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In this work, we study a phase-field model for curvature-driven pattern formation in biomembranes. The model is derived as a gradient flow of an energy functional that approximates the two-phase Canham--Helfrich energy. This leads to a Cahn--Hilliard-type equation with cross diffusion for the relative chemical concentration of one lipid phase, coupled to a fourth-order reaction-diffusion equation describing the height profile of the membrane. We first prove the existence of weak solutions for the case of regular double-well potentials, using a minimizing movement scheme to construct approximate solutions. The analysis is then extended to singular potentials, e.g., the Flory--Huggins potential, by approximating them with a Moreau--Yosida regularization. For both cases, we establish higher regularity, continuous dependence on the initial data, and consequently the uniqueness of weak solutions. Finally, we propose a well-posed finite element discretization of the model and present numerical experiments illustrating the effect of different physical parameters on the resulting membrane patterns. Depending on the parameter regime, we observe purely striped, dotted, or snake-like structur

We consider a one-dimensional version of a variational model for pattern formation in biological membranes. The driving term in the energy is a coupling between the order parameter and the local curvature of the membrane. We derive scaling laws for the minimal energy. As a main tool we present new nonlinear interpolation inequalities that bound fractional Sobolev seminorms in terms of a Cahn-Hillard/Modica-Mortola energy.

We present a numerical method to model the dynamics of inextensible biomembranes in a quasi-Newtonian incompressible flow, which better describes hemorheology in the small vasculature. We consider a level set model for the fluid-membrane coupling, while the local inextensibility condition is relaxed by introducing a penalty term. The penalty method is straightforward to implement from any Navier-Stokes/level set solver and allows substantial computational savings over a mixed formulation. A standard Galerkin finite element framework is used with an arbitrarily high order polynomial approximation for better accuracy in computing the bending force. The PDE system is solved using a partitioned strongly coupled scheme based on Crank-Nicolson time integration. Numerical experiments are provided to validate and assess the main features of the method.

We derive a macroscopic limit for a sharp interface version of a model proposed in [29] to investigate pattern formation due to competition of chemical and mechanical forces in biomembranes. We identify sub- and supercrital parameter regimes and show with the introduction of the autocorrelation function that the ground state energy leads to the isoperimetric problem in the subcritical regime, which is interpreted to not form fine scale patterns.

We review the theoretical analyses and simulations of the interactions between curvature-inducing proteins and biomembranes. Laterally isotropic proteins induce spherical budding, whereas anisotropic proteins, such as Bin/Amphiphysin/Rvs (BAR) superfamily proteins, induce tabulation. Both types of proteins can sense the membrane curvature. We describe the theoretical analyses of various transitions of protein binding accompanied by a change in various properties, such as the number of buds, the radius of membrane tubes, and the nematic order of anisotropic proteins. Moreover, we explain the membrane-mediated interactions and protein assembly. Many types of membrane shape transformations (spontaneous tubulation, formation of polyhedral vesicles, polygonal tubes, periodic bumps, and network structures, etc.) have been demonstrated by coarse-grained simulations. Furthermore, traveling waves and Turing patterns under the coupling of reaction-diffusion dynamics and membrane deformation are described.

The form and evolution of multi-phase biomembranes is of fundamental importance in order to understand living systems. In order to describe these membranes, we consider a mathematical model based on a Canham--Helfrich--Evans two-phase elastic energy, which will lead to fourth order geometric evolution problems involving highly nonlinear boundary conditions. We develop a parametric finite element method in an axisymmetric setting. Using a variational approach, it is possible to derive weak formulations for the highly nonlinear boundary value problems such that energy decay laws, as well as conservation properties, hold for spatially discretised problems. We will prove these properties and show that the fully discretised schemes are well-posed. Finally, several numerical computations demonstrate that the numerical method can be used to compute complex, experimentally observed two-phase biomembranes.

Cell plasma membranes display a dramatically rich structural complexity characterized by functional sub-wavelength domains with specific lipid and protein composition. Under favorable experimental conditions, patterned morphologies can also be observed in vitro on model systems such as supported membranes or lipid vesicles. Lipid mixtures separating in liquid-ordered and liquid-disordered phases below a demixing temperature play a pivotal role in this context. Protein-protein and protein-lipid interactions also contribute to membrane shaping by promoting small domains or clusters. Such phase separations displaying characteristic length-scales falling in-between the nanoscopic, molecular scale on the one hand and the macroscopic scale on the other hand, are named mesophases in soft condensed matter physics. In this review, we propose a classification of the diverse mechanisms leading to mesophase separation in biomembranes. We distinguish between mechanisms relying upon equilibrium thermodynamics and those involving out-of-equilibrium mechanisms, notably active membrane recycling. In equilibrium, we especially focus on the many mechanisms that dwell on an up-down symmetry breaking b

We examine the stability of a class of solitons, obtained from a generalization of the Boussinesq equation, which have been proposed to be relevant for pulse propagation in biomembranes and nerves. These solitons are found to be stable with respect to small amplitude fluctuations. They emerge naturally from non-solitonic initial excitations and are robust in the presence of dissipation.

We consider sharp interface asymptotics for a phase field model of two phase near spherical biomembranes involving a coupling between the local mean curvature and the local composition proposed by the first and second authors. The model is motivated by lipid raft formation. We introduce a reduced diffuse interface energy depending only on the membrane composition and derive the $Γ-$limit. We demonstrate that the Euler-Lagrange equations for the limiting functional and the sharp interface energy coincide. Finally, we consider a system of gradient flow equations with conserved Allen-Cahn dynamics for the phase field model. Performing a formal asymptotic analysis we obtain a system of gradient flow equations for the sharp interface energy coupling geodesic curvature flow for the phase interface to a fourth order PDE free boundary problem for the surface deformation.

Collective behavior of proteins on biomembranes is usually studied within the spontaneous curvature model. Here we consider an alternative phenomenological approach, which accounts consistently for partial ordering of proteins as well as the anchoring forces exerted on a membrane by layer of proteins. We show analytically that such anisotropic interactions can drive membrane bending, resulting in non-trivial equilibrium morphologies. The predicted instabilities can advance our conceptual understanding of physical mechanisms behind collective phenomena in biological systems, in particular those with inherent anisotropy.

Biomembranes and vesicles consisting of multiple phases can attain a multitude of shapes, undergoing complex shape transitions. We study a Cahn--Hilliard model on an evolving hypersurface coupled to Navier--Stokes equations on the surface and in the surrounding medium to model these phenomena. The evolution is driven by a curvature energy, modelling the elasticity of the membrane, and by a Cahn--Hilliard type energy, modelling line energy effects. A stable semidiscrete finite element approximation is introduced and, with the help of a fully discrete method, several phenomena occurring for two-phase membranes are computed.

Boussinesq-type wave equations involve nonlinearities and dispersion. In this paper a Boussinesq-type equation with amplitude-dependent nonlinearities is presented. Such a model was proposed by Heimburg and Jackson (2005) for describing longitudinal waves in biomembranes and later improved by Engelbrecht et al. (2015) taking into account the microinertia of a biomembrane. The steady solution in the form of a solitary wave is derived and the influence of nonlinear and dispersive terms over a large range of possible sets of coefficients demonstrated. The solutions emerging from arbitrary initial inputs are found using the numerical simulation. The properties of emerging trains of solitary waves waves are analysed. Finally, the interaction of solitary waves which satisfy the governing equation is studied. The interaction process is not fully elastic and after several interactions radiation effects may be significant. This means that for the present case the solitary waves are not solitons in the strict mathematical sense. However, like in other cases known in solid mechanics, such solutions may be conditionally called solitons.

Biomembranes play a central role in various phenomena like locomotion of cells, cell-cell interactions, packaging of nutrients, and in maintaining organelle morphology and functionality. During these processes, the membranes undergo significant morphological changes through deformation, scission, and fusion. Modeling the underlying mechanics of such morphological changes has traditionally relied on reduced order axisymmetric representations of membrane geometry and deformation. Axisymmetric representations, while robust and extensively deployed, suffer from their inability to model symmetry breaking deformations and structural bifurcations. To address this limitation, a 3D computational mechanics framework for high fidelity modeling of biomembrane deformation is presented. The proposed framework brings together Kirchhoff-Love thin-shell kinematics, Helfrich-energy based mechanics, and state-of-the-art numerical techniques for modeling deformation of surface geometries. Lipid bilayers are represented as spline-based surfaces immersed in a 3D space; this enables modeling of a wide spectrum of membrane geometries, boundary conditions, and deformations that are physically admissible in

We have studied biomembranes with grafted polymer chains using a coarse-grained membrane simulation, where a meshless membrane model is combined with polymer chains. We focus on the polymer-induced entropic effects on mechanical properties of membranes. The spontaneous curvature and bending rigidity of the membranes increase with increasing polymer density. Our simulation results agree with the previous theoretical predictions.

We prove solution uniqueness for the genus one Canham variational problem arising in the shape prediction of biomembranes. The proof builds on a result of Yu and Chen that reduces the variational problem to proving non-negativity of a sequence defined by a linear recurrence relation with polynomial coefficients. We combine rigorous numeric analytic continuation of D-finite functions with classic bounds from singularity analysis to derive an effective index where the asymptotic behaviour of the sequence, which is positive, dominates the sequence behaviour. Positivity of the finite number of remaining terms is then checked computationally.

In this paper we introduce a mathematical model for small deformations induced by external forces of closed surfaces that are minimisers of Helfrich-type energies. Our model is suitable for the study of deformations of cell membranes induced by the cytoskeleton. We describe the deformation of the surface as a graph over the undeformed surface. A new Lagrangian and the associated Euler-Lagrange equations for the height function of the graph are derived. This is the natural generalisation of the well known linearisation in the Monge gauge for initially flat surfaces. We discuss energy perturbations of point constraints and point forces acting on the surface. We establish existence and uniqueness results for weak solutions on spheres and on tori. Algorithms for the computation of numerical solutions in the general setting are provided. We present numerical examples which highlight the behaviour of the surface deformations in different settings at the end of the paper.

Transmembrane ion flow through channel proteins undergoing density fluctuations may cause lateral gradients of the electrical potential across the membrane giving rise to electrophoresis of charged channels. A model for the dynamics of the channel density and the voltage drop across the membrane (cable equation) coupled to a binding-release reaction with the cell skeleton (P. Fromherz and W. Zimmerman, Phys. Rev. E 51, R1659 (1995)) is analyzed in one and two spatial dimensions. Due to the binding release reaction spatially periodic modulations of the channel density with a finite wave number are favored at the onset of pattern formation, whereby the wave number decreases with the kinetic rate of the binding-release reaction. In a two-dimensional extended membrane hexagonal modulations of the ion channel density are preferred in a large range of parameters. The stability diagrams of the periodic patterns near threshold are calculated and in addition the equations of motion in the limit of a slow binding-release kinetics are derived.

Nonequilibrium dynamics of biomembranes with active inclusions is considered. The inclusions represent protein molecules which perform cyclic internal conformational motions driven by the energy brought with ATP ligands. As protein conformations cyclically change, this induces hydrodynamical flows and also directly affects the local curvature of a membrane. On the other hand, variations in the local curvature of the membrane modify the transitions rates between conformational states in a protein, leading to a feedback in the considered system. Moreover, active inclusions can move diffusively through the membrane so that surface concentration varies. The kinetic description of this system is constructed and the stability of the uniform stationary state is analytically investigated. We show that, as the rate of supply of chemical energy is increased above a certain threshold, this uniform state becomes unstable and stationary or traveling waves spontaneously develop in the system. Such waves are accompanied by periodic spatial variation of membrane curvature and inclusion density. For typical parameter values, their characteristic wavelengths are of the order of hundreds of nanometer

We study a physical model for the interaction between general inclusions bound to fluid membranes that possess finite tension, as well as the usual bending rigidity. We are motivated by an interest in proteins bound to cell membranes that apply forces to these membranes, due to either entropic or direct chemical interactions. We find an exact analytic solution for the repulsive interaction between two similar circularly symmetric inclusions. This repulsion extends over length scales of order tens of nanometers, and contrasts with the membrane-mediated contact attraction for similar inclusions on tensionless membranes. For non circularly symmetric inclusions we study the small, algebraically long-ranged, attractive contribution to the force that arises. We discuss the relevance of our results to biological phenomena, such as the budding of caveolae from cell membranes and the striations that are observed on their coats.