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 demonstrate our purely data-driven approach on experiments of 2D unconfined active nematics of extensile microtubule bundles, as well as on data from numerical simulations of active nematics.
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 perform dynamical simulations of a two-dimensional active nematic fluid in coexistence with an isotropic fluid. Drops of active nematic become elongated, and an effective anchoring develops at the nematic-isotropic interface. The activity also causes an undulatory instability of the interface. This results in defects of positive topological charge being ejected into the nematic, leaving the interface with a diffuse negative charge. Quenching the active lyotropic fluid results in a steady state in which phase-separating domains are elongated and then torn apart by active stirring.
Dense, active systems show active turbulence, a state characterised by flow fields that are chaotic, with continually changing velocity jets and swirls. Here we review our current understanding of active turbulence. The development is primarily based on the theory and simulations of active liquid crystals, but with accompanying summaries of related literature.
We perform energy spectrum analysis of the active turbulence in 3D bulk active nematic using continuum numerical modelling. Specifically, we calculate the spectra of two main energy contributions---kinetic energy and nematic elastic energy---and combine this with the geometrical analysis of the nematic order and flow fields, based on direct defect tracking and calculation of autocorrelations. We show that the active nematic elastic energy is concentrated at scales corresponding to the effective defect-to-defect separation, scaling with activity as $\simζ^{0.5}$, whereas the kinetic energy is l…
We consider a phenomenological continuum theory for an extensile, overdamped active nematic liquid crystal, applicable in the dense regime. Constructed from general principles, the theory is universal, with parameters independent of any particular microscopic realization. We show that it exhibits a bend instability similar to that seen in active suspensions, that leads to the proliferation of defects. We find three distinct nonequilibrium steady states: a defect-ordered nematic in which $+\frac{1}{2}$ disclinations develop polar ordering, an undulating nematic state with no defects, and a turb…
I put forward a continuum theory for active nematic gels, defined as fluids or suspensions of orientable rodlike objects endowed with active dynamics, that is based on symmetry arguments and compatibility with thermodynamics. The starting point is our recent theory that models (passive) nematic liquid crystals as relaxing nematic elastomers. The interplay between viscoelastic response and active dynamics of the microscopic constituents is naturally taken into account. By contrast with standard theories, activity is not introduced as an additional term of the stress tensor, but it is added as a…
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…
Living materials at different length scales manifest active nematic features such as orientational order, nematic topological defects, and active nematic turbulence. Using numerical simulations we investigate the impact of fluid inertia on the collective pattern formation in active nematics. We show that an incremental increase in inertial effects due to reduced viscosity results in gradual melting of nematic order with an increase in topological defect density before a discontinuous transition to a vortex-condensate state. The emergent vortex-condensate state at low enough viscosities coincid…
We investigate similarities in the micro-structural dynamics between externally driven and actively driven nematics. Walls, lines of strong deformations in the director field, and topological defects are characteristic features of an active nematic. Similar structures form in driven passive nematics when there are inhomogeneities in imposed velocity gradients due to non-linear flow fields or geometrical constraints. Specifically, pressure driven flow of a tumbling passive nematic in an expanding-contracting channel produces walls and defects similar to those seen in active nematics. We also st…
We consider a phenomenological continuum model for an active nematic fluid and show a universal, model independent, instability which renders the homogeneous nematic state unstable to order fluctuations. Using numerical and analytic tools we show that, for low energy excitations in the vicinity of a critical point, this instability leads to a phase separation in which the ordered regions form bands with the direction of nematic order being perpendicular to the direction of density gradient. We argue that this mechanism of phase separation, which we term self-regulation, is a universal feature …
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…
The problem of low Reynolds number turbulence in active nematic fluids is theoretically addressed. Using numerical simulations I demonstrate that an incompressible turbulent flow, in two-dimensional active nematics, consists of an ensemble of vortices whose areas are exponentially distributed within a range of scales. Building on this evidence, I construct a mean-field theory of active turbulence by which several measurable quantities, including the spectral densities and the correlation functions, can be analytically calculated. Because of the profound connection between the flow geometry and…
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…
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…
The growing interest in active nematics and the emerging evidence of the relevance of topological defects in biology asks for reliable data analysis tools to identify, classify and track such defects in simulation and microscopy data. We here provide such tools and demonstrate on two examples, on an active turbulent state in an active nematodynamic model and on emerging nematic order in a multi-phase field model, the possibility to compare statistical data on defect velocities with experimental results. The considered tools, which are physics based and data driven, are compared with each other…
We propose a simple deterministic dynamic equation and reveal the mechanism of large-scale endless evolvement of spatial density inhomogeneity in active nematic. We determine the phase regions analytically. The interplay of density, magnitude of nematic order, and nematic director is crucial for the long-wave-length instability and the emergence of seemingly fluctuated collective motions. Ordered nematic domains can absorb particles, grow and divide endlessly. The present finding extends our understanding of the large-scale and seemingly fluctuated organization in active fluids.
We study the dynamics of active nematic films on a substrate driven by active flows with or without the incompressible constraint.Through simulations and theoretical analysis, we show that arch patterns are stable in the compressible case, whilst they become unstable under the incompressibility constraint. For compressible flows at high enough activity, stable arches organize themselves into a smectic-like pattern, which induce an associated global polar ordering of $+1/2$ nematic defects. By contrast, divergence-free flows give rise to a local nematic order of the $+1/2$ defects, consisting o…
Defect dynamics in a thin active nematic layer is studied by asymptotic matching of solutions in the defect core and the far field. The analysis is facilitated by the correspondence between the 2D nematic and complex scalar field models. Self-propulsion and topological interactions are identified as the primary drivers of the defect motion, surpassing the influence of both passive backflow and active flow induced by other defects.
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…
We numerically investigate how spatial variations of extensile or contractile active stress affect bulk active nematic systems in two and three dimensions. In the absence of defects, activity gradients drive flows which re-orient the nematic director field and thus act as an effective anchoring force. At high activity, defects are created and the system transitions into active turbulence, a chaotic flow state characterized by strong vorticity. We find that in two-dimensional (2D) systems active torques robustly align $+1/2$ defects parallel to activity gradients, with defect heads pointing tow…
Topological defects are distinctive signatures of liquid crystals. They profoundly affect the viscoelastic behavior of the fluid by constraining the orientational structure in a way that inevitably requires global changes not achievable with any set of local deformations. In active nematic liquid crystals topological defects not only dictate the global structure of the director, but also act as local sources of motion, behaving as self-propelled particles. In this article we present a detailed analytical and numerical study of the mechanics of topological defects in active nematic liquid cryst…
In this work we present the first systematic framework to sculpt active nematics systems, using optimal control theory and a hydrodynamic model of active nematics. We demonstrate the use of two different control fields, (1) applied vorticity and (2) activity strength, to shape the dynamics of an extensile active nematic that is confined to a disk. In the absence of control inputs, the system exhibits two attractors, clockwise and counterclockwise circulating states characterized by two co-rotating topological $+\frac{1}{2}$ defects. We specifically seek spatiotemporal inputs that switch the sy…
We consider a microscopic field theoretical approach for interacting active nematic particles. With only steric interactions the self-propulsion strength in such systems can lead to different collective behaviour, e.g., synchronized self-spinning and collective translation. The different behaviour results from the delicate interplay between internal nematic structure, particle shape deformation and particle-particle interaction. For intermediate active strength an asymmetric shape emerges and leads to chirality and self-spinning crystals. For larger active strength the shape is symmetric and t…
We derive a fully covariant theory of the hydrodynamics of nematic and polar active surfaces, subjected to internal and external forces and torques. We study the symmetries of polar and nematic surfaces and find that in addition to 5 different types of in-plane isotropic surfaces, polar and nematic surfaces can be classified into 5 polar, 2 pseudopolar, 5 nematic and 2 pseudonematic types of surfaces. We give examples of physical realisations of the different types of surfaces we have identified. We obtain expressions for the equilibrium tensions, moments, and external forces and torques actin…
We study a continuum model of an extensile active nematic to show that mesoscale turbulence develops in two stages: (i) ordered regions undergo an intrinsic hydrodynamic instability generating walls, lines of stong bend deformations, (ii) the walls relax by forming oppositely charged pairs of defects. Both creation and annihilation of defect pairs reinstate nematic regions which undergo further instabilities, leading to a dynamic steady state. We compare this with the development of active turbulence in a contractile active nematic.
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…
Using a microscopic model of interacting polar biofilaments and motor proteins, we characterize the phase diagram of both homogeneous and inhomogeneous states in terms of experimental parameters. The polarity of motor clusters is key in determining the organization of the filaments in homogeneous isotropic, polarized and nematic states, while motor-induced bundling yields spatially inhomogeneous structures.
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…
Swimming bacteria in passive nematics in the form of lyotropic liquid crystals are defined as a new class of active matter known as living liquid crystals in recent studies. It has also been shown that liquid crystal solutions are promising candidates for trapping and detecting bacteria. We ask the question, can a similar class of matter be designed for background nematics which are also active? Hence, we developed a minimal model for the mixture of polar particles in active nematics. It is found that the active nematics in such a mixture are highly sensitive to the presence of polar particles…
We study numerically the rheological properties of a slab of active gel close o the isotropic-nematic transition. The flow behavior shows strong dependence on sample size, boundary conditions, and on the bulk constitutive curve, which, on entering the nematic phase, acquires an activity-induced discontinuity at the origin. The precursor of this within the metastable isotropic phase for contractile systems ({\em e.g.,} actomyosin gels) gives a viscosity divergence; its counterpart for extensile ({\em e.g.,} {\em B. subtilis}) suspensions admits instead a shear-banded flow with zero apparent vis…
We use active nematohydrodynamics to study the flow of an active fluid in a 3D microchannel, finding a transition between active turbulence and regimes where there is a net flow along the channel. We show that the net flow is only possible if the active nematic is flow aligning and that - in agreement with experiments - the appearance of the net flow depends on the aspect ratio of the channel cross-section. We explain our results in terms of the when hydrodynamic screening due to the channel walls allows the emergence of vortex rolls across the channel.
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…
We use numerical modelling to study the flow patterns of an active nematic confined in a cylindrical capillary, considering both planar and homeotropic boundary conditions. We find that active flow emerges not only along the capillary axis but also within the plane of the capillary, where radial vortices are formed. If topological defects are imposed by the boundary conditions, they act as local pumps driving the flow. At higher activity we demonstrate escape of the active defects and flow into the third dimension, indicating the importance of dimensionality in active materials. We argue that …
We characterize two-dimensional (2D) microtubule-based active nematics driven by light-responsive kinesin motor clusters. We assess two constructs of optogenetic kinesin: opto-K401, a processive motor, and opto-K365, a non-processive motor. Measurements reveal an order of magnitude improvement in the contrast of nematic flow speeds between maximally- and minimally-illuminated states for opto-K365 motors. Focusing on opto-K365 nematics, we characterize both the steady-state flow and defect density as a function of applied light and examine the transient behavior between steady-states. The stead…
In nature interactions between biopolymers and motor proteins give rise to biologically essential emergent behaviours. Besides cytoskeleton mechanics, active nematics arise from such interactions. Here we present a study on 3D active nematics made of microtubules, kinesin motors and depleting agent. It shows a rich behaviour evolving from a nematically ordered space-filling distribution of microtubule bundles toward a flattened and contracted 2D ribbon that undergoes a wrinkling instability and subsequently transitions into a 3D active turbulent state. The wrinkle wavelength is independent of …
We show that "dry" active nematics, e.g. collections of shaken elongated granular particles, exhibit large-scale spatiotemporal chaos made of interacting dense, ordered, band-like structures in a parameter region including the linear onset of nematic order. These results are obtained from the study of the relatively simple and well-known (deterministic) hydrodynamic equations describing these systems in a dilute limit, and of a self-propelled particle Vicsek-like model for this class of active matter. In this last case, revisiting the status of the strong fluctuations and long-range correlatio…
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.
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…
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 …
We show that the dominant mode of alignment plays an important role in dry active nematics,leading to two dynamical sub-classes defined by the nature of the instability of the nematic bands that characterize, in these systems, the coexistence phase separating the isotropic and fluctuating nematic states. In addition to the well-known instability inducing long undulations along the band, another stronger instability leading to the break-up of the band in many transversal segments may arise. We elucidate the origin of this strong instability for a realistic model of self-propelled rods and deter…
