Thomas Kwok-keung Au, Tom Yau-heng Wan
The Helfrich's shape equation of axisymmetric vesicles is studied. A sufficient condition on the physical parameters and some geometric properties are discovered for the biconcave shape.
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In this article, the formation of biconcave vesicle according to the minimization bending energy (Helfrich's model) is studied. We present that in order to have biconcave shape vesicle, there are necessary conditions on the parameters of the energy functional and there are implications on the geometric quantities of the vesicle. Furthermore, we also give a clear sufficient condition on the parameters to have the formation of biconcave shape vesicle.
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 …
The Helfrich bending energy plays an important role in providing a mechanism for the conformation of a lipid vesicle in theoretical biophysics, which is governed by the principle of energy minimization over configurations of appropriate topological characteristics. We will show that the presence of a quantity called the spontaneous curvature obstructs the existence of a minimizer of the Helfrich energy over the set of embedded ring tori. Besides, despite the well-realized knowledge that lipid vesicles may present themselves in a variety of shapes of complicated topology, there is a lack of top…
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.
(200 words) Oxidation of unsaturated lipids is a fundamental process involved in cell bioenergetics as well as in cell death. Using giant unilamellar vesicles and a chlorin photosensitizer, we asymmetrically oxidized the outer or inner monolayers of lipid membranes. We observed different shape transitions such as oblate to prolate and budding, which are typical of membrane curvature modifications. The asymmetry of the shape transitions is in accordance with a lowered effective spontaneous curvature of the leaflet being targeted. We interpret this effect as a decrease in the preferred area of t…
Morphologies of genus-1 and 2 toroidal vesicles are studied numerically by dynamically triangulated membrane models and experimentally by confocal laser microscopy. Our simulation results reproduce shape transformations observed in our experiments well. At large reduced volumes of the genus-1 vesicles, obtained vesicle shapes agree with the previous theoretical prediction, in which axisymmetric shapes are assumed: double-necked stomatocyte, discoidal toroid, and circular toroid. However, for small reduced volumes, it is revealed that a non-axisymmetric discoidal toroid and handled discocyte ex…
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…
Stationary shapes of axisymmetric vesicles beyond lowest-energy configurations
本文在Helfrich模型框架内通过最小化弹性能探索囊泡形态的能量景观，发现受压囊泡可呈现需极点施加点力的高能纺锤状构型，并在内部压力较低时预测多叶形状，合理解释了均匀AC场中实验观察到的大囊泡纺锤形状。
Motivated by recent advances in vesicle engineering, we consider theoretically the locomotion of shape-changing bilayer vesicles at low Reynolds number. By modulating their volume and membrane composition, the vesicles can be made to change shape quasi-statically in thermal equilibrium. When the control parameters are tuned appropriately to yield periodic shape changes which are not time-reversible, the result is a net swimming motion over one cycle of shape deformation. For two classical vesicle models (spontaneous curvature and bilayer coupling), we determine numerically the sequence of vesi…
The shape transformations of fluid membranes induced by curved protein rods are studied using meshless membrane simulations. The rod assembly at low rod density induces a flat membrane tube and oblate vesicle. It is found that the polyhedral shapes are stabilized at high rod densities. The discrete shape transition between triangular and buckled discoidal tubes is obtained and their curvature energies are analyzed by a simple geometric model. For vesicles, triangular hosohedron and elliptic-disk shapes are formed in equilibrium, whereas tetrahedral and triangular prism shapes are obtained as m…
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…
The morphologies of genus-2 to -8 fluid vesicles are studied by using dynamically triangulated membrane simulations with area-difference elasticity. It is revealed that the alignments of the membrane pores alter the vesicle shapes and the types of shape transitions for the genus $g \ge 3$. At a high reduced volume, a stomatocyte with a circular alignment of $g+1$ pores continuously transforms into a discocyte with a line of $g$ pores with increasing intrinsic area difference. In contrast, at a low volume, a stomatocyte transforms into a ($g+1$)-hedral shape and subsequently exhibits a discrete…
We systematize and extend the description of vesicle growth and shape change using linear nonequilibrium thermodynamics. By restricting the study to shape changes from spheres to axisymmetric ellipsoids, we are able to give a consistent formulation which includes the lateral tension of the vesicle membrane. This allows us to generalize and correct a previous calculation. Our present calculations suggest that, for small growing vesicles, a prolate ellipsoidal shape should be favored over oblate ellipsoids, whereas for large growing vesicles oblates should be favored over prolates. The validity …
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 shape equation and linking conditions for a vesicle with two-phase domains are derived. We refine the conjecture on the general neck condition for the limit shape of a budding vesicle proposed by Jülicher and Lipowsky [Phys. Rev. Lett. \textbf{70}, 2964 (1993); Phys. Rev. E \textbf{53}, 2670 (1996)], and then we use the shape equation and linking conditions to prove that this conjecture holds not only for axisymmetric budding vesicles, but also for asymmetric ones. Our study reveals that the mean curvature at any point on the membrane segments adjacent to the neck satisfies the general nec…
Cellular membranes exhibit a large variety of shapes, strongly coupled to their function. Many biological processes involve dynamic reshaping of membranes, usually mediated by proteins. This interaction works both ways: while proteins influence the membrane shape, the membrane shape affects the interactions between the proteins. To study these membrane-mediated interactions on closed and anisotropically curved membranes, we use colloids adhered to ellipsoidal membrane vesicles as a model system. We find that two particles on a closed system always attract each other, and tend to align with the…
The interaction of two types of vesicle systems was investigated: micrometer-sized, giant unilamellar vesicles (GUVs) formed from 1-palmitoyl-2-oleoyl-sn-glycero-3-phosphocholine (POPC) and submicrometer-sized, large unilamellar vesicles (LUVs) formed from oleic acid and oleate, both in a buffered aqueous solution (pH=8.8). Individual POPC GUVs were transferred with a micropipette into a suspension of oleic acid/oleate LUVs, and the shape changes of the GUVs were monitored using optical microscopy. The behavior of POPC GUVs upon transfer into a 0.8 mM suspension of oleic acid, in which oleic a…
We present a numerical study of the time-dependent motion of a two-dimensional vesicle in a channel under an imposed flow. In a Poiseuille flow the shape of the vesicle depends on the flow strength, the mechanical properties of the membrane, and the width of the channel as reported in the past. This study is focused on the centered snaking (CSn) shape, where the vesicle shows an oscillatory motion like a swimmer flagella even though the flow is stationary. We quantify this behavior by the amplitude and frequency of the oscillations of the vesicle's center of mass. We observe regions in paramet…
pH gradient-driven deformation of a crista-like vesicle
本文用Helfrich模型研究pH梯度驱动的嵴状囊泡形变，通过引入pH依赖自发曲率并构建功能性相图，揭示了嵴高效合成ATP的形态条件。
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 …
Amoeboid swimming of active vesicles
研究弱放气活性囊泡形状动力学与迁移，通过局部膜不可压缩性及小过剩面积假设分析其在自由空间的耦合动力学，依活性强度和频率，囊泡呈现多种动力学状态。
Dynamic shapes of floppy vesicles enclosing active Brownian particles with membrane adhesion
本文数值研究了包含活性布朗粒子且粒子与膜粘附的活性囊泡的动态行为，发现ABP活性和粒子体积分数共同决定囊泡形状相变，膜粘附显著改变囊泡行为并提供额外控制参数。
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, …
The assembly of curved protein rods on fluid membranes is studied using implicit-solvent meshless membrane simulations. As the rod curvature increases, the rods on a membrane tube assemble along the azimuthal direction first and subsequently along the longitudinal direction. Here, we show that both transition curvatures decrease with increasing rod stiffness. For comparison, curvature-inducing isotropic inclusions are also simulated. When the isotropic inclusions have the same bending rigidity as the other membrane regions, the inclusions are uniformly distributed on the membrane tubes and ves…
Soft bodies flowing in a channel often exhibit parachute-like shapes usually attributed to an increase of hydrodynamic constraint (viscous stress and/or confinement). We show that the presence of a fluid membrane leads to the reverse phenomenon and build a phase diagram of shapes --- which are classified as bullet, croissant and parachute --- in channels of varying aspect ratio. Unexpectedly, shapes are relatively wider in the narrowest direction of the channel. We highlight the role of flow patterns on the membrane in this response to the asymmetry of stress distribution.
Eukaryote cells have a flexible shape, which dynamically changes according to the function performed by the cell. One mechanism for deforming the cell membrane into the desired shape is through the expression of curved membrane proteins. Furthermore, these curved membrane proteins are often associated with the recruitment of the cytoskeleton, which then applies active forces that deform the membrane. This coupling between curvature and activity was previously explored theoretically in the linear limit of small deformations, and low dimensionality. Here we explore the unrestricted shapes of ves…
Despite the recent upsurge of theoretical reduced models for vesicle shape dynamics, comparisons with experiments have not been accomplished. We review the implications of some of the recently proposed models for vesicle dynamics, especially the Tumbling-Trembling domain regions of the phase plane and show that they all fail to capture the essential behavior of real vesicles for excess areas, Δ, greater than 0.4. We emphasize new observations of shape harmonics and the role of thermal fluctuations.
Sculpting vesicles with active particles: Less is more
本研究通过实验与模拟结合，发现将自推进粒子包裹在巨型囊泡中，在低粒子负载下可产生最显著的树突状等非平衡形状变化，揭示了局部内部力驱动细胞膜强变形的条件。
Coupling Anisotropic Curvature and Nematic Order: Mechanisms of Membrane Shape Remodeling
本研究理论探究各向异性弯曲膜组分通过曲率感知、向列作用等调控囊泡形态的机制，对比弧形与鞍形组分的不同效应，揭示拓扑缺陷定位规律，建立膜形状重塑的统一理解框架。
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…
Red blood cells (RBCs) are the major component of blood and the flow of blood is dictated by that of RBCs. We employ vesicles, which consist of closed bilayer membranes enclosing a fluid, as a model system to study the behavior of RBCs under a confined Poiseuille flow. We extensively explore two main parameters: i) the degree of confinement of vesicles within the channel, and ii) the flow strength. Rich and complex dynamics for vesicles are revealed ranging from steady-state shapes (in the form of parachute and slipper) to chaotic dynamics of shape. Chaos occurs through a cascade of multiple p…
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 …
As a model of soft confinement for polymers, we investigated equilibrium shapes of a flexible vesicle that contains a phase-separating polymer solution. To simulate such a system, we combined the phase field theory (PFT) for the vesicle and the self-consistent field theory (SCFT) for the polymer solution. We observed a transition from a symmetric prolate shape of the vesicle to an asymmetric pear shape induced by the domain structure of the enclosed polymer solution. Moreover, when a non-zero spontaneous curvature of the vesicle is introduced, a re-entrant transition between the prolate and th…
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…
Vesicles are soft elastic bodies with distinctive mechanical properties such as bending resistance, membrane fluidity, and their strong ability to deform, mimicking some properties of biological cells. While previous three-dimensional (3D) studies have identified stationary shapes such as slipper and axisymmetric ones, we report a complete phase diagram of 3D vesicle dynamics in a bounded Poiseuille flow with two more oscillatory dynamics, 3D snaking and swirling. 3D snaking is characterized by planar oscillatory motion of the mass center and shape deformations, which is unstable and leads to …
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…
A minimal model for poration induced electro deformation of Giant Vesicles
本研究提出一个极简解析模型，通过耦合电穿孔、电流体动力学与膜力学，揭示了强脉冲直流电场下巨型单层囊泡因膜穿孔产生额外表面积而呈现非椭球形的复杂变形机制。
Vesicles with internal active filaments: self-organized propulsion controls shape, motility, and dynamical response
本文提出“flexocytes”可变形囊泡模型，通过内部自推进丝的相互作用驱动膜形变，模拟生物细胞运动，并预测其在微图案基底和外部刺激下的动态响应，为软物质机器人构建提供新思路。
Unravelling the physical mechanisms behind the organisation of lipid domains is a central goal in cell biology and membrane biophysics. Previous studies on cells and model lipid bilayers featuring phase-separated domains found an intricate interplay between the membrane geometry and its chemical composition. However, the lack of a model system with simultaneous control over the membrane shape and conservation of its composition precluded a fundamental understanding of curvature-induced effects. Here, we present a new class of multicomponent vesicles supported by colloidal scaffolds of designed…
The Naito--Okuda--Ouyang Biconcave Red Blood Cell Profile as a Finite-Energy Unbranched Weak Immersion with Point-Force Residues
该研究分析 Naito--Okuda--Ouyang 双凹红细胞轮廓，证明其为有限能量弱浸入但非自由弱临界点，需添加点力势或固定点约束恢复平稳性，并澄清符号差异。
The growth, form, and division of prebiotic vesicles, membraneous bags of fluid of varying components and shapes is hypothesized to have served as the substrate for the origin of life. The dynamics of these out-of-equilibrium structures is controlled by physicochemical processes that include the intercalation of amphiphiles into the membrane, fluid flow across the membrane, and elastic deformations of the membrane. To understand prebiotic vesicular forms and their dynamics, we construct a minimal model that couples membrane growth, deformation, and fluid permeation, ultimately couched in terms…
Ohta-Kawasaki Model Reveals Patterns on Multicomponent Vesicles
本研究提出结合弹性弯曲模型与Ohta-Kawasaki模型的力化学框架，通过谱方法模拟多组分囊泡膜的耦合动力学，复现实验观测形态，为相关研究提供新见解与平台。
The role of membrane viscosity in the dynamics of fluid membranes
研究流体囊泡形态变化中膜内禀粘度与周围流体粘度的竞争机制，通过数值计算发现小囊泡中膜耗散主导且弛豫时间与尺寸平方成正比，大囊泡中体相耗散主导且弛豫时间与尺寸立方成正比。
We analyze the non-equilibrium shape fluctuations of giant unilamellar vesicles encapsulating motile bacteria. Owing to bacteria--membrane collisions, we experimentally observe a significant increase in the magnitude of membrane fluctuations at low wave numbers, compared to the well-known thermal fluctuation spectrum. We interrogate these results by numerically simulating membrane height fluctuations via a modified Langevin equation, which includes bacteria--membrane contact forces. Taking advantage of the length and time scale separation of these contact forces and thermal noise, we further c…
Only some special open surfaces satisfying the shape equation of lipid membranes can be compatible with the boundary conditions. As a result of this compatibility, the first integral of the shape equation should vanish for axisymmetric lipid membranes, from which two theorems of non-existence are verified: (i) There is no axisymmetric open membrane being a part of torus satisfying the shape equation; (ii) There is no axisymmetric open membrane being a part of a biconcave discodal surface satisfying the shape equation. Additionally, the shape equation is reduced to a second-order differential e…
Vesicles and many biological membranes are made of two monolayers of lipid molecules and form closed lipid bilayers. The dynamical behaviour of vesicles is very complex and a variety of forms and shapes appear. Lipid bilayers can be considered as a surface fluid and hence the governing equations for the evolution include the surface (Navier--)Stokes equations, which in particular take the membrane viscosity into account. The evolution is driven by forces stemming from the curvature elasticity of the membrane. In addition, the surface fluid equations are coupled to bulk (Navier--)Stokes equatio…
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