The Gaussian (saddle splay) rigidity of fluid membranes controls their equilibrium topology but is notoriously difficult to measure. In lipid mixtures, typical of living cells, linear interfaces separate liquid ordered (LO) from liquid disordered (LD) bilayer phases at subcritical temperatures. Here we consider such membranes supported by curved supports that thereby control the membrane curvatures. We show how spectral analysis of the fluctuations of the LO-LD interface provides a novel way of measuring the difference in Gaussian rigidity between the two phases. We provide a number of conditions for such interface fluctuations to be both experimentally measurable and sufficiently sensitive to the value of the Gaussian rigidity, whilst remaining in the perturbative regime of our analysis.
In this work I derive analytic expressions for the curvature dependent fluid-substrate surface tension of a hard sphere fluid on a hard curved wall. In a first step, the curvature thermodynamic properties are found as truncated power series in the activity in terms of the known second and third order cluster integral of the hard-sphere fluid near spherical and cylindrical walls. These results are then expressed as packing fraction power series and transformed to different reference regions which is equivalent to consider different positions of the dividing surface. All the obtained series expa…
We study thin self-assembled columns constrained to lie on a curved, rigid substrate. The curvature presents no local obstruction to equally spaced columns in contrast to curved crystals for which the crystalline bonds are frustrated. Instead, the vanishing compressional strain of the columns implies that their normals lie on geodesics which converge (diverge) in regions of positive (negative) Gaussian curvature, in analogy to the focussing of light rays by a lens. We show that the out of plane bending of the cylinders acts as an effective ordering field.
In the context of Rosenfeld's Fundamental Measure Theory, we show that the bending rigidity is not equal to zero for a hard-sphere fluid in contact with a curved hard wall. The implication is that the Hadwiger Theorem does not hold in this case and the surface free energy is given by the Helfrich expansion instead. The value obtained for the bending rigidity is (1) an order of magnitude smaller than the bending constant associated with Gaussian curvature, (2) changes sign as a function of the fluid volume fraction, (3) is independent of the choice for the location of the hard wall.
The preference of thin flat sheets to bend rather than stretch, combined with results from Geometry, mean that changes in a thin sheet's Gaussian curvature are prohibitively expensive. As a result, an imposed curvature in one principal direction inhibits bending in the other: so-called curvature-induced rigidity. Here, we study the buckling behaviour of a rectangular strip of finite thickness held horizontally in a gravitational field, but with a transverse curvature imposed at one end. The finite thickness of the sheet limits the efficacy of curvature-induced rigidity in two ways: (i) finite …
We argue that for complete wetting at a curved substrate (wall) the wall-fluid surface tension is non-analytic in $R_i^{-1}$, the curvature of the wall and that the density profile of the fluid near the wall acquires a contribution proportional to the gas-liquid surface tension $\times R_i^{-1}$ plus higher-order contributions which are non-analytic in $R_i^{-1}$. These predictions are confirmed by results of density functional calculations for the square-well model of a liquid adsorbed on a hard sphere and on a hard cylinder where complete wetting by gas (drying) occurs. The implications of o…
It is argued that to arrive at a quantitative description of the surface tension of a liquid drop as a function of its inverse radius, it is necessary to include the bending rigidity k and Gaussian rigidity k_bar in its description. New formulas for k and k_bar in the context of density functional theory with a non-local, integral expression for the interaction between molecules are presented. These expressions are used to investigate the influence of the choice of Gibbs dividing surface and it is shown that for a one-component system, the equimolar surface has a special status in the sense th…
We present a continuum model of the coupling between cells and substrate that accounts for some of the observed substrate-stiffness dependence of cell properties. The cell is modeled as an elastic active gel, adapting recently developed continuum theories of active viscoelastic fluids. The coupling to the substrate enters as a boundary condition that relates the cell's deformation field to local stress gradients. In the presence of activity, the coupling to the substrate yields spatially inhomogeneous contractile stresses and deformations in the cell and can enhance polarization, breaking the …
Using a mean field approach and simulation, we study the non-linear mechanical response of the vertex model (VM) of biological tissue under compression and dilation. The VM is known to exhibit a transition between rigid and fluid-like, or floppy, states driven by geometric incompatibility. Target perimeter and area set a target shape which may not be geometrically achievable, thereby engendering frustration. Previously, an asymmetry in the linear elastic response was identified at the rigidity transition between compression and dilation. Here we show and characterize how the asymmetry extends …
The sliding friction force exerted by stochastic linkers interacting with a moving filament is calculated. The elastic properties of the substrate on which the linkers are anchored are shown to strongly influence the friction force. In some cases, the force is maximal for a finite substrate rigidity. Collective effects give rise to a dynamical instability resulting in a stick-slip behaviour, which is substrate-sensitive. The relevance of these results for the motility of crawling cells powered by an actin retrograde flow is discussed.
We present a combined analytical approach and numerical study on the stability of a ring bound to an annular elastic substrate, which contains a circular cavity. The system is loaded by depressurizing the inner cavity. The ring is modeled as an Euler-Bernoulli beam and its equilibrium equations are derived from the mechanical energy which takes into account both stretching and bending contributions. The curvature of the substrate is considered explicitly to model the work done by its reaction force on the ring. We distinguish two different instabilities: periodic wrinkling of the ring or globa…
Coordinated and cooperative motion of cells is essential for embryonic development, tissue morphogenesis, wound healing and cancer invasion. A predictive understanding of the emergent mechanical behaviors in collective cell motion is challenging due to the complex interplay between cell-cell interactions, cell-matrix adhesions and active cell behaviors. To overcome this challenge, we develop a predictive cellular vertex model that can delineate the relative roles of substrate rigidity, tissue mechanics and active cell properties on the movement of cell collectives. We apply the model to the sp…
Motivated by recent experimental work on multicomponent lipid membranes supported by colloidal scaffolds, we report an exhaustive theoretical investigation of the equilibrium configurations of binary mixtures on curved substrates. Starting from the Jülicher-Lipowsky generalization of the Canham-Helfrich free energy to multicomponent membranes, we derive a number of exact relations governing the structure of an interface separating two lipid phases on arbitrarily shaped substrates and its stability. We then restrict our analysis to four classes of surfaces of both applied and conceptual interes…
We study the adsorption of semiflexible polymers such as polyelectrolytes or DNA on planar and curved substrates, e.g., spheres or washboard substrates via short-range potentials using extensive Monte-Carlo simulations, scaling arguments, and analytical transfer matrix techniques. We show that the adsorption threshold of stiff or semiflexible polymers on a planar substrate can be controlled by polymer stiffness: adsorption requires the highest potential strength if the persistence length of the polymer matches the range of the adsorption potential. On curved substrates, i.e., an adsorbing sphe…
Smectic order on arbitrary curved substrate can be described by a differential form of rank one (1-form), whose geometric meaning is the differential of the local phase field of the density modulation. The exterior derivative of 1-form is the local dislocation density. Elastic deformations are described by superposition of exact differential forms. We use the formalism of differential forms to systematically classify and characterize all low energy smectic states on torus as well as on sphere. A two dimensional smectic order confined on either manifold exhibits many topologically distinct low …
We present a novel mechanism for resolving the mechanical rigidity of nanoscopic circular polymers that flow in a complex environment. The emergence of a regime of negative differential mobility induced by topological interactions between the rings and the substrate is the key mechanism for selective sieving of circular polymers with distinct flexibilities. A simple model accurately describes the sieving process observed in molecular dynamics simulations and yields experimentally verifiable analytical predictions, which can be used as a reference guide for improving filtration procedures of ci…
Multi-component membranes such as ternary mixtures of lipids and cholesterol can exhibit coexistence regions between two liquid phases. When such membranes adhere to a corrugated substrate, the phase separation process strongly depends on the interplay between substrate topography, bending rigidities, and line tension of the membrane domains as we show theoretically via energy minimization and Monte Carlo simulations. For sufficiently large bending rigidity contrast between the two membrane phases, the corrugated substrate truncates the phase separation process and leads to a stable pattern of…
Eukaryotic cells adhere to extracellular matrix during the normal development of the organism, forming static adhesion as well as during cell motility. We study this process by considering a simplified coarse-grained model of a vesicle that has uniform adhesion energy with a flat substrate, mobile curved membrane proteins and active forces. We find that a high concentration of curved proteins alone increases the spreading of the vesicle, by the self-organization of the curved proteins at the high curvature vesicle-substrate contact line, thereby reducing the bending energy penalty at the vesic…
Flaps can be detached from a thin film glued on a solid substrate by tearing and peeling. For flat substrates, it has been shown that these flaps spontaneously narrow and collapse in pointy triangular shapes. Here we show that various shapes, triangular, elliptic, acuminate or spatulate, can be observed for the tears by adjusting the curvature of the substrate. From combined experiments and theoretical models, we show that the flap morphology is governed by simple geometric rules.
When thermal energies are weak, two dimensional lamellar structures confined on a curved substrate display complex patterns arising from the competition between layer bending and compression in the presence of geometric constraints. We present broad design principles to engineer the geometry of the underlying substrate so that a desired lamellar pattern can be obtained by self-assembly. Two distinct physical effects are identified as key factors that contribute to the interaction between the shape of the underlying surface and the resulting lamellar morphology. The first is a local ordering fi…
We investigate liquid layers adsorbed at spherical and corrugated cylindrical substrates. The effective Hamiltonians for the liquid-gas interfaces fluctuating in the presence of such curved substrates are derived via the mean-field density functional theory. Their structure is compared with the Helfrich Hamiltonian which is parametrized by the bending and Gaussian rigidity coefficients. For long-ranged interparticle interactions of van der Waals type these coefficients turn out to be non-universal functions of interfacial curvatures; their form varies from one interface to another. We discuss …
We study the global influence of curvature on the free energy landscape of two-dimensional binary mixtures confined on closed surfaces. Starting from a generic effective free energy, constructed on the basis of symmetry considerations and conservation laws, we identify several model-independent phenomena, such as a curvature-dependent line tension and local shifts in the binodal concentrations. To shed light on the origin of the phenomenological parameters appearing in the effective free energy, we further construct a lattice-gas model of binary mixtures on non-trivial substrates, based on the…
Nucleation and growth is the dominant relaxation mechanism driving first order phase transitions. In two-dimensional at systems nucleation has been applied to a wide range of problems in physics, chemistry and biology. Here we study nucleation and growth of two-dimensional phases lying on curved surfaces and show that curvature modify both, critical sizes of nuclei and paths towards the equilibrium phase. In curved space nucleation and growth becomes inherently inhomogeneous and critical nuclei form faster on regions of positive Gaussian curvature. Substrates of varying shape display complex e…
Based on energy conservation, we derive a critical condition theoretically for electrowettinginduced droplet detachment from a hydrophobic curved surface. Phase diagrams are constructed in terms of droplet volume, viscosity, Ohnesorge number, friction coefficient at contact line, surface curvature, surface wettability and electrowetting number. The deduced critical condition offers a general and quantitative prediction on when the detachment occurs, a criterion enabling us to gain more insights into how to accurately manipulate the electrowetting-induced detachment of an aqueous droplet on a c…
We investigate the turbulent dynamics of a two-dimensional active nematic liquid crystal con- strained on a curved surface. Using a combination of hydrodynamic and particle-based simulations, we demonstrate that the fundamental structural features of the fluid, such as the topological charge density, the defect number density, the nematic order parameter and defect creation and annihilation rates, are simple linear functions of the substrate Gaussian curvature, which then acts as a control parameter for the chaotic flow. Our theoretical predictions are then compared with experiments on microtu…
In this Letter, we consider a neutral system of mobile positive and negative charges confined on the surface of curved films. This may be an appropriate model for: i) a highly charged membrane whose counterions are confined to a sheath near its surface; ii) a membrane composed of an equimolar mixture of anionic and cationic surfactants in aqueous solution. We find that the charge fluctuations contribute a non-analytic term to the bending rigidity that varies logarithmically with the radius of curvature. This may lead to spontaneous vesicle formation, which is indeed observed in similar systems…
The properties of the interface in a phase-separated solution of polymers with different degrees of polymerization and Kuhn segment lengths are calculated. The starting point is the planar interface, the profile of which is calculated in the so-called 'blob model', which incorporates the solvent in an implicit way. The next step is the study of a metastable droplet phase formed by imposing a chemical potential different from that at coexistence. The pressure difference across the curved interface, which corresponds to this higher chemical potential, is used to calculate the curvature propertie…
We investigate a continuum mechanical model for an adherent cell on two dimensional adhesive micropatterned substrates. The cell is modeled as an isotropic and homogeneous elastic material subject to uniform internal contractile stresses. The build-up of tension from cortical actin bundles at the cell periphery is incorporated by introducing an energy cost for bending of the cell boundary, resulting to a resistance to changes in local curvature. Integrin-based adhesions are modeled as harmonic springs, that pin the cell to adhesive patches of a predefined geometry. Using Monte Carlo simulation…
A new set of mathematical identities is presented for axi-symmetric sessile drops on flat and curved substrates. The geometrical parameters, including the apex curvature and height, and the contact radius, are related by the identities. The validity of the identities are checked by various numerical solutions both for flat and curved substrates.
It is widely recognized that the shape of epithelial cells is determined by the tension generated by the actomyosin cortex and the adhesion of cells to the substrate and to each other. To account for these biological and structural contributions to cell shape, different physical models have been proposed. However, an experimental procedure that would allow a validation of a minimal physical model for the shape of epithelial cells in 3D has not yet been proposed. In this study, we cultured MDCK epithelial cells on substrates with a sinusoidal profile, allowing us to measure the shape of the cel…
We study the dynamics of a one-component liquid constrained on a spherical substrate, a 2-sphere, and investigate how the mode-coupling theory (MCT) can describe the new features brought by the presence of curvature. To this end we have derived the MCT equations in a spherical geometry. We find that, as seen from the MCT, the slow dynamics of liquids in curved space at low temperature does not qualitatively differ from that of glass-forming liquids in Euclidean space. The MCT predicts the right trend for the evolution of the relaxation slowdown with curvature but is dramatically off at a quant…
Cells move differently on substrates with different elasticities. In particular, the persistence time of their motion is higher on stiffer substrates. We show that this behavior will result in a net transport of cells directed up a soft-to-stiff gradient. Using simple random walk models with controlled persistence and stochastic simulations, we characterize this propensity to move in terms of the durotactic index measured in experiments. A one-dimensional model captures the essential features of this motion and highlights the competition between diffusive spreading and linear, wavelike propaga…
We present a systematic study of how vortices in superfluid films interact with the spatially varying Gaussian curvature of the underlying substrate. The Gaussian curvature acts as a source for a geometric potential that attracts (repels) vortices towards regions of negative (positive) Gaussian curvature independently of the sign of their topological charge. Various experimental tests involving rotating superfluid films and vortex pinning are first discussed for films coating gently curved substrates that can be treated in perturbation theory from flatness. An estimate of the experimental regi…
The adhesion of a stiff film onto a curved substrate often generates elastic stresses in the film that eventually give rise to its delamination. Here we predict that delamination of very thin films can be dramatically suppressed through tiny, smooth deformations of the substrate, dubbed here "wrinklogami", that barely affect the macroscale topography. This "pro-lamination" effect reflects a surprising capability of smooth wrinkles to suppress compression in elastic films even when spherical or other doubly-curved topography is imposed, in a similar fashion to origami folds that enable construc…
Ordered phases on curved substrates experience a complex interplay of ordering and intrinsic curvature, commonly producing frustration and singularities. This is an especially important issue in crystals as ever-smaller scale materials are grown on real surfaces; eventually, surface imperfections are on the same scale as the lattice constant. Here, we gain insights into this general problem by studying two-dimensional smectic order on substrates with highly localized intrinsic curvature, constructed from cones and their intersections with planes. In doing so we take advantage of fully tractabl…
