arXiv subjects

Mehrana R. Nejad

Publications and source records attributed to Mehrana R. Nejad.

16 recordsLinked to original sources

arXiv2026

Evaporation-Induced Pattern Formation and Wetting in Active Microtubule-Kinesin Droplets

Active networks composed of biopolymers and motor proteins provide versatile biomimetic systems that have advanced active matter physics and deepened our understanding of cytoskeletal dynamics and self-organization under diverse stimuli. In these systems, activity arises in aqueous solutions where motor proteins cross-link biopolymers and generate active stress driving the emergent network behavior. Here, we establish the active network in the form of a sessile, multi-component droplet on a substrate and investigate how evaporation influences its dynamics. We focus on how mass loss and compositional changes in the droplet reshape the behavior of the active suspension. We show that capillary and Marangoni flows drive the self-organization of microtubules into a distinctive radial arrangement within the droplet. The cross-linking ability of motor proteins gives rise to a striking non-monotonic wetting behavior, where the extensile stresses generated by the motor proteins strongly affect the characteristic timescale of the contact-line retracting and subsequent expansion. Using a combined experimental and theoretical approach, we demonstrate the crucial role of crosslinking in evaporating microtubule networks, and explain how active stresses together with evaporation-induced flows govern the dynamics of reconstituted microtubule systems and their wetting behavior. Evaporating droplets have recently attracted significant attention in the scientific community, and the findings of the setup presented in this study can have broad implications, ranging from self-organization and mechanical pattern formation in biological systems to questions about the origin of life.

cond-mat.soft↗

arXiv2025

Collective synchrony in confluent, pulsatile epithelia

Collective cell migration lies at the intersection of developmental biology and non-equilibrium physics, where active processes give rise to emergent patterns that are biologically relevant. Here, we investigate dilatational modes--cycles of expansion and contraction--in epithelial monolayers, and show that the divergence of the velocity field exhibits robust, large-scale temporal oscillations. These oscillatory patterns, reminiscent of excitable media and their biological analogs, emerge spontaneously from the coupled dynamics of actively pulsing cells. We find that the temporal persistence of these oscillations varies non-monotonically with cell density: synchrony initially increases with density, reaches a maximum at intermediate densities and is lost at higher values. This trend mirrors changes in the spatial correlation length of cell-cell interactions, and the density of topological defects in the system, suggesting a shared physical origin. We develop a continuum model in which a complex-valued Ginzburg-Landau-type field that governs the amplitude and phase of oscillations is coupled to local cell density. Simulations reproduce the observed behavior, revealing that local density adapts to phase patterns, reinforcing temporal coherence up to a critical density, and variations in the density of topological defects as a function of cell density. Extending our analysis to breast cancer cell lines with increasing invasiveness, we find that malignant cells exhibit longer phase persistence and fewer topological defects, suggesting a mechanistic link between temporal coherence and metastatic potential. Together, these results highlight the role of density-dependent synchrony dynamics as a fundamental, quantifiable mode of collective behavior in active epithelial matter, with implications for morphogenesis, cancer progression, and tissue diagnostics.

cond-mat.soft↗

arXiv2025

Thin active nematohydrodynamic layers: asymptotic theories and instabilities

Starting from a three-dimensional description of an active nematic layer, we employ an asymptotic theory to derive a series of low-dimensional continuum models that capture the coupled dynamics of flat and curved films, including variations in film thickness, shape deformations, internal velocity fields, and the dynamics of orientational order. Using this asymptotic theory, we investigate instabilities driven by activity in both the nematic and isotropic phases for cylindrical and flat films. In the flat case, we demonstrate that incorporating shape and thickness variations fundamentally alters the bend and splay nature of instabilities compared to conventional two dimensional nematic instabilities. In the isotropic phase, we find that both extensile and contractile activity can induce nematic order, in contrast with active nematics on fixed surfaces, where only extensile activity leads to ordering. For the case of curved geometries such as a cylindrical film, we reveal that thickness and shape instabilities are inherently coupled. In the isotropic phase, the emergence of nematic order triggers both thickness and shape instabilities. In the nematic phase, contractile activity induces thickness instabilities, which in turn drive geometric deformations. Our results highlight the crucial interplay between activity, thickness variations, and curvature, providing new insights into the behavior of active nematic films beyond the conventional two dimensional paradigm that has been studied to date.

cond-mat.soft↗

arXiv2025

Coarse-graining dense, deformable active particles

We coarse-grain a model of closely-packed ellipses that can vary their aspect ratio to derive continuum equations for materials comprising confluent deformable particles such as epithelial cell layers. We show that contractile nearest neighbour interactions between ellipses can lead to their elongation and nematic ordering. Adding flows resulting from active hydrodynamic stresses produced by the particles also affects the aspect ratio and can result in active turbulence. Our results, which agree well with multi-phase field simulations of deformable isotropic cells, provide a bridge between models which explicitly resolve cells and continuum theories of active matter.

cond-mat.soft↗

arXiv2025

Cellular dynamics emerging from turbulent flows steered by active filaments

We develop a continuum theory to describe the collective dynamics of deformable epithelial cells, using two tensor order parameters to distinguish the force-generating active filaments in the cells from their shape. The theory demonstrates how active flows create nematic domains of anisotropic cells, drive active turbulence, and create topological defects. We show that the filament flow-aligning parameter, $λ_{\tens{Q}}$, a rheological quantity that determines the response of the filaments to velocity gradients in the active flows, plays a significant, to date unappreciated, role in determining the pattern of extensional and compressional active flows. In a contractile cell layer, local flows are expected to align elongated cells perpendicular to the active filaments. However, with increasing $λ_{\tens{Q}}$, long-range correlations in the active turbulent flow field lead to extended regions where this alignment is parallel, as recently observed in experiments on confluent MDCK cell layers. Further, the two order parameter formalism allows us to distinguish defects in the filament director field, which contribute to the active driving, and those in the shape director field, measured in experiments, which are advected by the active flows. By considering the relative orientations of shape and filaments we are able to explain the surprising observation of defects moving towards their heads in contractile cell layers.

cond-mat.soft↗

arXiv2023

Spontaneous rotation of active droplets in two and three dimensions

We use numerical simulations and linear stability analysis to study active nematic droplets, in the regime where the passive phase is isotropic. We show that activity leads to the emergence of nematic order and of spontaneous rotation in both two and three dimensions. In 2D the rotation is caused by the formation of a chiral $+1$ defect at the center of the drop. With increasing activity the droplet deforms to an ellipse, and then to a rotating annulus. Growing droplets form extended active arms which loop around to produce holes. In 3D the rotation is due to a disclination which loops away from and back to the surface, defining the rotation axis. In the bulk the disclination loop ends at a skyrmion. Active extensile flows deform the droplet to an oblate ellipsoid, contractile flows elongate it along the rotation axis. We compare our results on rotation in two-dimensional droplets with experiments on microtubule and motor protein suspensions and find a critical radius $\sim 700 μm$ above which the spontaneous rotation gives way to active turbulence. Comparing the simulation parameters with experiments on epithelial cell colonies shows that the crossover radius for cell colonies could be as large as $2 mm$, in agreement with experiments.

cond-mat.soft↗

arXiv2023

Stress-shape misalignment in confluent cell layers

This study investigates the relationship between cell shape and cell-generated stresses in confluent cell layers. Using simultaneous measurements of cell shape orientation and cell-generated contractile forces in MDCK and LP-9 colonies, we report the emergence of correlated, dynamic domains in which misalignment between the directors defined by cell shape and by contractile forces reaches up to 90$^o$, effectively creating extensile domains in a monolayer of contractile cells. To understand this misalignment, we develop a continuum model that decouples the orientation of cell-generated active forces from the orientation of the cell shapes. This challenges the prevailing understanding that cells throughout a tissue create either contractile or extensile forces, and the validity of the usual active nematic models of cell motility where active forces are strictly slaved to cell shape orientation.

physics.bio-ph↗

arXiv2022

Collective rotational motion of freely-expanding T84 epithelial cell colonies

Coordinated rotational motion is an intriguing, yet still elusive mode of collective cell migration, which is relevant in pathological and morphogenetic processes. Most of the studies on this topic have been carried out on confined epithelial cells. The driver of collective rotation in such conditions has not been clearly elucidated, although it has been speculated that spatial confinement can play an essential role in triggering cell rotation. Here, we study the growth of epithelial cell colonies freely expanding (i.e., with no physical constraints) on the surface of cell culture plates, a case which has received scarce attention in the literature. We find that coordinated cell rotation spontaneously occurs in cell clusters in the free growth regime, thus implying that cell confinement is not necessary to elicit collective rotation as previously suggested. The collective rotation was size and shape dependent: a highly coordinated disk-like rotation was found in small cell clusters with a round shape, while collective rotation was suppressed in large irregular cell clusters generated by merging of different clusters in the course of their growth. The angular motion was persistent in the same direction, although clockwise and anticlockwise rotations were equally likely to occur among different cell clusters. Radial cell velocity was low as compared to the angular velocity. A clear difference in morphology was observed between cells at the periphery and the ones in the core of the clusters, the former being more elongated and spread out as compared to the latter. Overall, our results provide the first quantitative and systematic evidence that coordinated cell rotation does not require a spatial confinement and occurs spontaneously in freely expanding epithelial cell colonies.

cond-mat.soft↗

arXiv2022

Geometrical control of interface patterning underlies active matter invasion

Interaction between active materials and the boundaries of geometrical confinement is key to many emergent phenomena in active systems. For living active matter consisting of animal cells or motile bacteria, the confinement boundary is often a deformable interface, and it has been unclear how activity-induced interface dynamics might lead to morphogenesis and pattern formation. Here we studied the evolution of bacterial active matter confined by a deformable boundary. We discovered that an ordered morphological pattern emerged at the interface characterized by periodically-spaced interfacial protrusions; behind the interfacial protrusions, bacterial swimmers self-organized into multicellular clusters displaying +1/2 nematic defects. Subsequently, a hierarchical sequence of transitions from interfacial protrusions to creeping branches allowed the bacterial active drop to rapidly invade surrounding space with a striking self-similar branch pattern. We found that this interface patterning is controlled by the local curvature of the interface, a phenomenon we denote as collective curvature sensing. Using a continuum active model, we revealed that the collective curvature sensing arises from enhanced active stresses near high-curvature regions, with the active length-scale setting the characteristic distance between the interfacial protrusions. Our findings reveal a protrusion-to-branch transition as a novel mode of active matter invasion and suggest a new strategy to engineer pattern formation of active materials.

cond-mat.soft↗

arXiv2022

Active extensile stress promotes 3D director orientations and flows

We use numerical simulations and linear stability analysis to study an active nematic layer where the director is allowed to point out of the plane. Our results highlight the difference between extensile and contractile systems. Contractile stress suppresses the flows perpendicular to the layer and favours in-plane orientations of the director. By contrast, extensile stress promotes instabilities that can turn the director out of the plane, leaving behind a population of distinct, in-plane regions that continually elongate and divide. This supports extensile forces as a mechanism for the initial stages of layer formation in living systems, and we show that a planar drop with extensile (contractile) activity grows into three dimensions (remains in two dimensions). The results also explain the propensity of disclination lines in three dimensional active nematics to be of twist-type in extensile or wedge-type in contractile materials.

cond-mat.soft↗

arXiv2021

Quadrupolar active stress induces exotic phases of defect motion in active nematics

A wide range of living and artificial active matter exists in close contact with substrates and under strong confinement, where in addition to dipolar active stresses, quadrupolar active stresses can become important. Here, we numerically investigate the impact of quadrupolar non-equilibrium stresses on the emergent patterns of self-organisation in non-momentum conserving active nematics. Our results reveal that beyond having stabilising effects, the quadrupolar active forces can induce various modes of topological defect motion in active nematics. In particular, we find the emergence of both polar and nematic ordering of the defects, as well as new phases of self-organisation that comprise topological defect chains and topological defect asters. The results contribute to further understanding of emergent patterns of collective motion and non-equilibrium self-organisation in active matter.

cond-mat.soft↗

arXiv2020

Role of friction in multidefect ordering

We use continuum simulations to study the impact of friction on the ordering of defects in an active nematic. Even in a frictionless system, +1/2 defects tend to align side-by-side and orient antiparallel reflecting their propensity to form, and circulate with, flow vortices. Increasing friction enhances the effectiveness of the defect-defect interactions, and defects form dynamically evolving, large scale, positionally and orientationally-ordered structures which can be explained as a competition between hexagonal packing, preferred by the -1/2 defects, and rectangular packing preferred by the +1/2 defects.

cond-mat.soft↗

arXiv2020

The role of dimensionality and geometry in quench-induced nonequilibrium forces

We present an analytical formalism, supported by numerical simulations, for studying forces that act on curved walls following temperature quenches of the surrounding ideal Brownian fluid. We show that, for curved surfaces, the post-quench forces initially evolve rapidly to an extremal value, whereafter they approach their steady state value algebraically in time. In contrast to the previously-studied case of flat boundaries (lines or planes), the algebraic decay for the curved geometries depends on the dimension of the system. Specifically, the steady-state values of the force are approached in time as $t^{-d/2}$ in d-dimensional spherical (curved) geometries. For systems consisting of concentric circles or spheres, the exponent does not change for the force on the outer circle or sphere. However, the force exerted on the inner circle or sphere experiences an overshoot and, as a result, does not evolve towards the steady state in a simple algebraic manner. The extremal value of the force also depends on the dimension of the system, and originates from the curved boundaries and the fact that particles inside a sphere or circle are locally more confined, and diffuse less freely than particles outside the circle or sphere.

cond-mat.stat-mech↗

arXiv2019

Activity induced nematic order in isotropic liquid crystals

We use linear stability analysis to show that an isotropic phase of elongated particles with dipolar flow fields can develop nematic order as a result of their activity. We argue that ordering is favoured if the particles are flow-aligning and is strongest if the wavevector of the order perturbation is neither parallel nor perpendicular to the nematic director. Numerical solutions of the hydrodynamic equations of motion of an active nematic confirm the results. The instability is contrasted to the well-known hydrodynamic instability of an ordered active nematic.

cond-mat.soft↗

arXiv2019

Interplay of quenching temperature and drift in Brownian dynamics

We investigate the non-equilibrium evolution of ideal Brownian particles confined between two walls, following simultaneous quenches of the temperature and a constant external force. We compute (analytically and in numeric simulations) the post-quench dynamics of the density and the pressure exerted by the particles on the two walls perpendicular to the drift force. For identical walls, symmetry breaking associated with the drift gives rise to unequal particle densities and pressures on the two walls. While the pressure on one wall increases monotonically after the quench, on the other wall, depletion causes a non-monotonic dynamics with an overshooting at finite times, before the long-term steady-state value is reached. For walls immersed in a Brownian gas, the effective interaction force changes sign from repulsive at short times to attractive at late times. These findings have potential applications in various soft matter systems or fluids with charged Brownian particles, as well as carrier dynamics in semiconducting structures.

cond-mat.stat-mech↗

arXiv2018

Self-generated gradients stabilize the hydrodynamic instabilities in active suspensions

Ordered phases emerged in active suspensions of polar swimmers are under long-wavelength hydrodynamic mediated instabilities. In this letter, we show that chemical molecules dissolved in aqueous suspensions, as an unavoidable part of most wet active systems, can mediate long-range interactions and subsequently stabilize the ordered phases. Chemoattractant in living suspensions and dissolved molecules producing phoretic forces in synthesized Janus suspensions are reminiscent of such molecules. Communication between swimmers through the gradients of such chemicals generated by individual swimmers, is the foundation of this stabilization mechanism. To classify the stable states of such active systems, we investigate the detailed phase diagrams for two classes of systems with momentum conserving and non-conserving dynamics. Our linear stability analysis shows how the stabilization mechanism can work for swimmers with different dynamical properties, e.g., pushers or pullers and with various static characteristics, e.g., spherical, oblate or prolate geometries.

cond-mat.stat-mech↗