We calculate the Casimir stresses in a thin layer of active fluid with nematic order. By using a stochastic hydrodynamic approach for an active fluid layer of finite thickness $L$, we generalize the Casimir stress for nematic liquid crystals in thermal equilibrium to active systems. We show that the active Casimir stress differs significantly from its equilibrium counterpart. For contractile activity, the active Casimir stress, although attractive like its equilibrium counterpart, diverges logarithmically as $L$ approaches a threshold of the spontaneous flow instability from below. In contrast, for small extensile activity, it is repulsive, has no divergence at any $L$ and has a scaling with $L$ different from its equilibrium counterpart.
We investigate the pseudo-Casimir force in a slab of material composed of nematically ordered long polymers. We write the total mesoscopic energy together with the constraint connecting the local density and director fluctuations and evaluate the corresponding fluctuation free energy by standard methods. It leads to a pseudo-Casimir force of a different type than in the case of standard, short molecule nematic. We investigate its separation dependence and its magnitude and explicitly derive the relevant limiting cases.
The elastic interactions between objects embedded in a nematic liquid crystal are usually caused by the average distorsion-rather than by the fluctuations-of the nematic orientational field. We argue that for sufficiently small particles, the nematic-mediated interaction originates purely from the fluctuations of the nematic director. This Casimir interaction decays as d^(-6), d being the distance between the particles, and it dominates van der Waals interactions close to the isotropic-to-nematic transition. Considering the nematic as a polymer solvent, we show that the onset of this Casimir i…
Fluctuation-induced interactions between anisotropic objects immersed in a nematic liquid crystal are shown to depend on the relative orientation of these objects. The resulting long-range ``Casimir'' torques are explicitely calculated for a simple geometry where elastic effects are absent. Our study generalizes previous discussions restricted to the case of isotropic walls, and leads to new proposals for experimental tests of Casimir forces and torques in nematics.
We consider the effective pseudo-Casimir interaction forces mediated by a nematic liquid-crystalline film bounded by two planar surfaces, one of which imposes a random (disordered) distribution of the preferred anchoring axis in the so-called easy direction. We consider both the case of a quenched as well as an annealed disorder for the easy direction on the disordered surface and analyze the resultant fluctuation-induced interaction between the surfaces. In the case of quenched disorder, we show that the disorder effects appear additively in the total interaction and are dominant at intermedi…
We consider a nematic liquid crystal confined by two parallel planar interfaces, one being laterally homogeneous and the other provided by a substrate endowed with a periodic chemical stripe pattern. Based on continuum theory we analyze the influence of the lateral pattern on the liquid-crystalline Casimir force acting on the interfaces of the nematic cell at distance d due to the thermal fluctuations of the nematic director. For d much larger than the pattern periodicity, the influence of the patterned substrate can be described by a homogeneous, effective anchoring energy. By tuning this par…
Recent experiments on monolayers of spindle-like cells plated on adhesive stripe-shaped domains have provided a convincing demonstration that certain types of collective phenomena in epithelia are well described by active nematic hydrodynamics. While recovering some of the hallmark predictions of this framework, however, these experiments have also revealed a number of unexpected features that could be ascribed to the existence of chirality over length scales larger than the typical size of a cell. In this article we elaborate on the microscopic origin of chiral stresses in nematic cell monola…
We examine fluctuation-induced (pseudo-Casimir) interactions in nematic liquid-crystalline films confined between two surfaces, where one of the surfaces imposes a strong homeotropic anchoring (ensuring a uniform mean director profile), while the other one is assumed to be a chemically disordered substrate exhibiting an annealed, random distribution of anchoring energies. We employ a saddle-point approximation to evaluate the free energy of interaction mediated between the two surfaces and investigate how the interaction force is influenced by the presence of disordered surface anchoring energ…
Ordering configurations of a director field on a curved membrane induce stress. In this work, we present a theoretical framework to calculate the stress tensor and the torque as a consequence of the nematic ordering; we use the variational principle and invariance of the energy under Euclidean motions. Euler-Lagrange equations of the membrane as well as the corresponding boundary conditions also appear as natural results. The stress tensor found includes attraction-repulsion forces between defects; likewise, defects are attracted to patches with the same sign in gaussian curvature. These force…
Topological defects are increasingly being identified in various biological systems, where their characteristic flow fields and stress patterns are associated with continuous active stress generation by biological entities. Here, using numerical simulations of continuum fluctuating nematohydrodynamics we show that even in the absence of any specific form of active stresses associated with self-propulsion, mesoscopic fluctuations in either orientational alignment or hydrodynamics can independently result in flow patterns around topological defects that resemble the ones observed in active syste…
Investigations of the elastic behavior of bulk elastomers have traditionally proceeded on the basis of classical rubber elasticity, which regards chains as thermally fluctuating but disregards the thermal fluctuations of the cross-links. Here, we consider an incompressible and flat elastomer film of an axisymmetric shape confined between two large hard co-planar substrates, with the axis of the film perpendicular to the substrates. We address the impact that thermal fluctuations of the cross-links have on the free energy of elastic deformation of the system, subject to the requirement that the…
The topological properties of many materials are central to their behavior, with the dynamics of topological defects being particularly important to intrinsically out-of-equilibrium, active materials. In this paper, local manipulation of the ordering, dynamics, and topological properties of microtubule-based extensile active nematic films is demonstrated in a joint experimental and simulation study. Hydrodynamic stresses created by magnetically actuated rotation of disk-shaped colloids in proximity to the films compete with internal stresses in the active nematic, enabling local control of the…
Recent experiments in various cell types have shown that two-dimensional tissues often display local nematic order, with evidence of extensile stresses manifest in the dynamics of topological defects. Using a mesoscopic model where tissue flow is generated by fluctuating traction forces coupled to the nematic order parameter, we show that the resulting tissue dynamics can spontaneously produce local nematic order and an extensile internal stress. A key element of the model is the assumption that in the presence of local nematic alignment, cells preferentially crawl along the nematic axis, resu…
With quenched disorder, we introduce two-dimensional active nematics suspended in an incompressible fluid. We write the coarse-grained hydrodynamic equations of motion for slow variables, viz. density, orientation and flow fields. The quenched disorder is introduced such that it interacts with the local orientation at every point with some strength. Disorder strength is tuned from zero to large values. We numerically study the defect dynamics and system's kinetics and find that the finite disorder slows the ordering. The presence of fluid induces large fluctuation in the orientation field, fur…
An elastic membrane with embedded nematic molecules is considered as a model of anisotropic fluid membrane with internal ordering. By considering the geometric coupling between director field and membrane curvature, the nematic texture is shown to induce anisotropic stresses additional to Canham-Helfrich elasticity. Building upon differential geometry, analytical expressions are found for the membrane stress and torque induced by splaying, twisting and bending of the nematic director as described by the Frank energy of liquid crystals. The forces induced by prototypical nematic textures are vi…
A continuum description is essential for understanding a variety of collective phenomena in active matter. However, building quantitative continuum models of active matter from first principles can be extremely challenging due to both the gaps in our knowledge and the complicated structure of nonlinear interactions. Here we use a novel physically-informed data-driven approach to construct a complete mathematical model of an active nematic from experimental data describing kinesin-driven microtubule bundles confined to an oil-water interface. We find that the structure of the model is similar t…
We investigate the mechanics of an active droplet endowed with internal nematic order and surrounded by an isotropic Newtonian fluid. Using numerical simulations we demonstrate that, due to the interplay between the active stresses and the defective geometry of the nematic director, this system exhibits two of the fundamental functions of living cells: spontaneous division and motility, by means of self-generated hydrodynamics flows. These behaviors can be selectively activated by controlling a single physical parameter, namely, an active variant of the capillary number.
Active nematics are the nonequilibrium analog of passive liquid crystals in which anisotropic units consume free energy to drive emergent behavior. Similar to liquid crystal (LC) molecules in displays, ordering and dynamics in active nematics are sensitive to boundary conditions; however, unlike passive liquid crystals, active nematics, such as those composed of living matter, have the potential to regulate their boundaries through self-generated stresses. Here, using bacterial biofilms confined by a hydrogel as a model system, we show how a three-dimensional, living nematic can actively shape…
We develop a formal analogy between configurational stresses in two distinct physical systems, and study the flows that they induce when the configurations of interest include topological de- fects. The two systems in question are electrokinetic flows in a nematic fluid under an applied electrostatic field, and the motion of self propelling or active particles in a nematic matrix (a living liquid crystal). The mapping allows the extension, within certain limits, of existing results on trans- port in electrokinetic systems to self propelled transport. We study motion induced by a pair of point …
We study the effects of externally applied shear flow on a model of suspensions of motors and filaments, via the equations of active hydrodynamics [PRL {\bf 89} (2002) 058101; {\bf 92} (2004) 118101]. In the absence of shear, the orientationally ordered phase of {\it both} polar and apolar active particles is always unstable at zero-wavenumber. An imposed steady shear large enough to overcome the active stresses stabilises both apolar and moving polar phases. Our work is relevant to {\it in vitro} studies of active filaments, the reorientation of endothelial cells subject to shear flow and she…
We develop a description of defect loops in three-dimensional active nematics based on a multipole expansion of the far-field director and show how this leads to a self-dynamics dependent on the loop's geometric type. The dipole term leads to active stresses that generate a global self-propulsion for splay and bend loops. The quadrupole moment is non-zero only for non-planar loops and generates a net `active torque', such that defect loops are both self-motile and self-orienting. Our analysis identifies right- and left-handed twist loops as the only force and torque free geometries, suggesting…
Point-like motile topological defects control the universal dynamics of diverse two-dimensional active nematics ranging from shaken granular rods to cellular monolayers. A comparable understanding in higher dimensions has yet to emerge. We report the creation of three-dimensional active nematics by dispersing extensile microtubule bundles in a passive colloidal liquid crystal. Light-sheet microscopy reveals the millimeter-scale structure of active nematics with a single bundle resolution and the temporal evolution of the associated nematic director field. The dominant excitations of three-dime…
Cytoskeletal active nematics exhibit striking non-equilibrium dynamics that are powered by energy-consuming molecular motors. To gain insight into the structure and mechanics of these materials, we design programmable clusters in which kinesin motors are linked by a double-stranded DNA linker. The efficiency by which DNA-based clusters power active nematics depends on both the stepping dynamics of the kinesin motors and the chemical structure of the polymeric linker. Fluorescence anisotropy measurements reveal that the motor clusters, like filamentous microtubules, exhibit local nematic order.…
We examine the dynamics of a compressible active nematic liquid crystal on a frictional substrate. When frictional damping dominates over viscous dissipation, we eliminate flow in favor of active stresses to obtain a minimal dynamical model for the nematic order parameter, with elastic constants renormalized by activity. The renormalized elastic constants can become negative at large activity, leading to the selection of spatially inhomogeneous patterns via a mechanism analogous to that responsible for modulated phases arising at an equilibrium Lifshitz point. Tuning activity and the degree of…
From the mitotic spindle up to tissues and biofilms, many biological systems behave as active droplets, which often break symmetry and change shape spontaneously. Here, I show that active nematic droplets can experience a fingering instability. I consider an active fluid that acquires nematic order through anchoring at the droplet interface, and I predict its morphological stability in terms of three dimensionless parameters: the anchoring angle, the penetration length of nematic order compared to droplet size, and an active capillary number. Droplets with extensile (contractile) stresses and …
Suspensions of active agents with nematic interactions exhibit complex spatio-temporal dynamics such as mesoscale turbulence. Since the Reynolds number of microscopic flows is very small on the scale of individual agents, inertial effects are typically excluded in continuum theories of active nematic turbulence. Whether active stresses can collectively excite inertial flows is currently unclear. To address this question, we investigate a two-dimensional continuum theory for active nematic turbulence. In particular, we compare mesoscale turbulence with and without the effects of advective inert…
We examine the scaling with activity of the emergent length scales that control the nonequilibrium dynamics of an active nematic liquid crystal, using two popular hydrodynamic models that have been employed in previous studies. In both models we find that the chaotic spatio-temporal dynamics in the regime of fully developed active turbulence is controlled by a single active scale determined by the balance of active and elastic stresses, regardless of whether the active stress is extensile or contractile in nature. The observed scaling of the kinetic energy and enstropy with activity is consist…
We use a continuum model to examine the effect of activity on a phase separating mixture of an extensile active nematic and a passive fluid. We highlight the distinct role of previously considered interfacial active stresses and bulk active stresses that couple to liquid crystalline degrees of freedom. Interfacial active stresses can arrest phase separation, as previously demonstrated. Bulk extensile active stresses can additionally strongly suppress phase separation by sustained self-stirring of the fluid, substantially reducing the size of the coexistence region in the temperature/concentrat…
We describe the basic properties and consequences of introducing active stresses, with principal direction along the local director, in cholesteric liquid crystals. The helical ground state is found to be linearly unstable to extensile stresses, without threshold in the limit of infinite system size, whereas contractile stresses are hydrodynamically screened by the cholesteric elasticity to give a finite threshold. This is confirmed numerically and the non-linear consequences of instability, in both extensile and contractile cases, are studied. We also consider the stresses associated to defec…
Living active collectives have evolved with remarkable self-patterning ability to meet the physical and biological constraints for growth and survival. However, how complex multicellular patterns emerge from a single founder cell remains elusive. Here, by recourse to an agent-based model, we track the three-dimensional (3D) morphodynamics and cell orientational order of growing bacterial biofilms encased by agarose gels. Confined growth causes spatiotemporally heterogeneous stress buildup in the biofilm. High hydrostatic and low shear stresses at the core of the biofilm promote viscous-to-elas…
We analyze one of the simplest active suspensions with complex dynamics: a suspension of immotile "Extensor" particles that exert active extensile dipolar stresses on the fluid in which they are immersed. This is relevant to several experimental systems, such as recently studied tripartite rods that create extensile flows by consuming a chemical fuel. We first describe the system through a Doi-Onsager kinetic theory based on microscopic modeling. This theory captures the active stresses produced by the particles that can drive hydrodynamic instabilities, as well as the steric interactions of r…
Order and alignment are ubiquitous in growing colonies of rod-shaped bacteria due to the nematic properties of the constituent particles. These effects are the result of the active stresses generated by growth, passive mechanical interactions between cells, and flow-induced effects due to the shape of the confining container. However, how these contributing factors interact to give rise to the observed global alignment patterns remains elusive. Here, we study, in-silico, colonies of growing rod-shaped particles of different aspect ratios confined in channel-like geometries. A spatially resolve…
We report on the emergence of stable self-propelled bound defects in monolayers of active nematics, which form virtual full-integer topological defects in the form of vortices and asters. Through numerical simulations and analytical arguments, we identify the phase-space of the bound defect formation in active nematic monolayers. It is shown that an intricate synergy between the nature of active stresses and the flow-aligning behaviour of active particles can stabilise the motion of self-propelled positive half-integer defects into specific bound structures. Our findings uncover new complexiti…
Efficient nutrient mixing is crucial for the survival of bacterial colonies and other living systems. This raises the question of whether the optimization of mixing through the emergence of active turbulent motion in bacterial swarms played a role in the evolution of bacterial shapes. Here, to address this question, we solve the hydrodynamic equation for active nematics coupled with an advection-diffusion equation for the nutrients. The latter models a conserved activity field and mimics the conservation of nutrients in bacterial swarms. At the interface between active and passive nematic phas…
Starting from the stochastic equation for the density operator, we formulate the exact (instantaneous) stress tensor for interacting Brownian particles, whose average value agrees with expressions derived previously. We analyze the relation between the stress tensor and forces on external potentials, and observe that, out of equilibrium, particle currents give rise to extra forces. Next, we derive the stress tensor for a Landau-Ginzburg theory in non-equilibrium situations, finding an expression analogous to that of the exact microscopic stress tensor, and discuss the computation of out-of-equ…
Active materials are those in which individual, uncoordinated local stresses drive the material out of equilibrium on a global scale. Examples of such assemblies can be seen across scales from schools of fish to the cellular cytoskeleton and underpin many important biological processes. Synthetic experiments that recapitulate the essential features of such active systems have been the object of study for decades as their simple rules allow us to elucidate the physical underpinnings of collective motion. One system of particular interest has been active nematic liquid crystals (LCs). Because of…
We construct a hydrodynamic theory of noisy, apolar active smectics, in bulk suspension or on a substrate. Our predictions include: quasi-long-ranged smectic order in dimension d = 2, and long- ranged in d = 3, extending previously published results to all dynamical regimes; Kosterlitz-Thouless melting to an active nematic at high and low concentrations in d = 2; nonzero second-sound speed parallel to the layers; the suppression of giant number fluctuations by smectic elasticity; instability to spontaneous undulation and flow in bulk contractile smectics; a layer spacing instability, possibly …
Topological defects are at the root of the large-scale organization of liquid crystals. In two-dimensional active nematics, two classes of topological defects of charges $\pm 1/2$ are known to play a major role due to active stresses. Despite this importance, few analytical results have been obtained on the flow-field and active-stress patterns around active topological defects. Using the generic hydrodynamic theory of active systems, we investigate the flow and stress patterns around these topological defects in unbounded, two-dimensional active nematics. Under generic assumptions, we derive …
