arXiv subjects

Julia M. Yeomans

Publications and source records attributed to Julia M. Yeomans.

At least 19 recordsLinked to original sources

arXiv2026

Competing routes to spontaneous flow in confined active nematics

Active nematics can spontaneously develop flow beyond a critical activity in confined geometries. We show analytically that the onset of flow is governed by two competing instabilities: a long-wavelength mode leading to unidirectional flow and a finite-wavelength instability producing transverse rolls. A reduced description reveals how activity, flow alignment, and nematic elasticity control the competition between these modes. We identify regimes in which the finite-wavelength instability has a lower critical activity than the long-wavelength instability, causing vortices to emerge before unidirectional flow. This establishes wavelength selection as an intrinsic feature of the onset of spontaneous flow in active nematics.

cond-mat.soft↗

arXiv2026

Active flows drive anchoring of nematics at rigid walls

Although confinement strongly influences flows in active materials, it remains unclear how active particles align at rigid boundaries when no thermodynamic anchoring is imposed. We address this question using continuum simulations of active nematics, together with analytical arguments based on a reduced near-wall description. In the flow-tumbling regime, extensile systems align parallel to the boundary, whereas contractile systems align perpendicular to it, consistent with active anchoring observed at active-passive interfaces. In the flow-aligning regime, the preferred orientation depends on the sign of activity and of the flow aligning parameter: either the shear-like flow generated near the wall selects the Leslie angle, or no unique alignment is established. These results provide a unified framework for activity-induced anchoring at rigid walls, demonstrating that boundary alignment in dense active matter can emerge solely from the interplay between self-generated flows and orientational dynamics.

cond-mat.soft↗

arXiv2026

Bridging Elastic and Active Turbulence

Remarkably, even under negligible inertia, the addition of microstructural agents can generate chaotic flow fields. Such behavior can arise in polymer solutions, leading to elastic turbulence, or from active, self-driven particles, which generate active turbulence. Here, we demonstrate a close and hitherto unrecognized connection between these two classes of turbulence. Specifically, we reveal that their continuum descriptions are analogous at the macroscopic level, such that polymeric fluids can be interpreted as a deformable analogue of contractile active matter. Moreover, our numerical results for Kolmogorov flow demonstrate that the transition into the well-known traveling arrowhead structures in elastic turbulence is marked by the emergence of $\pm 1/2$ topological defects, long recognized as a defining feature of active turbulence, in the polymer director field. Importantly, these coherent structures originate from a transverse instability driven by activity-like gradients generated by anisotropically stretched, contractile polymers. At sufficiently strong activity, the system undergoes a transition into a flow-suppressed state characterized by weak polymer stretching and ordering, a behavior that can be explained by analogy with the spontaneous-flow transition observed in channel-confined active nematics.

physics.flu-dyn↗

arXiv2025

Low-Pass Filtering of Active Turbulent Flows to Liquid Substrates

To study the impact of active systems on their surroundings, we introduce a model that couples an active nematic fluid to an isotropic substrate fluid via friction. We numerically show that as the active layer develops turbulence, the substrate inherits the chaotic behaviour, exhibiting a novel form of turbulence driven by locally generated stochastic forcing from the active layer. In particular, the short-length-scale flow structures in the active layer are filtered out, so the system behaves as a de facto low-pass filter. We derive analytically the transfer function between the two layers and use it to predict the large-q decay of the substrate energy spectrum, and to investigate how tensorial quantities, such as the strain rate and the active stresses, are transmitted between the active layer and the substrate. Our analysis agrees with recent experiments measuring velocity-velocity correlations in mixtures of active and passive microtubules, and it may have implications for traction force microscopy measurements in cellular layers.

cond-mat.soft↗

arXiv2025

From Equilibrium Multistability to Spatiotemporal Chaos in Channel Flows of Nematic Fluids

We investigate channel-confined, nematic liquid crystals using the Beris-Edwards model of nematohydrodynamics. Using strong homeotropic anchoring at the walls, we find multistability i.e. multiple coexisting states where the uniform nematic state coexists with states having spatially varying scalar nematic order and director fields. When a pressure gradient is applied, flows develop, and the inherent multistability of the system organizes a variety of complex dynamics. For low pressure gradients, steady flows are established, and the director fields that emerge from the multistable states at equilibrium correspond to Bowser and Dowser configurations similar to those reported in experiments. An increasing pressure-gradient destabilizes steady Bowser and Dowser flow states sequentially, leading to unsteady periodic and chaotic regimes featuring cyclical topological transitions, pulsating flows, advecting defects and spatiotemporal chaos. These findings demonstrate that modest variations in the scalar nematic order, as captured by the Beris-Edwards model, can qualitatively modify equilibrium structures and give rise to complex nonequilibrium behaviour in confined nematics-contrasting with the Ericksen-Leslie model, which assumes a constant scalar order parameter. Our key model predictions - multistability, periodically oscillating states and advecting defect-mediated turbulence can be experimentally investigated in pressure-driven channel flows of nematic fluids.

cond-mat.soft↗

arXiv2025

Why Extensile and Contractile Tissues Could be Hard to Tell Apart

Active nematic models explain the topological defects and flow patterns observed in epithelial tissues, but the nature of active stress-whether it is extensile or contractile, a key parameter of the theory-is not well established experimentally. Individual cells are contractile, yet tissue-level behavior often resembles extensile nematics. To address this discrepancy, we use a continuum theory with two-tensor order parameters that distinguishes cell shape from active stress. We show that correlating cell shape and flow, whether in coherent flows in channels, near topological defects, or at rigid boundaries, cannot unambiguously determine the type of active stress. Our results demonstrate that simultaneous measurements of stress and cell shape are essential to fully interpret experiments investigating the nature of the physical forces acting within epithelial cell layers.

cond-mat.soft↗

arXiv2025

Active sorting to boundaries in active nematic -- passive isotropic fluid mixtures

We use a two-fluid model to study a confined mixture of an active nematic fluid and a passive isotropic fluid. We find that an extensile active fluid preferentially accumulates at a boundary if the anchoring is planar, whereas its boundary concentration decreases for homeotropic anchoring. These tendencies are reversed if the active fluid is contractile. We argue that the sorting results from gradients in the nematic order, and show that the behaviour can be driven by either imposed boundary anchoring or spontaneous anchoring induced by active flows. Our results can be tested by experiments on microtubule-kinesin motor networks, and may be relevant to sorting to the boundary in cell colonies or cancer spheroids.

cond-mat.soft↗

arXiv2025

Channel flows of deformable nematics

We describe channel flows in a continuum model of deformable nematic particles. In a simple shear flow, deformability leads to a nonlinear coupling of strain rate and vorticity, and results in shape oscillations or flow alignment. The final steady state can depend on initial conditions, and we explain this behaviour by considering a phase space representation of the dynamics. In Poiseuille flow, particle deformability and nematic elasticity induce banding, where particles near the walls are aligned, and those near the centre of the channel oscillate in direction and shape. Our results show that particle deformability can lead to complex behaviour even in simple flows, suggesting new microfluidic experiments.

cond-mat.soft↗

arXiv2025

Spontaneous Hole Formation in Cell Monolayers Emerges from Collective Cell Motion

Although cell monolayers typically remain confluent, they can spontaneously develop persistent holes as a result of collective cellular motion. Recent studies on MDCK monolayers cultured on soft substrates have revealed that cells can align to create regions of local nematic order, and topological defects that generate localised mechanical stresses which can spontaneously trigger hole formation. To investigate this process, we develop a continuum multi-phase field model that incorporates internal dissipation and active dipolar forces that drive cell shape anisotropy. Our simulations show that reducing substrate friction enhances cell-cell velocity correlations. In this low-friction regime, topological defects give rise to spiral flow patterns that concentrate stress and can trigger hole formation. We further demonstrate that the number and stability of the holes, whether they close or persist, depends on both substrate friction and cellular activity. These findings underscore the critical role of collective cell dynamics in maintaining tissue integrity.

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↗

arXiv2024

Vertex model with internal dissipation enables sustained flows

Complex tissue flows in epithelia are driven by intra- and inter-cellular processes that generate, maintain, and coordinate mechanical forces. There has been growing evidence that cell shape anisotropy, manifested as nematic order, plays an important role in this process. Here we extend an active nematic vertex model by replacing substrate friction with internal viscous dissipation, dominant in epithelia not supported by a substrate or the extracellular matrix, which are found in many early-stage embryos. When coupled to cell shape anisotropy, the internal viscous dissipation allows for long-range velocity correlations and thus enables the spontaneous emergence of flows with a large degree of spatiotemporal organisation. We demonstrate sustained flow in epithelial sheets confined to a channel, providing a link between the cell-level vertex model of tissue dynamics and continuum active nematics, whose behaviour in a channel is theoretically understood and experimentally realisable. Our findings also show a simple mechanism that could account for collective cell migration correlated over distances large compared to the cell size, as observed during morphogenesis.

cond-mat.soft↗

arXiv2024

Self organisation of invasive breast cancer driven by the interplay of active and passive nematic dynamics

In invasive breast cancer, cell clusters of varying sizes and shapes are embedded in the fibrous extracellular matrix (ECM). Although the prevailing view attributes this structure to increasing disorder resulting from loss of function and dedifferentiation, our findings reveal that it arises through a process of active self-organization driven by cancer cell motility. Simulations and histological analyses of tumours from over 2,000 breast cancer patients reveal that motile, aligned cancer cells within clusters move as active nematic aggregates through the surrounding highly aligned ECM fibres, which form a confining, passive nematic phase. Cellular motion leads to cluster splitting and coalescence. The degree of cluster activity, combined with heterogeneity in cell motility, is reflected in specific scaling behaviours for cluster shape, size distribution, and the distance between cluster boundaries and nematic defects in ECM alignment. Increased activity estimates correlate with tumour progression and are associated with a poorer prognosis for patients.

physics.bio-ph↗

arXiv2024

Cell Sorting in an Active Nematic Vertex Model

We study a mixture of extensile and contractile cells using a vertex model extended to include active nematic stresses. The two cell populations phase separate over time. While phase separation strengthens monotonically with an increasing magnitude of contractile activity, the dependence on extensile activity is non-monotonic, so that sufficiently high values reduce the extent of sorting. We interpret this by showing that extensile activity renders the system motile, enabling cells to undergo neighbour exchanges. Contractile cells that come into contact as a result are then more likely to stay connected due to an effective attraction arising from contractile activity.

cond-mat.soft↗

arXiv2024

Phase Ordering in Binary Mixtures of Active Nematic Fluids

We use a continuum, two-fluid approach to study a mixture of two active nematic fluids. Even in the absence of thermodynamically-driven ordering, for mixtures of different activities we observe turbulent microphase separation, where domains form and disintegrate chaotically in an active turbulent background. This is a weak effect if there is no elastic nematic alignment between the two fluid components, but is greatly enhanced in the presence of an elastic alignment or substrate friction. We interpret the results in terms of relative flows between the two species which result from active anchoring at concentration gradients. Our results may have relevance in interpreting epithelial cell sorting and the dynamics of multi-species bacterial colonies.

cond-mat.soft↗

arXiv2024

Cell sorting by active forces in a phase-field model of cell monolayers

Cell sorting, the segregation of cells with different properties into distinct domains, is a key phenomenon in biological processes such as embryogenesis. We use a phase-field model of a confluent cell layer to study the role of activity in cell sorting. We find that a mixture of cells with extensile or contractile dipolar activity, and which are identical apart from their activity, quickly sort into small, elongated patches which then grow slowly in time. We interpret the sorting as driven by the different diffusivity of the extensile and contractile cells, mirroring the ordering of Brownian particles connected to different hot and cold thermostats. We check that the free energy is not changed by either partial or complete sorting, thus confirming that activity can be responsible for the ordering even in the absence of thermodynamic mechanisms.

physics.bio-ph↗

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↗

arXiv2023

Active nematics with deformable particles

The hydrodynamic theory of active nematics has been often used to describe the spatio-temporal dynamics of cell flows and motile topological defects within soft confluent tissues. Those theories, however, often rely on the assumption that tissues consist of cells with a fixed, anisotropic shape and do not resolve dynamical cell shape changes due to flow gradients. In this paper we extend the continuum theory of active nematics to include cell shape deformability. We find that circular cells in tissues must generate sufficient active stress to overcome an elastic barrier to deforming their shape in order to drive tissue-scale flows. Above this threshold the systems enter a dynamical steady-state with regions of elongated cells and strong flows coexisting with quiescent regions of isotropic cells.

cond-mat.soft↗