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, resulting in anisotropy of fluctuations. Our work shows that activity can drive either extensile or contractile stresses in tissue, depending on the relative strength of the contractility of the cortical cytoskeleton and tractions by cells on the extracellular matrix.
How can a collection of motile cells, each generating contractile nematic stresses in isolation, become an extensile nematic at the tissue-level? Understanding this seemingly contradictory experimental observation, which occurs irrespective of whether the tissue is in the liquid or solid states, is not only crucial to our understanding of diverse biological processes, but is also of fundamental interest to soft matter and many-body physics. Here, we resolve this cellular to tissue level disconnect in the small fluctuation regime by using analytical theories based on hydrodynamic descriptions o…
We use a computational phase-field model together with analytical analysis to study how inter-cellular active forces can mediate individual cell morphology and collective motion in a confluent cell monolayer. Contractile inter-cellular interactions lead to cell elongation, nematic ordering and active turbulence, characterised by motile topological defects. Extensile interactions result in frustration, and perpendicular cell orientations become more prevalent. Furthermore, we show that contractile behaviour can change to extensile behaviour if anisotropic fluctuations in cell shape are consider…
We report numerical results for the hydrodynamics of inhomogeneous lyotropic and extensile active nematic gels. By simulating the coupled Cahn-Hilliard, Navier-Stokes, and Beris-Edwards equation for the evolution of the composition, flow and orientational order of an active nematic, we ask whether composition variations are important to determine its emergent physics. As in active gels of uniform composition, we find that increasing either activity or nematic tendency (e.g., overall active matter concentration) triggers a transition between an isotropic passive phase and an active nematic one.…
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 appear to destroy long-range order. Here, we study these dynamics in experimental a…
In microtubule-based active nematics, motor-driven extensile motion of microtubule bundles powers chaotic large-scale dynamics. We quantify the interfilament sliding motion both in isolated bundles and in a dense active nematic. The extension speed of an isolated microtubule pair is comparable to the molecular motor stepping speed. In contrast, the net extension in dense 2D active nematics is significantly slower; the interfilament sliding speeds are widely distributed about the average and the filaments exhibit both contractile and extensile relative motion. These measurements highlight the c…
In active nematic liquid crystals activity is able to drive chaotic spatiotemporal flows referred to as active turbulence. Active turbulence has been characterized through theoretical and experimental work as a low Reynolds number phenomenon. We show that, in two-dimensions, the active forcing alone is able to trigger hydrodynamic turbulence leading to the coexistence of active and inertial turbulence. This type of flows develops for sufficiently active and extensile flow-aligning nematics. We observe that the combined effect of an extensile nematic and large values of the flow-aligning parame…
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…
We use numerical simulations to investigate the hydrodynamic behavior of the interface between nematic (N) and isotropic (I) phases of a confined active liquid crystal. At low activities, a stable interface with constant shape and velocity is observed separating the two phases. For nematics in homeotropic channels, the velocity of the interface at the NI transition increases from zero (i) linearly with the activity for contractile systems and (ii) quadratically for extensile ones. Interestingly, the nematic phase expands for contractile systems while it contracts for extensile ones, as a resul…
Liquid crystals inevitably possess topological defect excitations generated through boundary conditions, 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 topolo…
Solitary waves, dubbed "solitons", are special types of waves that propagate for an infinite distance under ideal conditions. These waves are ubiquitously found in nature such as typhoon or neuron signals. Yet, their artificial generation and the control of their propagation remain outstanding challenges in materials science owing to an insufficient understanding of the experimental conditions and theoretical aspects. Herein, a generic strategy for forming particle-like solitons and controlling their kinetics in nematic fluid media is reported. The key to the realisation of the generation of s…
Essentially all biology is active and dynamic. Biological entities autonomously sense, com- pute, and respond using energy-coupled ratchets that can produce force and do work. The cytoskeleton, along with its associated proteins and motors, is a canonical example of biological active matter, which is responsible for cargo transport, cell motility, division, and morphol- ogy. Prior work on cytoskeletal active matter systems showed either extensile or contractile dynamics. Here, we demonstrate a cytoskeletal system that can control the direction of the network dynamics to be either extensile, co…
We investigate the effects of extrinsic curvature on the turbulent behavior of a 2D active nematic confined to the surface of a cylinder. The surface of a cylinder has no intrinsic curvatrue and only extrinsic curvature. A nematic field reacts to the extrinsic curvature by trying to align with the lowest principle curvature, in this case parallel to the long axis of the cylinder. When nematics are sufficiently active, there is a proliferation of defects arising from a bend or splay instability depending on the nature of the active stress. The extrinsic curvature of the cylinder beaks the rotat…
We numerically investigate the morphology and disclination line dynamics of active nematic droplets in three dimensions. Although our model only incorporates the simplest possible form of achiral active stress, active nematic droplets display an unprecedented range of complex morphologies. For extensile activity finger-like protrusions grow at points where disclination lines intersect the droplet surface. For contractile activity, however, the activity field drives cup-shaped droplet invagination, run-and-tumble motion or the formation of surface wrinkles. This diversity of behaviour is explai…
Topological defects play a key role in two-dimensional active nematics, and a transient role in two-dimensional active polar fluids. In this paper, we study both the transient and long-time behavior of defects in two-dimensional active polar fluids in the limit of strong order and overdamped, compressible flow, and compare the defect dynamics with the corresponding active nematics model studied recently. One result is non-central interactions between defect pairs for active polar fluids, and by extending our analysis to allow orientation dynamics of defects, we find that the orientation of $+1…
Active nematics are a class of far-from-equilibrium materials characterized by local orientational order of force-generating, anisotropic constitutes. Traditional methods for predicting the dynamics of active nematics rely on hydrodynamic models, which accurately describe idealized flows and many of the steady-state properties, but do not capture certain detailed dynamics of experimental active nematics. We have developed a deep learning approach that uses a Convolutional Long-Short-Term-Memory (ConvLSTM) algorithm to automatically learn and forecast the dynamics of active nematics. We demonst…
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…
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…
Rod-shaped bacteria, such as Escherichia coli, commonly live forming mounded colonies. They initially grow two-dimensionally on a surface and finally achieve three-dimensional growth. While it was recently reported that three-dimensional growth is promoted by topological defects of winding number $+1/2$ in populations of motile bacteria, how cellular alignment plays a role in non-motile cases is largely unknown. Here, we investigate the relevance of topological defects in colony formation processes of non-motile E. coli populations, and found that both $\pm 1/2$ topological defects contribute …
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…
Growing tissue and bacterial colonies are active matter systems where cell divisions and cellular motion generate active stress. Although they operate in the non-equilibrium regime, these biological systems can form large-scale ordered structures such as nematically aligned cells, topological defects, and fingerings. Mechanical instabilities also play an essential role during growth by generating large structural folding. How active matter dynamics and mechanical instabilities together develop large-scale order in growing tissue is not well understood. Here, we use chain forming Bacillus subti…
Confining a liquid crystal imposes topological constraints on the orientational order, allowing global control of equilibrium systems by manipulation of anchoring boundary conditions. In this article, we investigate whether a similar strategy allows control of active liquid crystals. We study a hydrodynamic model of an extensile active nematic confined in containers, with different anchoring conditions that impose different net topological charges on the nematic director. We show that the dynamics are controlled by a complex interplay between topological defects in the director and their induc…
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…
In vitro epithelial monolayers exhibit a rich repertoire of collective dynamical behaviours according to microenvironment mechanical features. We investigate the statistical and dynamical properties of human bronchial epithelial cell (HBEC) flows in bulk, by analysing their large length-scale and long time-scale spatiotemporal dynamics. Activity-driven spontaneous collective flows consist of an ensemble of vortices randomly positioned in space. By analysing a large population of vortices, we show that their area follows an exponential law with a constant mean value and their rotational frequen…
We introduce the notion of cell division-induced activity and show that the cell division generates extensile forces and drives dynamical patterns in cell assemblies. Extending the hydrodynamic models of lyotropic active nematics we describe turbulent-like velocity fields that are generated by the cell division in a confluent monolayer of cells. We show that the experimentally measured flow field of dividing Madin-Darby Canine Kidney (MDCK) cells is reproduced by our modeling approach. Division-induced activity acts together with intrinsic activity of the cells in extensile and contractile cel…
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…
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…
Active nematic fluids confined in narrow channels generate spontaneous flows when the activity is sufficiently intense. Recently, it was shown that if the molecular anchoring at the channel walls is conflicting flows are initiated even in the zero activity limit. An analytical laminar velocity profile for this specific configuration was derived within a simplified nematohydrodynamic model in which the nematic order parameter is a fixed-magnitude unit vector n. In this study we explore systematically active flows in this confined geometry with a more general theoretical model that uses a second…
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 …
Recent studies have shown that packings of cells, both eukaryotic cellular tissues and growing or swarming bacterial colonies, can often be understood as active nematic fluids. A key property of volume-conserving active nematic model systems is chaotic self-mixing characterized by motile topological defects. However, for active nematics driven by growth rather than motility, less is understood about mixing and defect motion. Mixing could affect evolutionary outcomes in bacterial colonies by counteracting the tendency to spatially segregate into monoclonal sectors, which reduces the local genet…
Non-motile elongated bacteria confined in two-dimensional open micro-channels can exhibit collective motion and form dense monolayers with nematic order if the cells proliferate, i.e., grow and divide. Using soft molecular dynamics simulations of a system of rods interacting through short range mechanical forces, we study the effects of the cell growth rate, the cell aspect ratio and of the sliding friction on nematic ordering and on pressure fluctuations in confined environments. Our results indicate that rods with aspect ratio >3.0 reach quasi-perfect nematic states at low sliding friction. …
It is increasingly being realized that liquid-crystalline features can play an important role in the properties and dynamics of cell monolayers. Here, we present a cell-based model of cell layers, based on the phase-field formulation, that connects mechanical properties at the single cell level to large-scale nematic and hydrodynamic properties of the tissue. In particular, we present a minimal formulation that reproduces the well-known bend and splay hydrodynamic instabilities of the continuum nemato-hydrodynamic formulation of active matter, together with an analytical description of the ins…
Suspensions of swimming micro-organisms provide examples of coordinated active dynamics. That has stimulated the study of a phenomenological theory combining synchronization and polar order in active matter. Here, we consider another example inspired by the traction forces of migrating cells. The novelty, in this case, is the global force-free nature of the traction force field. Such a constraint is absent in the case where the vector field describes swimming speeds in micro-organisms suspensions. Cell traction is characterized by means of a complex tensor quantity, that generalizes the nemati…
