous st…
Micron-size self-propelling particles are often proposed as synthetic models for biological microswimmers, yet they lack internally regulated adaptation, which is central to the autonomy of their biological counterparts. Conversely, adaptation and autonomy can be encoded in larger-scale soft-robotic devices, but transferring these capabilities to the colloidal scale remains elusive. Here, we create a new class of responsive microswimmers, powered by induced-charge electrophoresis, which can adapt their motility to external stimuli via an internal feedback. Using sequential capillary assembly, …
A gold-capped Janus particle suspended in a near-critical binary liquid mixture can self-propel under illumination. We have immobilized such a particle in a narrow channel and studied the nonequilibrium dynamics of a binary solvent around it, using experiment and numerical simulations. For the latter we consider both a purely diffusive and a hydrodynamic model. All approaches indicate that the early time dynamics is purely diffusive and characterized by composition layers traveling with a constant speed from the surface of the colloid into the bulk. Subsequently, hydrodynamic effects set in an…
Many cell types display random motility on two-dimensional substrates, but crawl persistently in a single direction when confined in a microchannel or on an adhesive micropattern. Does this imply that the motility mechanism of confined cells is fundamentally different from that of unconfined cells? We argue that both free- and confined- cell migration may be described by a generic model of cells as "velocity aligning" active Brownian particles previously proposed to solve a completely separate problem in collective cell migration. Our model can be mapped to a diffusive escape over a barrier an…
We study the collective dynamics of self-propelled rods in an inhomogeneous motility field. At the interface between two regions of constant but different motility, a smectic rod layer is spontaneously created through aligning interactions between the active rods, reminiscent of an artificial, semi-permeable membrane. This "active membrane" engulfes rods which are locally trapped in low-motility regions and thereby further enhances the trapping efficiency by self-organization, an effect which we call "self-encapsulation". Our results are gained by computer simulations of self-propelled rod mod…
Using Brownian dynamics (BD) simulations we investigate the self-organization of a monolayer of chiral active particles with dipolar interactions. Each particle is driven by both, translational and rotational self-propulsion, and carries a permanent point dipole moment at its center. The direction of the translational propulsion for each particle is chosen to be parallel to its dipole moment. Simulations are performed at high dipolar coupling strength and a density below that related to motility-induced phase separation in simple active Brownian particles. Despite this restriction, we observe …
We introduce and study a model of active Brownian motion with multiplicative noise describing fluctuations in the self-propulsion or activity. We find that the standard picture of density accumulation in slow regions is qualitatively modified by active fluctuations, as stationary density profiles are generally not determined only by the mean self-propulsion speed landscape. As a result, activity gradients generically correlate the particle self-propulsion speed and orientation, leading to emergent polarization at interfaces pointing either towards dense or dilute regions depending on the amoun…
Micromotor and nanomotor particles are typically made using dense solid particles that can sediment or be trapped in confined flow environments. Creation of much larger motors should be possible if a very low-density system is used with sufficient strength to carry liquid and still experience propulsive motion. Light, dense millimotors should also be able to deform more than dense solid ones in constrictions. Millimotors are created from permeable capsules of bacterial cellulose that are coated with catalase-containing metal-organic frameworks, enabling reactive propulsion in aqueous hydrogen …
