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Motivated by microfluidic applications, we investigate drag reduction in laminar pressure-driven flows in channels with streamwise-periodic superhydrophobic surfaces (SHSs) that are contaminated with soluble surfactant. We develop a model in the long-wave and weak-diffusion limit, where the streamwise SHS period is large compared to the channel height and the P\'eclet number is large. Employing asymptotic and numerical techniques, we determine the drag due to surfactant in terms of the relative strength of advection, diffusion, Marangoni effects and bulk-surface partitioning and exchange. In scenarios with strong bulk-surface exchange, the drag reduction exhibits a complex dependence on the thickness of the bulk-concentration boundary layer and surfactant strength. Strong Marangoni effects immobilise the interface through a linear surfactant distribution, whereas weak Marangoni effects yield a quasi-stagnant cap. The quasi-stagnant cap distribution has an intricate asymptotic structure with an upstream slip region followed by intermediate inner regions and a quasi-stagnant region that is mediated by weak bulk diffusion. The quasi-stagnant region differs from the immobile region of a classical stagnant cap, observed for instance in surfactant-laden air bubbles in water, by displaying weak slip. As bulk-surface exchange weakens, the bulk and interface decouple: the surfactant distribution is linear when the surfactant is strong, whilst it forms a classical stagnant cap when the surfactant is weak. The asymptotic solutions offer closed-form predictions of drag reduction across much of the parameter space, providing practical utility and enhancing understanding of surfactant dynamics in flows over SHSs.

Airway constriction and blockage in obstructive lung diseases cause ventilation heterogeneity and create barriers to effective drug deposition. Established computational particle-deposition models have not accounted for these impacts of disease. We present a new particle-deposition model that calculates ventilation based on the resistance of each airway, such that ventilation responds to airway constriction. The model incorporates distal airway constrictions representative of cystic fibrosis, allowing us to investigate the resulting impact on patterns of deposition. Unlike previous models, our model predicts how constrictions affect deposition in airways throughout the lungs, not just in the constricted airways. Deposition is reduced in airways directly distal and proximal to constrictions. When constrictions are clustered together, central-airways deposition can increase significantly in regions away from constrictions, but distal-airways deposition in those regions remains largely unchanged. We use our model to calculate lung clearance index (LCI), a clinical measure of ventilation heterogeneity, after applying constrictions of varying severities in one lobe. We find an increase in LCI coinciding with significantly reduced deposition in the affected lobe. Our results show how the model provides a framework for development of computational tools that capture the impacts of airway disease, which could significantly affect predictions of regional dosing., Comment: 24 pages, 9 figures

We investigate the viscoelastic relaxation to equilibrium of a disordered planar epithelium described using the cell vertex model. In its standard form, the model is formulated as coupled evolution equations for the locations of vertices of confluent polygonal cells. Exploiting the model's gradient-flow structure, we use singular-value decomposition to project modes of deformation of vertices onto modes of deformation of cells. We show how eigenmodes of discrete Laplacian operators (specified by constitutive assumptions related to dissipation and mechanical energy) provide a spatial basis for evolving fields, and demonstrate how the operators can incorporate approximations of conventional spatial derivatives. We relate the spectrum of relaxation times to the eigenvalues of the Laplacians, modified by corrections that account for the fact that the cell network (and therefore the Laplacians) evolve during relaxation to an equilibrium prestressed state, providing the monolayer with geometric stiffness. While dilational modes of the Laplacians capture rapid relaxation in some circumstances, showing diffusive dynamics, geometric stiffness is typically a dominant source of monolayer rigidity, as we illustrate for monolayers exposed to unsteady stretching deformations., Comment: 8 figures

This report relates to a study group hosted by the EPSRC funded network, Integrating data-driven BIOphysical models into REspiratory MEdicine (BIOREME), and supported by The Insigneo Institute and The Knowledge Transfer Network. The BIOREME network hosts events, including this study group, to bring together multi-disciplinary researchers, clinicians, companies and charities to catalyse research in the applications of mathematical modelling for respiratory medicine. The goal of this study group was to provide an interface between companies, clinicians, and mathematicians to develop mathematical tools to the problems presented. The study group was held at The University of Sheffield on the 17 - 20 April 2023 and was attended by 24 researchers from 13 different institutions. This report relates to a challenge presented by Arete Medical Technologies relating to impulse oscillometry (IOS), whereby a short pressure oscillation is imposed at a person's mouth during normal breathing, usually by a loudspeaker. The resulting pressure and flow rate changes can be used to the impedance of the airways, which in turn can provide proxy measurements for (patho)physiological changes in the small airways. Disentangling the signal so that airway mechanics can be measured accurately (and device properties/environmental effects can be accounted for) remains an open challenge that has the potential to significantly improve the device and its translation to clinic. In this report, several approaches to this problem, and the wider problem of interpreting oscillometry resuts are explored., Comment: 17 pages, 9 figures, Study group report

During cell division, the mitotic spindle moves dynamically through the cell to position the chromosomes and determine the ultimate spatial position of the two daughter cells. These movements have been attributed to the action of cortical force generators which pull on the astral microtubules to position the spindle, as well as pushing events by these same microtubules against the cell cortex and plasma membrane. Attachment and detachment of cortical force generators working antagonistically against centring forces of microtubules have been modelled previously (Grill et al. 2005, Phys. Rev. Lett. 94:108104) via stochastic simulations and mean-field Fokker-Planck equations (describing random motion of force generators) to predict oscillations of a spindle pole in one spatial dimension. Using systematic asymptotic methods, we reduce the Fokker-Planck system to a set of ordinary differential equations (ODEs), consistent with a set proposed by Grill et al., which can provide accurate predictions of the conditions for the Fokker-Planck system to exhibit oscillations. In the limit of small restoring forces, we derive an algebraic prediction of the amplitude of spindle-pole oscillations and demonstrate the relaxation structure of nonlinear oscillations. We also show how noise-induced oscillations can arise in stochastic simulations for conditions in which the mean-field Fokker-Planck system predicts stability, but for which the period can be estimated directly by the ODE model and the amplitude by a related stochastic differential equation that incorporates random binding kinetics., Comment: 7 figures, 2 tables, 2 online resources (movies)

Experiments have shown that surfactant introduced to a liquid-filled maze can find the solution path. We reveal how the maze-solving dynamics arise from interactions between the added surfactant and endogenous surfactant present at the liquid surface. We simulate the dynamics using a nonlinear model solved with a discrete mimetic scheme on a graph. Endogenous surfactant transforms local spreading into a non-local problem with an omniscient view of the maze geometry, key to the maze-solving dynamics. Our results offer insight into surfactant-driven transport in complex networks such as lung airways., Comment: 5 pages, 4 figures

Recognising that surfactants can impede the drag reduction resulting from superhydrophobic surfaces (SHSs), we investigate the impact of spatio-temporal fluctuations in surfactant concentration on the drag-reduction properties of SHSs. We model the unsteady transport of soluble surfactant in a channel flow bounded by two SHSs. The flow is laminar, pressure-driven, and the SHSs are periodic in the streamwise and spanwise directions. We assume that the channel length is much longer than the streamwise period, the streamwise period is much longer than the channel height and spanwise period, and bulk diffusion is sufficiently strong for cross-channel concentration gradients to be small. By combining long-wave and homogenisation theories, we derive an unsteady advection-diffusion equation for surfactant-flux transport over the length of the channel, which is coupled to a quasi-steady advection-diffusion equation for surfactant transport over individual plastrons. As diffusion over the length of the channel is typically small, the surfactant flux is governed by a nonlinear wave equation. In the fundamental case of the transport of a bolus of surfactant, we predict its propagation speed and describe its nonlinear evolution via interaction with the SHS. The propagation speed can fall below the average streamwise velocity as the surfactant adsorbs and rigidifies the plastrons. Smaller concentrations of surfactant are advected faster than larger ones, so that wave-steepening effects can lead to shock formation in the surfactant-flux distribution. Our asymptotic results reveal how unsteady surfactant transport can affect the spatio-temporal evolution of the slip velocity, drag reduction and effective slip length in SHS channels.

Successive drops of coloured ink mixed with surfactant are deposited onto a thin film of water to create marbling patterns in the Japanese art technique of Suminagashi. To understand the physics behind this and other applications where surfactant transports adsorbed passive matter at gas-liquid interfaces, we investigate the Lagrangian trajectories of material particles on the surface of a thin film of a confined viscous liquid under Marangoni-driven spreading by an insoluble surfactant. We study a model problem in which several deposits of exogenous surfactant simultaneously spread on a bounded rectangular surface containing a pre-existing endogenous surfactant. We derive Eulerian and Lagrangian formulations of the equations governing the Marangoni-driven surface flow. Both descriptions show how confinement can induce drift and flow reversal during spreading. The Lagrangian formulation captures trajectories without the need to calculate surfactant concentrations; however, concentrations can still be inferred from the Jacobian of the map from initial to current particle position. We explore a link between thin-film surfactant dynamics and optimal transport theory to find the approximate equilibrium locations of material particles for any given initial condition by solving a Monge-Amp\`ere equation. We find that, as the endogenous surfactant concentration $\delta$ vanishes, the equilibrium shapes of deposits using the Monge-Amp\`ere approximation approach polygons with corners curving in a self-similar manner over lengths scaling as $\delta^{1/2}$. We explore how Suminagashi patterns may be produced by using computationally efficient successive solutions of the Monge-Amp\`ere equation., Comment: 26 pages without appendices, 10 figures in main text, 2 figures in appendices, 7 figures in supplement

To assess how the presence of surfactant in lung airways alters the flow of mucus that leads to plug formation and airway closure, we investigate the effect of insoluble surfactant on the instability of a viscoplastic liquid coating the interior of a cylindrical tube. Evolution equations for the layer thickness using thin-film and long-wave approximations are derived that incorporate yield-stress effects and capillary and Marangoni forces. Using numerical simulations and asymptotic analysis of the thin-film system, we quantify how the presence of surfactant slows growth of the Rayleigh-Plateau instability, increases the size of initial perturbation required to trigger instability and decreases the final peak height of the layer. When the surfactant strength is large, the thin-film dynamics coincide with the dynamics of a surfactant-free layer but with time slowed by a factor of four and the capillary Bingham number, a parameter proportional to the yield stress, exactly doubled. By solving the long-wave equations numerically, we quantify how increasing surfactant strength can increase the critical layer thickness for plug formation to occur and delay plugging. The previously established effect of the yield stress in suppressing plug formation [Shemilt et al., J. Fluid Mech., 2022, vol. 944, A22] is shown to be amplified by introducing surfactant. We discuss the implications of these results for understanding the impact of surfactant deficiency and increased mucus yield stress in obstructive lung diseases., Comment: 35 pages, 11 figures

Superhydrophobic surfaces (SHSs) can reduce the friction drag in turbulent flows. In the laminar regime, it has been shown that trace amounts of surfactant can negate this drag reduction, at times rendering these surfaces no better than solid walls (Peaudecerf et al., Proc. Natl. Acad. Sci. USA 114(28), 7254-9, 2017). However, surfactant effects on the drag-reducing properties of SHSs have not yet been studied under turbulent flow conditions, where predicting the effects of surfactant in direct numerical simulations remains expensive by today's standards. We present a model for turbulent flow inclusive of surfactant, in either a channel or boundary-layer configuration, over long but finite-length streamwise ridges that are periodic in the spanwise direction, with period $P$ and gas fraction $\varphi$. We adopt a technique based on a shifted log law to acquire an expression for the drag reduction. The average streamwise and spanwise slip lengths are derived by introducing a local laminar model within the viscous sublayer, whereby the effect of surfactant is modelled by modifying the average streamwise and spanwise slip lengths. Our model agrees with available laboratory experimental data from the literature when conditions are clean (surfactant-free), or when there are low surfactant levels. However, we find an appreciable drag increase for larger background surfactant concentrations that are characteristic of turbulent flows over SHSs for marine applications.

While superhydrophobic surfaces (SHSs) show promise for drag reduction applications, their performance can be compromised by traces of surfactant, which generate Marangoni stresses that increase drag. This question is addressed for soluble surfactant in a three-dimensional laminar channel flow, with periodic SHSs on both walls. We assume that diffusion is sufficiently strong for cross-channel concentration gradients to be small. Exploiting a long-wave theory that accounts for a rapid transverse Marangoni-driven flow, we derive a one-dimensional model for surfactant evolution, which allows us to predict the drag reduction across the parameter space. The system exhibits multiple regimes, involving competition between Marangoni effects, bulk and interfacial diffusion, advection and shear dispersion. We map out asymptotic regions in the high-dimensional parameter space, deriving approximations of the drag reduction in each region and comparing them to numerical simulations. Our atlas of maps provides a comprehensive analytical guide for designing surfactant-contaminated channels with SHSs, to maximise the drag reduction in applications.

The vertex model is widely used to simulate the mechanical properties of confluent epithelia and other multicellular tissues. This inherently discrete framework allows a Cauchy stress to be attributed to each cell, and its symmetric component has been widely reported, at least for planar monolayers. Here we consider the stress attributed to the neighbourhood of each tricellular junction, evaluating in particular its leading-order antisymmetric component and the associated couple stresses, which characterise the degree to which individual cells experience (and resist) in-plane bending deformations. We develop discrete potential theory for localised monolayers having disordered internal structure and use this to derive the analogues of Airy and Mindlin stress functions. These scalar potentials typically have broad-banded spectra, highlighting the contributions of small-scale defects and boundary-layers to global stress patterns. An affine approximation attributes couple stresses to pressure differences between cells sharing a trijunction, but simulations indicate an additional role for non-affine deformations., Comment: 8 figures, 1 table

One mechanism for airway closure in the lung is the surface-tension-driven instability of the mucus layer which lines the airway wall. We study the instability of an axisymmetric layer of viscoplastic Bingham liquid coating the interior of a rigid tube, which is a simple model for an airway that takes into account the yield stress of mucus. An evolution equation for the thickness of the liquid layer is derived using long-wave theory, from which we also derive a simpler thin-film evolution equation. In the thin-film case, we show that two branches of marginally-yielded static solutions of the evolution equation can be used to both predict the size of initial perturbation required to trigger instability and quantify how increasing the capillary Bingham number (a parameter measuring yield stress relative to surface tension) reduces the final deformation of the layer. Using numerical solutions of the long-wave evolution equation, we quantify how the critical layer thickness required to form a liquid plug in the tube increases as the capillary Bingham number is increased. We discuss the significance of these findings for modelling airway closure in obstructive conditions such as cystic fibrosis, where the mucus layer is often thicker and has a higher yield stress., Comment: 36 pages, 12 figures; revised version after response from J. Fluid Mech. reviewers and relevant revisions made

We investigate the transport of a solute past isolated sinks in a bounded domain when advection is dominant over diffusion, evaluating the effectiveness of homogenization approximations when sinks are distributed uniformly randomly in space. Corrections to such approximations can be non-local, non-smooth and non-Gaussian, depending on the physical parameters (a P\'eclet number Pe, assumed large, and a Damk\"ohler number Da) and the compactness of the sinks. In one spatial dimension, solute distributions develop a staircase structure for large Pe, with corrections being better described with credible intervals than with traditional moments. In two and three dimensions, solute distributions are near-singular at each sink (and regularized by sink size), but their moments can be smooth as a result of ensemble averaging over variable sink locations. We approximate corrections to a homogenization approximation using a moment-expansion method, replacing the Green's function by its free-space form, and test predictions against simulation. We show how, in two or three dimensions, the leading-order impact of disorder can be captured in a homogenization approximation for the ensemble mean concentration through a modification to Da that grows with diminishing sink size., Comment: 5 figures

We present theoretical and numerical results that demonstrate the sensitivity of the shape of a static meniscus in a rectangular channel to localised geometric perturbations in the form of narrow ridges and grooves imposed on the channel walls. The Young--Laplace equation is solved for a gas/liquid interface with fixed contact angle using computations, analytical arguments and semi-analytical solutions of a linearised model for small-amplitude perturbations. We find that the local deformation of the meniscus's contact line near a ridge or groove is strongly dependent on the shape of the perturbation. In particular, small-amplitude perturbations that change the channel volume induce a change in the pressure difference across the meniscus, resulting in long-range curvature of its contact line. We derive an explicit expression for this induced pressure difference directly in terms of the boundary data. We show how contact lines can be engineered to assume prescribed patterns using suitable combinations of ridges and grooves., Comment: 25 pages, 10 figures

We model the advective Marangoni spreading of insoluble surfactant at the free surface of a viscous fluid that is confined within a two-dimensional rectangular cavity. Interfacial deflections are assumed small, with contact lines pinned to the walls of the cavity, and inertia is neglected. Linearizing the surfactant transport equation about the equilibrium state allows a modal decomposition of the dynamics, with eigenvalues corresponding to decay rates of perturbations. Computation of the family of mutually orthogonal two-dimensional eigenfunctions reveals singular flow structures near each contact line, resulting in spatially oscillatory patterns of wall shear stress and a pressure field that diverges logarithmically. These singularities at a stationary contact line are associated with dynamic compression of the surfactant monolayer; we show how they can be regularized by weak surface diffusion. Their existence highlights the need for careful treatment in computations of unsteady advection-dominated surfactant transport in confined domains., Comment: 23 pages, 6 figures

The vertex model is a popular framework for modelling tightly packed biological cells, such as confluent epithelia. Cells are described by convex polygons tiling the plane and their equilibrium is found by minimizing a global mechanical energy, with vertex locations treated as degrees of freedom. Drawing on analogies with granular materials, we describe the force network for a localized monolayer and derive the corresponding discrete Airy stress function, expressed for each $N$-sided cell as $N$ scalars defined over kites covering the cell. We show how a torque balance (commonly overlooked in implementations of the vertex model) requires each internal vertex to lie at the orthocentre of the triangle formed by neighbouring edge centroids. Torque balance also places a geometric constraint on the stress in the neighbourhood of cellular trijunctions, and requires cell edges to be orthogonal to the links of a dual network that connect neighbouring cell centres and thereby triangulate the monolayer. We show how the Airy stress function depends on cell shape when a standard energy functional is adopted, and discuss implications for computational implementations of the model.

The primary exchange units in the human placenta are terminal villi, in which fetal capillary networks are surrounded by a thin layer of villous tissue, separating fetal from maternal blood. To understand how the complex spatial structure of villi influences their function, we use an image-based theoretical model to study the effect of tissue metabolism on the transport of solutes from maternal blood into the fetal circulation. For solute that is taken up under first-order kinetics, we show that the transition between flow-limited and diffusion-limited transport depends on two new dimensionless parameters defined in terms of key geometric quantities, with strong solute uptake promoting flow-limited transport conditions. We present a simple algebraic approximation for solute uptake rate as a function of flow conditions, metabolic rate and villous geometry. For oxygen, accounting for nonlinear kinetics using physiological parameter values, our model predicts that villous metabolism does not significantly impact oxygen transfer to fetal blood, although the partitioning of fluxes between the villous tissue and the capillary network depends strongly on the flow regime.

Across mammalian species, solute exchange takes place in complex microvascular networks. In the human placenta, the primary exchange units are terminal villi that contain disordered networks of fetal capillaries and are surrounded externally by maternal blood. Here we show how the irregular internal structure of a terminal villus determines its exchange capacity for a wide range of solutes. Distilling geometric features into three scalar parameters, obtained from image analysis and computational fluid dynamics, we capture archetypal features of the the structure-function relationship of terminal villi using a simple algebraic approximation, revealing transitions between flow- and diffusion-limited transport at vessel and network levels. Our theory accommodates countercurrent effects, incorporates nonlinear blood rheology and offers an efficient method for testing network robustness. Our results show how physical estimates of solute transport, based on carefully defined geometrical statistics, provide a viable method for linking placental structure and function, and offer a framework for assessing transport in other microvascular systems.

Microtubules are filamentous tubular protein polymers which are essential for a range of cellular behaviour, and are generally straight over micron length scales. However, in some gliding assays, where microtubules move over a carpet of molecular motors, individual microtubules can also form tight arcs or rings, even in the absence of crosslinking proteins. Understanding this phenomenon may provide important explanations for similar highly curved microtubules which can be found in nerve cells undergoing neurodegeneration. We propose a model for gliding assays where the kinesins moving the microtubules over the surface induce ring formation through differential binding, substantiated by recent findings that a mutant version of the motor protein kinesin applied in solution is able to lock-in microtubule curvature. For certain parameter regimes, our model predicts that both straight and curved microtubules can exist simultaneously as stable steady-states, as has been seen experimentally. Additionally, unsteady solutions are found, where a wave of differential binding propagates down the microtubule as it glides across the surface, which can lead to chaotic motion. Whilst this model explains two-dimensional microtubule behaviour in an experimental gliding assay, it has the potential to be adapted to explain pathological curling in nerve cells., Comment: 24 pages, 6 figures. Accepted for publication in the Bulletin of Mathematical Biology

We consider a cellular monolayer, described using a vertex-based model, for which cells form a spatially disordered array of convex polygons that tile the plane. Equilibrium cell configurations are assumed to minimize a global energy defined in terms of cell areas and perimeters; energy is dissipated via dynamic area and length changes, as well as cell neighbour exchanges. The model captures our observations of an epithelium from a Xenopus embryo showing that uniaxial stretching induces spatial ordering, with cells under net tension (compression) tending to align with (against) the direction of stretch, but with the stress remaining heterogeneous at the single-cell level. We use the vertex model to derive the linearized relation between tissue-level stress, strain and strain-rate about a deformed base state, which can be used to characterize the tissue's anisotropic mechanical properties; expressions for viscoelastic tissue moduli are given as direct sums over cells. When the base state is isotropic, the model predicts that tissue properties can be tuned to a regime with high elastic shear resistance but low resistance to area changes, or vice versa., Comment: 9 figures

Transport in biological systems often occurs in complex spatial environments involving random structures. Motivated by such applications, we investigate an idealised model for solute transport past an array of point sinks, randomly distributed along a line, which remove solute via first-order kinetics. Random sink locations give rise to long-range spatial correlations in the solute field and influence the mean concentration. We present a non-standard approach to evaluating these features based on rationally approximating integrals of a suitable Green's function, which accommodates contributions varying on short and long lengthscales and has deterministic and stochastic components. We refine the results of classical two-scale methods for a periodic sink array (giving more accurate higher-order corrections with non-local contributions) and find explicit predictions for the fluctuations in concentration and disorder-induced corrections to the mean for both weakly and strongly disordered sink locations. Our predictions are validated across a large region of parameter space., Comment: 28 pages, 7 figures and 2 tables

A liquid drop spreading over a thin heterogeneous precursor film (such as an inhaled droplet on the mucus-lined wall of a lung airway) will experience perturbations in shape and location as its advancing contact line encounters regions of low or high film viscosity. Prior work on spatially one-dimensional spreading over a precursor film having a random viscosity field [Xu & Jensen 2016, Proc. Roy. Soc. A 472, 20160270] has demonstrated how viscosity fluctuations are swept into a narrow region behind the contact line, where they can impact drop dynamics. Here we investigate two-dimensional drops, seeking to understand the relationship between the statistical properties of the precursor film and those of the spreading drop. Assuming the precursor film is much thinner than the drop and viscosity fluctuations are weak, we use asymptotic methods to derive explicit predictions for the mean and variance of drop area and the drop's lateral drift. For larger film variability, we use Gaussian process emulation to estimate the variance of outcomes from a restricted set of simulations. Stochastic drift of the droplet is predicted to be greatest when the initial drop diameter is comparable to the correlation length of viscosity fluctuations., Comment: 23 pages, 5 figures

A liquid film wetting the interior of a long circular cylinder redistributes under the action of surface tension to form annular collars or occlusive plugs. These equilibrium structures are invariant under axial translation within a perfectly smooth uniform tube and therefore can be displaced axially by very weak external forcing. We consider how this degeneracy is disrupted when the tube wall is rough, and determine threshold conditions under which collars or plugs resist displacement under forcing. Wall roughness is modelled as a non-axisymmetric Gaussian random field of prescribed correlation length and small variance, mimicking some of the geometric irregularities inherent in applications such as lung airways. The thin film coating this surface is modelled using lubrication theory. When the roughness is weak, we show how the locations of equilibrium collars and plugs can be identified in terms of the azimuthally averaged tube radius; we derive conditions specifying equilibrium collar locations under an externally imposed shear flow, and plug locations under an imposed pressure gradient. We use these results to determine the probability of external forcing being sufficient to displace a collar or plug from a rough-walled tube, when the tube roughness is defined only in statistical terms.

Using a popular vertex-based model to describe a spatially disordered planar epithelial monolayer, we examine the relationship between cell shape and mechanical stress at the cell and tissue level. Deriving expressions for stress tensors starting from an energetic formulation of the model, we show that the principal axes of stress for an individual cell align with the principal axes of shape, and we determine the bulk effective tissue pressure when the monolayer is isotropic at the tissue level. Using simulations for a monolayer that is not under peripheral stress, we fit parameters of the model to experimental data for Xenopus embryonic tissue. The model predicts that mechanical interactions can generate mesoscopic patterns within the monolayer that exhibit long-range correlations in cell shape. The model also suggests that the orientation of mechanical and geometric cues for processes such as cell division are likely to be strongly correlated in real epithelia. Some limitations of the model in capturing geometric features of Xenopus epithelial cells are highlighted., Comment: 29 pages, 10 figures, revision

We examine theoretically the spreading of a viscous liquid drop over a thin film of uniform thickness, assuming the liquid's viscosity is regulated by the concentration of a solute that is carried passively by the spreading flow. The solute is assumed to be initially heterogeneous, having a spatial distribution with prescribed statistical features. To examine how this variability influences the drop's motion, we investigate spreading in a planar geometry using lubrication theory, combining numerical simulations with asymptotic analysis. We assume diffusion is sufficient to suppress solute concentration gradients across but not along the film. The solute field beneath the bulk of the drop is stretched by the spreading flow, such that the initial solute concentration immediately behind the drop's effective contact lines has a long-lived influence on the spreading rate. Over long periods, solute swept up from the precursor film accumulates in a short region behind the contact line, allowing patches of elevated viscosity within the precursor film to hinder spreading. A low-order model provides explicit predictions of the variances in spreading rate and drop location, which are validated against simulations.

Motivated by uncertainty quantification in natural transport systems, we investigate an individual-based transport process involving particles undergoing a random walk along a line of point sinks whose strengths are themselves independent random variables. We assume particles are removed from the system via first-order kinetics. We analyse the system using a hierarchy of approaches when the sinks are sparsely distributed, including a stochastic homogenization approximation that yields explicit predictions for the extrinsic disorder in the stationary state due to sink strength fluctuations. The extrinsic noise induces long-range spatial correlations in the particle concentration, unlike fluctuations due to the intrinsic noise alone. Additionally, the mean concentration profile, averaged over both intrinsic and extrinsic noise, is elevated compared with the corresponding profile from a uniform sink distribution, showing that the classical homogenization approximation can be a biased estimator of the true mean., Comment: 16 pages, 8 figures

During pregnancy, oxygen diffuses from maternal to fetal blood through villous trees in the placenta. In this paper, we simulate blood flow and oxygen transfer in feto-placental capillaries by converting three-dimensional representations of villous and capillary surfaces, reconstructed from confocal laser scanning microscopy, to finite-element meshes, and calculating values of vascular flow resistance and total oxygen transfer. The relationship between the total oxygen transfer rate and the pressure drop through the capillary is shown to be captured across a wide range of pressure drops by physical scaling laws and an upper bound on the oxygen transfer rate. A regression equation is introduced that can be used to estimate the oxygen transfer in a capillary using the vascular resistance. Two techniques for quantifying the effects of statistical variability, experimental uncertainty and pathological placental structure on the calculated properties are then introduced. First, scaling arguments are used to quantify the sensitivity of the model to uncertainties in the geometry and the parameters. Second, the effects of localized dilations in fetal capillaries are investigated using an idealized axisymmetric model, to quantify the possible effect of pathological placental structure on oxygen transfer. The model predicts how, for a fixed pressure drop through a capillary, oxygen transfer is maximized by an optimal width of the dilation. The results could explain the prevalence of fetal hypoxia in cases of delayed villous maturation, a pathology characterized by a lack of the vasculo-syncytial membranes often seen in conjunction with localized capillary dilations., Comment: Final version of manuscript