The statistical physics of active matter: from self-catalytic colloids to living cells

These lecture notes are designed to provide a brief introduction into the phenomenology of active matter and to present some of the analytical tools used to rationalize the emergent behavior of active systems. Such systems are made of interacting agents able to extract energy stored in the environment to produce sustained directed motion. The local conversion of energy into mechanical work drives the system far from equilibrium, yielding new dynamics and phases. The emerging phenomena can be classified depending on the symmetry of the active particles and on the type of microscopic interactions. We focus here on steric and aligning interactions, as well as interactions driven by shape changes. The models that we present are all inspired by experimental realizations of either synthetic, biomimetic or living systems. Based on minimal ingredients, they are meant to bring a simple and synthetic understanding of the complex phenomenology of active matter.Comment: Lecture notes for the international summer school "Fundamental Problems in Statistical Physics" 2017 in Brunec

Hydrodynamics of Turning Flocks

We present a hydrodynamic model of flocking that generalizes the familiar Toner-Tu equations to incorporate turning inertia of well-polarized flocks. The continuum equations controlled by only two dimensionless parameters, orientational inertia and alignment strength, are derived by coarse graining the inertial spin model recently proposed by Cavagna et al. The interplay between orientational inertia and bend elasticity of the flock yields anisotropic spin waves that mediate the propagation of turning information throughout the flock. The coupling between spin current density to the local vorticity field through a nonlinear friction gives rise to a hydrodynamic mode with angular-dependent propagation speed at long wavelength. This mode goes unstable as a result of the growth of bend and splay deformations augmented by the spin wave, signaling the transition to complex spatio-temporal patterns of continuously turning and swirling flocks.Comment: 12 pages, 3 figure

Hydrodynamic and rheology of active polar filaments

The cytoskeleton provides eukaryotic cells with mechanical support and helps them perform their biological functions. It is a network of semiflexible polar protein filaments and many accessory proteins that bind to these filaments, regulate their assembly, link them to organelles and continuously remodel the network. Here we review recent theoretical work that aims to describe the cytoskeleton as a polar continuum driven out of equilibrium by internal chemical reactions. This work uses methods from soft condensed matter physics and has led to the formulation of a general framework for the description of the structure and rheology of active suspension of polar filaments and molecular motors.Comment: 30 pages, 5 figures. To appear in "Cell Motility", Peter Lenz, ed. (Springer, New York, 2007

Organization and instabilities of entangled 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.Comment: 4pages, 2 eps figure, revtex

Active Jamming: Self-propelled soft particles at high density

We study numerically the phases and dynamics of a dense collection of self-propelled particles with soft repulsive interactions in two dimensions. The model is motivated by recent in vitro experiments on confluent monolayers of migratory epithelial and endothelial cells. The phase diagram exhibits a liquid phase with giant number fluctuations at low packing fraction and high self-propulsion speed and a jammed phase at high packing fraction and low self-propulsion speed. The dynamics of the jammed phase is controlled by the low frequency modes of the jammed packing.Comment: 4 pages, 4 figure

The low noise phase of a 2d active nematic

We consider a collection of self-driven apolar particles on a substrate that organize into an active nematic phase at sufficiently high density or low noise. Using the dynamical renormalization group, we systematically study the 2d fluctuating ordered phase in a coarse-grained hydrodynamic description involving both the nematic director and the conserved density field. In the presence of noise, we show that the system always displays only quasi-long ranged orientational order beyond a crossover scale. A careful analysis of the nonlinearities permitted by symmetry reveals that activity is dangerously irrelevant over the linearized description, allowing giant number fluctuations to persist though now with strong finite-size effects and a non-universal scaling exponent. Nonlinear effects from the active currents lead to power law correlations in the density field thereby preventing macroscopic phase separation in the thermodynamic limit.Comment: 17 pages, 5 figure

Controlling cell-matrix traction forces by extracellular geometry

We present a minimal continuum model of strongly adhering cells as active contractile isotropic media and use the model to study the effect of the geometry of the adhesion patch in controlling the spatial distribution of traction and cellular stresses. Activity is introduced as a contractile, hence negative, spatially homogeneous contribution to the pressure. The model shows that patterning of adhesion regions can be used to control traction stress distribution and yields several results consistent with experimental observations. Specifically, the cell spread area is found to increase with substrate stiffness and an analytic expression for the dependence is obtained for circular cells. The correlation between the magnitude of traction stresses and cell boundary curvature is also demonstrated and analyzed.Comment: 12 pages, 4 figure