Aparna Baskaran

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.

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.

Orientational order of motile defects in active nematics.

The study of liquid crystals at equilibrium has led to fundamental insights into the nature of ordered materials, as well as to practical applications such as display technologies. Active nematics are a fundamentally different class of liquid crystals, driven away from equilibrium by the autonomous motion of their constituent rod-like particles. This internally generated activity powers the continuous creation and annihilation of topological defects, which leads to complex streaming flows whose chaotic dynamics seem to destroy long-range order. Here, we study these dynamics in experimental and computational realizations of active nematics. By tracking thousands of defects over centimetre-scale distances in microtubule-based active nematics, we identify a non-equilibrium phase characterized by a system-spanning orientational order of defects. This emergent order persists over hours despite defect lifetimes of only seconds. Similar dynamical structures are observed in coarse-grained simulations, suggesting that defect-ordered phases are a generic feature of active nematics.

Dynamics of self-propelled particles under strong confinement

We develop a statistical theory for the dynamics of non-aligning, non-interacting self-propelled particles confined in a convex box in two dimensions. We find that when the size of the box is small compared to the persistence length of a particles trajectory (strong confinement), the steady-state density is zero in the bulk and proportional to the local curvature on the boundary. Conversely, the theory may be used to construct the box shape that yields any desired density distribution on the boundary, thus offering a general tool to understand and design such confinements. When the curvature variations are small, we also predict the distribution of orientations at the boundary and the exponential decay of pressure as a function of box size recently observed in simulations in a spherical box.

Reentrant phase behavior in active colloids with attraction

Motivated by recent experiments, we study a system of self-propelled colloids that experience short-range attractive interactions and are confined to a surface. Using simulations we find that the phase behavior for such a system is reentrant as a function of activity: phase-separated states exist in both the low- and high-activity regimes, with a homogeneous active fluid in between. To understand the physical origins of reentrance, we develop a kinetic model for the systems steady-state dynamics whose solution captures the main features of the phase behavior. We also describe the varied kinetics of phase separation, which range from the familiar nucleation and growth of clusters to the complex coarsening of active particle gels.

Spontaneous segregation of self-propelled particles with different motilities

We study mixtures of self-propelled and passive rod-like particles in two dimensions using Brownian dynamics simulations. The simulations demonstrate that the two species spontaneously segregate to generate a rich array of dynamical domain structures whose properties depend on the propulsion velocity, density, and composition. In addition to presenting phase diagrams as a function of the system parameters, we investigate the mechanisms driving segregation. We show that the difference in collision frequencies between self-propelled and passive rods provides a driving force for segregation, which is amplified by the tendency of the self-propelled rods to swarm or cluster. Finally, both self-propelled and passive rods exhibit giant number fluctuations for sufficient propulsion velocities.

Structure and Dynamics of a Phase-Separating Active Colloidal Fluid

We examine a minimal model for an active colloidal fluid in the form of self-propelled Brownian spheres that interact purely through excluded volume with no aligning interaction. Using simulations and analytic modeling, we quantify the phase diagram and separation kinetics. We show that this nonequilibrium active system undergoes an analog of an equilibrium continuous phase transition, with a binodal curve beneath which the system separates into dense and dilute phases whose concentrations depend only on activity. The dense phase is a unique material that we call an active solid, which exhibits the structural signatures of a crystalline solid near the crystal-hexatic transition point, and anomalous dynamics including superdiffusive motion on intermediate time scales.

Nonequilibrium statistical mechanics of self-propelled hard rods

Using tools of nonequilibrium mechanics, we study a model of self-propelled hard rods on a substrate in two dimensions to quantify the interplay of self-propulsion and excluded-volume effects. We derive a Smoluchowski equation for the configurational probability density of self-propelled rods that contains several modifications as compared to the familiar Smoluchowski equation for thermal rods. As a side-product of our work, we also present a purely dynamical derivation of the Onsager form of the mean-field excluded-volume interaction among thermal hard rods.