M. Cristina Marchetti

Athermal phase separation of self-propelled particles with no alignment

We study numerically and analytically a model of self-propelled polar disks on a substrate in two dimensions. The particles interact via isotropic repulsive forces and are subject to rotational noise, but there is no aligning interaction. As a result, the system does not exhibit an ordered state. The isotropic fluid phase separates well below close packing and exhibits the large number fluctuations and clustering found ubiquitously in active systems. Our work shows that this behavior is a generic property of systems that are driven out of equilibrium locally, as for instance by self-propulsion.

Statistical mechanics and hydrodynamics of bacterial suspensions

Unicellular living organisms, such as bacteria and algae, propel themselves through a medium via cyclic strokes involving the motion of cilia and flagella. Dense populations of such “active particles” or “swimmers” exhibit a rich collective behavior at large scales. Starting with a minimal physical model of a stroke-averaged swimmer in a fluid, we derive a continuum description of a suspension of active organisms that incorporates fluid-mediated, long-range hydrodynamic interactions among the swimmers. Our work demonstrates that hydrodynamic interactions provide a simple, generic origin for several nonequilibrium phenomena predicted or observed in the literature. The continuum model derived here does not depend on the microscopic physical model of the individual swimmer. The details of the large-scale physics do, however, differ for “shakers” (particles that are active but not self-propelled, such as melanocytes) and “movers” (self-propelled particles), “pushers” (most bacteria) and “pullers” (algae like Chlamydomonas). Our work provides a classification of the large-scale behavior of all these systems.

Topology and dynamics of active nematic vesicles

Liquid crystals on a deformable substrate The orientation of the molecules in a liquid crystalline material will change in response to either changes in the substrate or an external field. This is the basis for liquid crystalline devices. Vesicles, which are fluid pockets surrounded by lipid bilayers, will change size or shape in response to solvent conditions or pressure. Keber et al. report on the rich interactions between nematic liquid crystals placed on the surface of a vesicle. Changes to the vesicle size, for example, can “tune” the liquid crystal molecules. But conversely, the shape of the vesicles can also change in response to the activity of the nematic molecules. Science, this issue p. 1135 Dynamical shape-changing materials result from merging active liquid crystals with soft deformable vesicles. Engineering synthetic materials that mimic the remarkable complexity of living organisms is a fundamental challenge in science and technology. We studied the spatiotemporal patterns that emerge when an active nematic film of microtubules and molecular motors is encapsulated within a shape-changing lipid vesicle. Unlike in equilibrium systems, where defects are largely static structures, in active nematics defects move spontaneously and can be described as self-propelled particles. The combination of activity, topological constraints, and vesicle deformability produces a myriad of dynamical states. We highlight two dynamical modes: a tunable periodic state that oscillates between two defect configurations, and shape-changing vesicles with streaming filopodia-like protrusions. These results demonstrate how biomimetic materials can be obtained when topological constraints are used to control the non-equilibrium dynamics of active matter.

Cadherin-based intercellular adhesions organize epithelial cell–matrix traction forces

Cell–cell and cell–matrix adhesions play essential roles in the function of tissues. There is growing evidence for the importance of cross talk between these two adhesion types, yet little is known about the impact of these interactions on the mechanical coupling of cells to the extracellular matrix (ECM). Here, we combine experiment and theory to reveal how intercellular adhesions modulate forces transmitted to the ECM. In the absence of cadherin-based adhesions, primary mouse keratinocytes within a colony appear to act independently, with significant traction forces extending throughout the colony. In contrast, with strong cadherin-based adhesions, keratinocytes in a cohesive colony localize traction forces to the colony periphery. Through genetic or antibody-mediated loss of cadherin expression or function, we show that cadherin-based adhesions are essential for this mechanical cooperativity. A minimal physical model in which cell–cell adhesions modulate the physical cohesion between contractile cells is sufficient to recreate the spatial rearrangement of traction forces observed experimentally with varying strength of cadherin-based adhesions. This work defines the importance of cadherin-based cell–cell adhesions in coordinating mechanical activity of epithelial cells and has implications for the mechanical regulation of epithelial tissues during development, homeostasis, and disease.

Instabilities of isotropic solutions of active polar filaments.

We study the dynamics of an entangled, isotropic solution of polar filaments coupled by molecular motors which generate relative motion of the filaments in two and three dimensions. We investigate the stability of the homogeneous state for constant motor concentration taking into account excluded volume and entanglement. At low filament density the system develops a density instability, while at high filament density entanglement effects drive the instability of orientational fluctuations.

Defect annihilation and proliferation in active nematics.

Liquid crystals inevitably possess topological defect excitations generated through boundary conditions, through applied fields, or in quenches to the ordered phase. In equilibrium, pairs of defects coarsen and annihilate as the uniform ground state is approached. Here we show that defects in active liquid crystals exhibit profoundly different behavior, depending on the degree of activity and its contractile or extensile character. While contractile systems enhance the annihilation dynamics of passive systems, extensile systems act to drive defects apart so that they swarm around in the manner of topologically well-characterized self-propelled particles. We develop a simple analytical model for the defect dynamics which reproduces the key features of both the numerical solutions and recent experiments on microtubule-kinesin assemblies.

Geometry Regulates Traction Stresses in Adherent Cells

Cells generate mechanical stresses via the action of myosin motors on the actin cytoskeleton. Although the molecular origin of force generation is well understood, we currently lack an understanding of the regulation of force transmission at cellular length scales. Here, using 3T3 fibroblasts, we experimentally decouple the effects of substrate stiffness, focal adhesion density, and cell morphology to show that the total amount of work a cell does against the substrate to which it is adhered is regulated by the cell spread area alone. Surprisingly, the number of focal adhesions and the substrate stiffness have little effect on regulating the work done on the substrate by the cell. For a given spread area, the local curvature along the cell edge regulates the distribution and magnitude of traction stresses to maintain a constant strain energy. A physical model of the adherent cell as a contractile gel under a uniform boundary tension and mechanically coupled to an elastic substrate quantitatively captures the spatial distribution and magnitude of traction stresses. With a single choice of parameters, this model accurately predicts the cells mechanical output over a wide range of cell geometries.

Fluctuations and Pattern Formation in Self-Propelled Particles

We consider a coarse-grained description of a collection of self-propelled particles given by hydrodynamic equations for the density and polarization fields. We find that the ordered moving or flocking state of the system is unstable to spatial fluctuations beyond a threshold set by the self-propulsion velocity of the individual units. In this region, the system organizes itself into an inhomogeneous state of well-defined propagating stripes of flocking particles interspersed with low-density disordered regions. Further, we find that even in the regime where the homogeneous flocking state is stable, the system exhibits large fluctuations in both density and orientational order. We study the hydrodynamic equations analytically and numerically to characterize both regimes.

Hydrodynamics of self-propelled hard rods.

Motivated by recent simulations and by experiments on aggregation of gliding bacteria, we study a model of the collective dynamics of self-propelled hard rods on a substrate in two dimensions. The rods have finite size, interact via excluded volume, and their dynamics is overdamped by the interaction with the substrate. Starting from a microscopic model with nonthermal noise sources, a continuum description of the system is derived. The hydrodynamic equations are then used to characterize the possible steady states of the systems and their stability as a function of the particles packing fraction and the speed of self-propulsion.

Enhanced Diffusion and Ordering of Self-Propelled Rods

Starting from a minimal physical model of self-propelled hard rods on a substrate in two dimensions, we derive a modified Smoluchowski equation for the system. Self-propulsion enhances longitudinal diffusion and modifies the mean-field excluded volume interaction. From the Smoluchowski equation we obtain hydrodynamic equations for rod concentration, polarization and nematic order parameter. New results at large scales are a lowering of the density of the isotropic-nematic transition and a strong enhancement of boundary effects in confined self-propelled systems.