With tangent angle perturbation approach the axial symmetry deformation of a spherical vesicle in large under the pressure changes is studied by the elasticity theory of Helfrich spontaneous curvature model.Three main results in axial symmetry shape: biconcave shape, peanut shape, and one type of myelin are obtained. These axial symmetry morphology deformations are in agreement with those observed in lipsome experiments by dark-field light microscopy [Hotani, J. Mol. Biol. 178, (1984) 113] and in the red blood cell with two thin filaments (myelin) observed in living state (see, Bessis, Living Blood Cells and Their Ultrastructure, Springer-Verlag, 1973). Furthermore, the biconcave shape and peanut shape can be simulated with the help of a powerful software, Surface Evolver [Brakke, Exp. Math. 1, 141 (1992) 141], in which the spontaneous curvature can be easy taken into account.
By means of Surface Evolver (Exp. Math,1,141 1992), a software package of brute-force energy minimization over a triangulated surface developed by the geometry center of University of Minnesota, we have numerically searched the non-axisymmetric shapes under the Helfrich spontaneous curvature (SC) energy model. We show for the first time there are abundant mechanically stable non-axisymmetric vesicles in SC model, including regular ones with intrinsic geometric symmetry and complex irregular ones. We report in this paper several interesting shapes including a corniculate shape with six corns, a…
We propose a free energy expression accounting for the formation of spherical vesicles from planar lipidic membranes and derive a Fokker-Planck equation for the probability distribution describing the dynamics of vesicle formation. We found that formation may occur as an activated process for small membranes and as a transport process for sufficiently large membranes. We give explicit expressions for the transition rates and the characteristic time of vesicle formation in terms of the relevant physical parameters.
We investigate the morphology of a toroidal fluid membrane vesicle confined inside a spherical container. The equilibrium shapes are assembled in a geometrical phase diagram as a function of scaled area and reduced volume of the membrane. For small area the vesicle can adopt its free form. When increasing the area, the membrane cannot avoid contact and touches the confining sphere along a circular contact line, which extends to a zone of contact for higher area. The elastic energies of the equilibrium shapes are compared to those of their confined counterparts of spherical topology to predict …
We develop an analytical theory to explain the experimentally-observed morphological transitions of giant vesicles induced by AC electric fields (1). The model treats the inner and suspending media as lossy dielectrics, while the membrane as an ion-impermeable flexible incompressible-fluid sheet. The vesicle shape is obtained by balancing electric, hydrodynamic, and bending stresses exerted on the membrane. Considering a nearly spherical vesicle, the solution to the electrohydrodynamic problem is obtained as a regular perturbation expansion in the excess area. The theory predicts that stationa…
The morphology of spherically confined flexoelectric fluid membrane vesicles in an external uniform electric field is studied numerically. Due to the deformations induced by the confinement, the membrane becomes polarized resulting in an interaction with the external field. The equilibrium shapes of the vesicle without electric field can be classified in a geometrical phase diagram as a function of scaled area and reduced volume [1,2]. When the area of the membrane is only slightly larger than the area of the confining sphere, a single axisymmetric invagination appears. A non-vanishing electri…
We investigate the electrostatics of a partially formed, charged spherical shell in a salt solution. We solve the problem numerically at the Poisson-Boltzmann level and analytically in the Debye-Huckel regime. From the results on energetics of partially formed shells we examine the stability of tethered (crystalline) and fluid shells towards rupture. We clearly delineate different regimes of stability towards rupture, where, for fluid shells, we also include the effects of bending elasticity of the shells. Our analysis shows how charging of the shell induces its instability towards rupture but…
A model of vesicle electrodeformation is described which obtains a parametrized vesicle shape by minimizing the sum of the membrane bending energy and the energy due to the electric field. Both the vesicle membrane and the aqueous media inside and outside the vesicle are treated as leaky dielectrics, and the vesicle itself is modelled as a nearly spherical shape enclosed within a thin membrane. It is demonstrated (a) that the model achieves a good quantitative agreement with the experimentally determined prolate-to-oblate transition frequencies in the kHz range, and (b) that the model can expl…
Nuclear pores have an approximately uniform distribution in the nuclear envelope of most living cells. Hence, the morphology of the nuclear envelope is a spherical stomatocyte with a high genus. We have investigated the morphology of high-genus vesicles under pore-size constraint using dynamically triangulated membrane simulations. Bending-energy minimization without volume or other constraints produces a circular-cage stomatocyte, where the pores are aligned in a circular line on an oblate bud. As the pore radius is reduced, the circular-pore alignment is more stabilized than a random pore di…
We present a model of bi-phasic vesicle in the limit of large surface tension. In this regime, the vesicle is completely stretched and well described by two spherical caps with a fold which concentrates the membrane stress. The conservation laws and geometric constraints restrict the space of possible shapes to a pair of solutions labeled by a parameter $τ$ given by {\it line tension/pressure}. For a given $τ$ value, the two solutions differ by the length of the interface between domains. For a critical value $τ\_c$, the two vesicle shapes become identical and no solution exists above this cri…
An analytical theory is developed to describe the dynamics of a closed lipid bilayer membrane (vesicle) freely suspended in a general linear flow. Considering a nearly spherical shape, the solution to the creeping-flow equations is obtained as a regular perturbation expansion in the excess area. The analysis takes into account the membrane fluidity, incompressibility and resistance to bending. The constraint for a fixed total area leads to a non-linear shape evolution equation at leading order. As a result two regimes of vesicle behavior, tank-treading and tumbling, are predicted depending on …
The disk-to-vesicle transition of a fluid membrane with no spontaneous curvature is well described by the competition between edge line and curvature energies. However, the transition of asymmetric membranes with spontaneous curvatures is not yet understood. In this study, the shape of the fluid membrane patch with a constant spontaneous curvature and its closing transition to a vesicle is investigated using theory and meshless membrane simulations. It is confirmed that the (meta)stable and transient membranes are well approximated by spherical caps. The membrane Gaussian modulus can be estima…
A tethered vesicle, which consists of a cylindrical membrane tube and a spherical vesicle, is produced by a mechanical force that is experimentally imposed by optical tweezers and a micropipette. This tethered vesicle is employed for examining the curvature sensing of curvature-inducing proteins. In this study, we clarify how the binding of proteins with a laterally isotropic spontaneous curvature senses and generates the membrane curvatures of the tethered vesicle using mean-field theory and meshless membrane simulation. The force-dependence curves of the protein density in the membrane tube …
Narrow membrane tubes are commonly pulled out from the surface of phospholipid vesicles using forces applied either through laser or magnetic tweezers or through the action of processive motor proteins. Recent examples have emerged where such tubes spontaneously grow from vesicles coated with bioactive cytoskeletal filaments (e.g. FtsZ, microtubule) in the presence GTP. We show how a soft vesicle deforms due to the interplay between its topology, local curvature and the forces due to the active filaments. We present results from Dynamically Triangulated Monte Carlo simulations of a spherical c…
Vesicle budding induced by protein binding that generates an isotropic spontaneous curvature is studied using a mean-field theory. Many spherical buds are formed via protein binding. As the binding chemical potential increases, the proteins first bind to the buds and then to the remainder of the vesicle. For a high spontaneous curvature and/or high bending rigidity of the bound membrane, it is found that a first-order transition occurs between a small number of large buds and a large number of small buds. These two states coexist around the transition point. The proposed scheme is simple and e…
Biological membranes constantly change their shape in response to external stimuli, and understanding the remodeling and stability of vesicles in heterogeneous environments is therefore of fundamental importance for a range of cellular processes. One crucial question is how vesicles respond to external osmotic stresses, imposed by differences in solute concentrations between the vesicle interior and exterior. Previous analyses of the membrane bending energy have predicted that micron-sized giant unilamellar vesicles (GUVs) should become globally deformed already for nanomolar concentration dif…
We report the self-assembly of thin actin shells beneath the membranes of giant vesicles. Ion-carrier mediated influx of Mg2+ induces actin polymerization in the initially spherical vesicles. Buckling of the vesicles and the formation of blisters after thermally induced bilayer expansion is demonstrated. Bilayer flickering is dominated by tension generated by its coupling to the actin cortex. Quantitative flicker analysis suggests the bilayer and the actin cortex are separated by 0.4 \mum to 0.5 \mum due to undulation forces.
Lipid vesicles play a key role in fundamental biological processes. Despite recent progress, we lack a complete understanding of the non-equilibrium dynamics of vesicles due to challenges associated with long-time observation of shape fluctuations in strong flows. In this work, we present a flow-phase diagram for vesicle shape and conformational transitions in planar extensional flow using a Stokes trap, which enables control over the center-of-mass position of single or multiple vesicles in precisely defined flows [Shenoy, Rao, Schroeder, \textit{PNAS}, 113(15):3976-3981, 2016]. In this way, …
We investigate the stability of growing vesicles using the formalism of nonequilibrium thermodynamics. The vesicles are growing due to the accretion of lipids to the bilayer which forms the vesicle membrane. The thermodynamic description is based on the hydrodynamics of a water{/}lipid mixture together with a model of the vesicle as a discontinuous system in the sense of linear nonequilibrium thermodynamics. This formulation allows the forces and fluxes relevant to the dynamic stability of the vesicle to be identified. The method is used to analyze the stability of a spherical vesicle against …
Membrane budding has been extensively studied as an equilibrium process attributed to the formation of coexisting domains or changes in the vesicle area to volume ratio (reduced volume). In contrast, non-equilibrium budding remains experimentally widely unexplored especially when time scales fall well below the characteristic diffusion time of lipidsτ . We show that localized mechanical perturbations, initiated by driving giant unilamellar vesicles (GUVs) through their lipid phase transition, leads to the immediate formation of rapidly growing, multiply localized, non-equilibrium buds, when th…
Phagocytosis is the process of engulfment and internalization of comparatively large particles by the cell, that plays a central role in the functioning of our immune system. We study the process of phagocytosis by considering a simplified coarse grained model of a three-dimensional vesicle, having uniform adhesion interaction with a rigid particle, in the presence of curved membrane proteins and active cytoskeletal forces. Complete engulfment is achieved when the bending energy cost of the vesicle is balanced by the gain in the adhesion energy. The presence of curved (convex) proteins reduces…
Whilst many vertebrates appear externally left-right symmetric, the arrangement of internal organs is asymmetric. In zebrafish, the breaking of left-right symmetry is organised by Kupffer's Vesicle (KV): an approximately spherical, fluid-filled structure that begins to form in the embryo 10 hours post fertilisation. A crucial component of zebrafish symmetry breaking is the establishment of a cilia-driven fluid flow within KV. However, it is still unclear (a) how dorsal, ventral and equatorial cilia contribute to the global vortical flow, and (b) if this flow breaks left-right symmetry through …
Curvature-inducing proteins containing a Bin/Amphiphysin/Rvs domain often have intrinsically disordered domains. Recent experiments have shown that these disordered chains enhance curvature sensing and generation. Here, we report on the modification of protein-membrane interactions by disordered chains using meshless membrane simulations. The protein and bound membrane are modeled together as a chiral crescent protein rod with two excluded-volume chains. As the chain length increases, the repulsion between them reduces the cluster size of the proteins. It induces spindle-shaped vesicles and a …
Spatial organisation is a hallmark of all living cells, and recreating it in model systems is a necessary step in the creation of synthetic cells. It is therefore of both fundamental and practical interest to better understand the basic mechanisms underlying spatial organisation in cells. In this work, we use a continuum model of membrane and protein dynamics to study the behaviour of curvature-inducing proteins on membranes of spherical shape, such as living cells or lipid vesicles. We show that the interplay between curvature energy, entropic forces, and the geometric constraints on the memb…
The curvature sensitive localization of proteins on membranes is vital for many cell biological processes. Coarse-grained models are routinely employed to study the curvature sensing phenomena and membrane morphology at the length scale of few micrometers. Two prevalent phenomenological models exist for modeling experimental observations of curvature sensing, (1) the spontaneous curvature model and (2) the curvature mismatch model, which differ in their treatment of the change in elastic energy due to the binding of proteins on the membrane. In this work, the prediction of sensing and generati…
A micelle consists of monolayer of lipid molecules containing hydrophilic head and hydrophobic tail. These amphiphilic molecules in aqueous environment aggregate spontaneously into monomolecular layer held together due to hydrophobic effect by weak non-covalent forces. Micelles are flexible surfaces that show variety of shapes of different topology, but remarkably in mechanical equilibrium conditions they are spherical in shape. The shape and size of a micelle are functions of many variables such as lipid concentration, temperature, ionic strength, etc. Addressing the question, why the shape o…
The dynamics of a spherical elastic capsule, containing a Newtonian fluid bounded by an elastic membrane and immersed in another Newtonian fluid, in a uniform DC electric field is investigated. Discontinuity of electrical properties such as conductivities of the internal and external fluid media as well as capacitance and conductance of the membrane lead to a net interfacial Maxwell stress which can cause the deformation of such an elastic capsule. We investigate this problem considering well established membrane laws for a thin elastic membrane, with fully resolved hydrodynamics in the Stokes…
