Meso-scale turbulence was originally observed experimentally in various suspensions of swimming bacteria, as well as in the collective motion of active colloids. The corresponding large-scale dynamical patterns were reproduced in a simple model of a polar fluid, assuming a constant density of active particles. Recent, more detailed experimental studies revealed additional interesting aspects, such as anomalous velocity statistics and clustering phenomena. Those phenomena cannot be explained by currently available models for active polar fluids. Herein, we extend the continuum model suggested by Dunkel et al. to include density variations and a feedback between the local density and self-propulsion speed of the active particles. If the velocity decreases strong enough with the density, a linear stability analysis of the resulting model shows that, in addition to the short-wavelength instability of the original model, a long-wavelength instability occurs. This is typically observed for high densities of polar active particles and is analogous to the well-known phenomenon of motility-induced phase separation (MIPS) in scalar active matter. We determine a simple phase diagram indicating the linear instabilities and perform systematic numerical simulations for the various regions in the corresponding parameter space. The interplay between the well understood short-range instability and the long-range instability leads to interesting dynamics and novel phenomena concerning nucleation and coarsening processes. Our simulation results display a rich variety of novel patterns, including phase separation into domains with dynamically changing irregularly shaped boundaries. Anomalous velocity statistics are observed in all phases where the system segregates into regions of high and low densities. This offers a simple explanation for their occurrence in recent experiments with bacterial suspensions.
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 analyse a generic motility model, with the motility mechanism arising by contractile stress due to the interaction of myosin and actin. A hydrodynamic active polar gel theory is used to model the cytoplasm of a cell and is combined with a Helfrich-type model to account for membrane properties. The overall model allows to consider motility without the necessity for local adhesion. Besides a detailed numerical approach together with convergence studies for the highly nonlinear free boundary problem, we also compare the induced flow field of the motile cell with that of classical squirmer mode…
Combining model experiments and theory, we investigate the dense phases of polar active matter beyond the conventional flocking picture. We show that above a critical density flocks assembled from self-propelled colloids arrest their collective motion, lose their orientational order and form solids that actively rearrange their local structure while continuously melting and freezing at their boundaries. We establish that active solidification is a first-order dynamical transition: active solids nucleate, grow, and slowly coarsen until complete phase separation with the polar liquids they coexi…
We develop a dynamic mean-field theory for polar active particles that interact through a self-generated field, in particular one generated through emitting a chemical signal. While being a form of chemotactic response, it is different from conventional chemotaxis in that particles discontinuously change their motility when the local concentration surpasses a threshold. The resulting coupled equations for density and polarization are linear and can be solved analytically for simple geometries, yielding inhomogeneous density profiles. Specifically, here we consider a planar and circular interfa…
Active-particle suspensions exhibit distinct polarization-density patterns in activity landscapes, even without anisotropic particle interactions. Such polarization without alignment forces is at work in motility-induced phase separation and betrays intrinsic microscopic activity to mesoscale observers. Using stable long-term confinement of a single hot microswimmer in a dedicated force-free particle trap, we examine the polarized interfacial layer at a motility step and confirm that it does not exert pressure onto the bulk. Our observations are quantitatively explained by an analytical theory…
We study the phase behavior of polar Active Brownian Particles moving in two-spatial dimensions and interacting through volume exclusion and velocity alignment. We combine particle-based simulations of the microscopic model with a simple mean-field kinetic model to understand the impact of velocity alignment on the motility-induced phase separation of self-propelled disks. We show that, as the alignment strength is increased, approaching the onset of collective motion from below, orientational correlations grow, rendering the diffusive reorientation dynamics slower. As a consequence, the tende…
Motility-induced phase separation (MIPS) is a purely non-equilibrium phenomenon in which self-propelled particles phase separate without any attractive interactions. One surprising feature of MIPS is the emergence of polar, nematic, and higher order structures at the interfacial region, whose underlying physics remains poorly understood. Starting with a model of MIPS in which all many-body interactions are captured by an effective speed function and an effective pressure function that depend solely on the local particle density, I derive analytically an infinite set of integral formulae (IF) f…
We explore a generic mechanism whereby a droplet of active matter acquires motility by the spontaneous breakdown of a discrete symmetry. The model we study offers a simple representation of a "cell extract" comprising, e.g., a droplet of actomyosin solution. (Such extracts are used experimentally to model the cytoskeleton.) Actomyosin is an active gel whose polarity describes the mean sense of alignment of actin fibres. In the absence of polymerization and depolymerization processes ('treadmilling'), the gel's dynamics arises solely from the contractile motion of myosin motors; this should be …
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 (…
As the constituent particles of a flock are polar and in a driven state, their interactions must, in general, be fore-aft asymmetric and non-reciprocal. Within a model that explicitly retains the classical spin angular momentum field of the particles we show that the resulting asymmetric contribution to interparticle torques, if large enough, leads to a buckling instability of the flock. Precisely this asymmetry also yields a natural mechanism for a difference between the speed of advection of information along the flock and the speed of the flock itself, concretely establishing that the absen…
Many biological systems consist of self-motile and passive agents both of which contribute to overall functionality. However, there are very few studies of the properties of such mixtures. Here we formulate a model for mixtures of self-motile and passive agents and show that the model gives rise to three different dynamical phases: a disordered mesoturbulent phase, a polar flocking phase, and a vortical phase characterized by large-scale counterrotating vortices. We use numerical simulations to construct a phase diagram and compare the statistical properties of the different phases of the mode…
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…
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 present an experimental realization of the collective trapping phase transition [Kaiser et al., PRL 108, 268307 (2012)], using motile polar granular rods in the presence of a V-shaped obstacle. We offer a theory of this transition based on the interplay of motility-induced condensation and liquid-crystalline ordering and show that trapping occurs when persistent influx overcomes the collective expulsion of smectic defect structures. In agreement with the theory, our experiments find that a trap fills to the brim when the trap angle $θ$ is below a threshold $θ_c$, while all particles escape …
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…
We use coarse-grained molecular dynamics simulations to study the motility of a 2D vesicle containing self-propelled rods, as a function of the vesicle bending rigidity and the number density, length, and activity of the enclosed rods. Above a threshold value of the rod length, distinct dynamical regimes emerge, including a dramatic enhancement of vesicle motility characterized by a highly persistent random walk. These regimes are determined by clustering of the rods within the vesicle; the maximum motility state arises when there is one long-lived polar cluster. We develop a scaling theory th…
We present a numerical study of the phase behavior of repulsively interacting active polar particles that align their active velocities nematically. The amplitude of the active velocity, and the noise in its orientational alignment control the active nature of the system. At high values of orientational noise, the structural fluid undergoes a continuous nematic-isotropic transition in active orientation. This transition is well separated from an active phase separation, characterized by the formation of high density hexatic clusters, observed at lower noise strengths. With increasing activity,…
Active matter physics has been developed with various types of self-propelled particles, including those with polar and bipolar motility and beyond. However, the bipolar motions experimentally realized so far have been either random along the axis or periodic at intrinsic frequencies. Here we report another kind of bipolar active particles, whose periodic bipolar self-propulsion is set externally at a controllable frequency. We used Quincke rollers -- dielectric particles suspended in a conducting liquid driven by an electric field -- under an AC electric field instead of the usually used DC f…
A large class of mesoscopic or macroscopic flocking theories are coarse-grained from microscopic models that feature binary interactions as the chief aligning mechanism. However while such theories seemingly predict the existence of polar order with just binary interactions, actomyosin motility assay experiments show that binary interactions are insufficient to obtain polar order, especially at high densities. To resolve this paradox, here we introduce a solvable one-dimensional flocking model and derive its stochastic hydrodynamics. We show that two-body interactions are insufficient to gener…
We study the role of hydrodynamic interactions in the collective behaviour of collections of microscopic active particles suspended in a fluid. We introduce a novel calculational framework that allows us to separate the different contributions to their collective dynamics from hydrodynamic interactions on different length scales. Hence we are able to systematically show that lubrication forces when the particles are very close to each other play as important a role as long-range hydrodynamic interactions in determining their many-body behaviour. We find that motility-induced phase separation i…
The behavior of polar columnar phases made of bent shaped molecules (B_1Rev and B_1RevTilted) was studied under applied electric field. There are two competing mechanisms of switching in polar B_1RevTilted columnar phase: the collective rotation around the long molecular axis and the collective rotation around the tilt cone. Respectively monostable and bistable optical switching is observed for antiferroelectric B_1RevTilted phase. The main factor discriminating the type of the switching is a width of the column cross-section.
Many types of mammalian cells exert active contractile forces and mechanically deform their elastic substrate, to accomplish biological functions such as cell migration. These substrate deformations provide a mechanism by which cells can sense other cells, leading to long range mechanical intercell interactions and possible self organization. Here, we treat cells as noisy motile particles that exert contractile dipolar stresses on elastic substrates as they move. By combining this minimal model for the motility of individual cells with a linear elastic model that accounts for substrate mediate…
We study numerically the hydrodynamics of a self-propelled particle system, consisting of spherical squirmers sedimented on a flat surface. We observe the emergence of dynamic structures, due to the interplay of particle-particle and particle-wall hydrodynamic interactions. At low coverages, our results demonstrate the formation of small chiral spinners: two or three particles are bound together via near-field hydrodynamic interactions and form a rotating dimer or trimer respectively. The stability of the self-organised spinners can be tuned by the strength of the sedimentation. Increasing the…
