We examine the interplay of motility and information exchange in a model of run-and-tumble active particles where the particle's motility is encoded as a bit of information that can be exchanged upon contact according to the rules of AND and OR logic gates in a circuit. Motile AND particles become non-motile upon contact with a non-motile particle. Conversely, motile OR particles remain motile upon collision with their non-motile counterparts. AND particles that have become non-motile additionally "reawaken", i.e., recover their motility, at a fixed rate $μ$, as in the SIS (Susceptible, Infected, Susceptible) model of epidemic spreading, where an infected agent can become healthy again, but keeps no memory of the recent infection, hence it is susceptible to a renewed infection. For $μ=0$, both AND and OR particles relax irreversibly to absorbing states of all non-motile or all motile particles, respectively. The relaxation kinetics is, however, faster for OR particles that remain active throughout the process. At finite $μ$, the AND dynamics is controlled by the interplay between reawakening and collision rates. The system evolves to a state of all motile particles (an absorbing state in the language of absorbing phase transitions) for $μ>μ_c$ and to a mixed state with coexisting motile and non-motile particles (an active state in the language of absorbing phase transitions) for $μ<μ_c$. The final state exhibits a rich structure controlled by motility-induced aggregation. Our work can be relevant to biochemical signaling in motile bacteria, the spreading of epidemics and of social consensus, as well as light-controlled organization of active colloids.
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 migration behaviors of cancer cells are known to be heterogeneous. However, the interplay between the adhesion interactions, dynamical shape changes and fluid flows in regulating cell migration heterogeneity and plasticity during cancer metastasis is still elusive. To further quantitative understanding of cell motility and morphology, we develop a theory using stochastic quantization method that describes the role of biophysical cues in regulating diverse cell motility. We show that the cumulative effect of time dependent adhesion interactions that determine the structural rearrangements a…
Collectives of actively-moving particles can spontaneously separate into dilute and dense phases -- a fascinating phenomenon known as motility-induced phase separation (MIPS). MIPS is well-studied for randomly-moving particles with no directional bias. However, many forms of active matter exhibit collective chemotaxis, directed motion along a chemical gradient that the constituent particles can generate themselves. Here, using theory and simulations, we demonstrate that collective chemotaxis strongly competes with MIPS -- in some cases, arresting or completely suppressing phase separation, or …
Collective cell migration in cohesive units is vital for tissue morphogenesis, wound repair, and immune response. While the fundamental driving forces for collective cell motion stem from contractile and protrusive activities of individual cells, it remains unknown how their balance is optimized to maintain tissue cohesiveness and the fluidity for motion. Here we present a cell-based computational model for collective cell migration during wound healing that incorporates mechanochemical coupling of cell motion and adhesion kinetics with stochastic transformation of active motility forces. We s…
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…
Dense active systems are widespread in nature, examples range from bacterial colonies to biological tissues. Dense clusters of active particles can be obtained by increasing the packing fraction of the system or taking advantage of a peculiar phenomenon named motility-induced phase separation (MIPS). In this work, we explore the phase diagram of a two-dimensional model of active glass and show that disordered active materials develop a rich collective behaviour encompassing both MIPS and glassiness. We find that, although the glassy state is almost indistinguishable from that of equilibrium gl…
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 (…
We study a generic model for quorum-sensing bacteria in circular confinement. Every bacterium produces signaling molecules, the local concentration of which triggers a response when a certain threshold is reached. If this response lowers the motility then an aggregation of bacteria occurs, which differs fundamentally from standard motility-induced phase separation due to the long-ranged nature of the concentration of signal molecules. We analyze this phenomenon analytically and by numerical simulations employing two different protocols leading to stationary cluster and ring morphologies, respe…
Epithelial cell clusters often move collectively on a substrate. Mechanical signals play a major role in organizing this behavior. There are a number of experimental observations in these systems which await a comprehensive explanation. These include: the internal strains are tensile even for clusters that expand by proliferation; the tractions on the substrate are often confined to the edges of the cluster; there can exist density waves within the cluster; clusters can exhibit collective durotaxis when individual cells show no effect; and for cells in an annulus there is a transition between …
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…
Contact Forces in Motility-Regulated Active Matter
Massachusetts Institute of Technology（麻省理工学院）；Stanford University（斯坦福大学）；University of Geneva（日内瓦大学）；Université Paris Cité（巴黎西岱大学）；University of Oxford（牛津大学）
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…
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 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…
Substrate-based cell motility is essential for fundamental biological processes, such as tissue growth, wound healing and immune response. Even if a comprehensive understanding of this motility mode remains elusive, progress has been achieved in its modeling using a whole cell physical model. The model takes into account the main mechanisms of cell motility - actin polymerization, substrate mediated adhesion and actin-myosin dynamics and combines it with steric cell-cell and hydrodynamic interactions. The model predicts the onset of collective cell migration, which emerges spontaneously as a r…
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…
Active matter consists of self-propelled elements exhibits fascinating collective motions ranging from biological to artificial systems. Among wide varieties of active matter systems, reconstituted bio-filaments moving on molecular motor turf interacting purely by physical interactions provides the fundamental test ground for understanding biological motility. However, until now, multi-filament collisions,depletion agents or binding molecules has been required for the emergence of ordered patterns in motility assay. Thus, whether simple physical interactions during collisions such as steric ef…
Collective cell migration underlies important biological processes, such as embryonic development, wound healing and cancer invasion. While many aspects of single cell movements are now well established, the mechanisms leading to displacements of cohesive cell groups are still poorly understood. To elucidate the emergence of collective migration in mechanosensitive cells, we examine a self-propelled Voronoi (SPV) model of confluent tissues with an orientational feedback that aligns a cell's polarization with its local migration velocity. While shape and motility are known to regulate a density…
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…
One of the most intriguing phenomena in active matter has been the gas-liquid like motility induced phase separation (MIPS) observed in repulsive active particles. However, experimentally no particle can be a perfect sphere, and the asymmetric shape, mass distribution or catalysis coating can induce an active torque on the particle, which makes it a chiral active particle. Here using computer simulations and dynamic mean-field theory, we demonstrate that the large enough torque of circle active Brownian particles (cABPs) in two dimensions generates a dynamical clustering state interrupting the…
We show that memory, in the form of underdamped angular dynamics, is a crucial ingredient for the collective properties of self-propelled particles. Using Vicsek-style models with an Ornstein-Uhlenbeck process acting on angular velocity, we uncover a rich variety of collective phases not observed in usual overdamped systems, including vortex lattices and active foams. In a model with strictly nematic interactions the smectic arrangement of Vicsek waves giving rise to global polar order is observed. We also provide a calculation of the effective interaction between vortices in the case where a …
The dynamics of active matter driven by interacting molecular motors has a non-potential structure at the local scale. However, we show that there exists a quasi-potential effectively describing the collective self-organization of the motors propelling a cell at a continuum active gel level. Such a model allows us to understand cell motility as an active phase transition problem between the static and motile steady state configurations that minimize the quasi-potential. In particular both configurations can coexist in a metastable fashion and a small stochastic disorder in the gel is sufficien…
Active matter deals with systems whose particles consume energy at the individual level in order to move. To unravel features such as the emergence of collective structures several models have been suggested, such as the on-lattice model of run-and-tumble particles implemented via the Persistent Exclusion Process (PEP). In our work, we study a one dimensional system of run-and-tumble repulsive or attractive particles, both on and off lattice. Additionally, we implement a cluster motility dynamics in the on-lattice case (since in the off-lattice case cluster motility arises from the individual …
Active matter systems provide fascinating examples of pattern formation and collective motility without counterparts in equilibrium systems. Here, we employ Brownian dynamics simulations to study the collective motion and self-organization in systems of self-propelled semiflexible filaments, inspired by the gliding motility of \textit{filamentous Cyanobacteria}. Specifically, we investigate the influence of stochastic direction reversals on the patterns. We explore pattern formation and dynamics by modulating three relevant physical parameters, the bending stiffness, the activity, and the reve…
We investigate theoretically the collective dynamics of soft active particles living in a viscous fluid. We focus on a minimal model for active but non-motile particles consisting of $N>1$ elastic dimers deformed by active stresses and interacting hydrodynamically. We first derive a set of effective equations of motion for the positions of the particles. We then exploit these equations in two experimentally-relevant cases: uncorrelated random internal stresses, and uniform monochromatic external shaking. In both cases, we show that small groups of intrinsically non-motile particles can display…
Active matter comprises individually driven units that convert locally stored energy into mechanical motion. Interactions between driven units lead to a variety of non-equilibrium collective phenomena in active matter. One of such phenomena is anomalously large density fluctuations, which have been observed in both experiments and theories. Here we show that, on the contrary, density fluctuations in active matter can also be greatly suppressed. Our experiments are carried out with marine algae ($\it{Effrenium\ voratum}$) which swim in circles at the air-liquid interfaces with two different euk…
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 b…
The creation of synthetic systems that emulate the defining properties of living matter, such as motility, gradient-sensing, signalling and replication, is a grand challenge of biomimetics. Such imitations of life crucially contain active components that transform chemical energy into directed motion. These artificial realizations of motility point in the direction of a new paradigm in en- gineering, through the design of emergent behavior by manipulating properties at the scale of the individual components. Catalytic colloidal swimmers are a particularly promising example of such systems. Her…
In this work, we study the dynamics of a single active Brownian particle, as well as the collective behavior of interacting active Brownian particles, in a fluctuating heterogeneous environment. We employ a variant of the diffusing diffusivity model where the equation of motion of the active particle involves a time-dependent motility and diffusivities. Within our model, those fluctuations are coupled to each other. Using analytical methods, we obtain the probability distribution function of particle displacement and its moments for a single particle. We then investigate the impact of the envi…
We investigate the nonequilibrium dynamics of spherical active Brownian particles in three spatial dimensions that interact via a pair potential. The investigation is based on a predictive local field theory that is derived by a rigorous coarse-graining starting from the overdamped Langevin dynamics of the particles. This field theory is highly accurate and applicable even for the highest activities. It includes configurational order parameters and derivatives up to infinite orders. We present also three finite reduced models that result from the general field theory by suitable approximations…
We discuss recent progress in the theoretical description of chemotaxis by coupling the diffusion equation of a chemical species to equations describing the motion of sensing microorganisms. In particular, we discuss models for autochemotaxis of a single microorganism which senses its own secretion leading to phenomena such as self-localization and self-avoidance. For two heterogeneous particles, chemotactic coupling can lead to predator-prey behavior including chase and escape phenomena, and to the formation of active molecules, where motility spontaneously emerges when the particles approach…
Tracking experiments in dense biological tissues reveal a diversity of sources f or local energy injection at the cell scale. The effect of cell motility has been largely studied, but much less is known abo ut the effect of the observed volume fluctuations of individual cells. We devise a simple microscopic model of `actively-deforming' particles where local fluctuations of the particle size constitute a unique source of motion. We demonstrate that collective motion can emerge under the sole influence of such active volume fluctuations. We interpret the onset of diffusive motion as a nonequili…
Recently, non-reciprocal systems have become a focus of growing interest. Examples occur in soft and active matter, but also in engineered quantum materials and neural (brain) networks. Here, we investigate the impact of non-reciprocity on the collective behavior of a system of (dry) chiral active matter. Specifically, we consider a mixture of "circle swimmers" with steric interactions and non-reciprocal alignment couplings. Based on hydrodynamic equations which we derive from a set of Langevin equations, we explore the interplay of non-reciprocity, finite size, and chirality. We first conside…
Collective cell migration is a highly regulated process involved in wound healing, cancer metastasis and morphogenesis. Mechanical interactions among cells provide an important regulatory mechanism to coordinate such collective motion. Using a Self-Propelled Voronoi (SPV) model that links cell mechanics to cell shape and cell motility, we formulate a generalized mechanical inference method to obtain the spatio-temporal distribution of cellular stresses from measured traction forces in motile tissues and show that such traction-based stresses match those calculated from instantaneous cell shape…
Here we characterize the motility of athermal swimming droplets within the framework of active rotational diffusion. Just like active colloids, their trajectories can be modeled with a constant velocity $V$ and a slow angular diffusion, but the random changes in direction are not thermally driven. Instead, $V$ is determined by the interfacial tension gradient along the droplet surface, while local micellar fluctuations lead to droplet reorientation with a persistence time $τ$. We show that the origin of locomotion is the difference in the critical micellar concentration $Δ$CMC in the front and…
