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Register a new userMembrane Stretching Elasticity of Lipid Vesicle and Thermal Shape Fluctuations
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One of the most widely used methods for determination of the bending elasticity modulus of model lipid membranes is the analysis of the shape fluctuations of nearly spherical lipid vesicles. The theoretical basis of this analysis is given by Milner and Safran. In their theory the stretching effects are not considered. In the present study we generalized their approach including the stretching effects deduced after an application of statistical mechanics of vesicles.
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The mechanical properties of biological membranes play an important role in the structure and the functioning of living organisms. One of the most widely used methods for determination of the bending elasticity modulus of the model lipid membranes (simplified models of the biomembranes with similar mechanical properties) is analysis of the shape fluctuations of the nearly spherical lipid vesicles. A theoretical basis of such an analysis is developed by Milner and Safran. In the present studies we analyze their results using an approach based on the Bogoljubov inequalities and the approximating Hamiltonian method. This approach is in accordance with the principles of statistical mechanics and is free of contradictions. Our considerations validate the results of Milner and Safran if the stretching elasticity K_s of the membrane tends to zero.
We have studied the mesoscopic shape fluctuations in aligned multilamellar stacks of DMPC bilayers using the neutron spin-echo technique. The corresponding in plane dispersion relation $tau^{-1}$(q$_{||}$) at different temperatures in the gel (ripple, P$_{beta}$) and the fluid (L$_{alpha}$) phase of this model system has been determined. Two relaxation processes, one at about 10ns and a second, slower process at about 100ns can be quantified. The dispersion relation in the fluid phase is fitted to a smectic hydrodynamic theory, with a correction for finite q$_z$ resolution. We extract values for, the bilayer bending rigidity $kappa$, the compressional modulus of the stacks $B$, and the effective sliding viscosity $eta_3$. The softening of a mode which can be associated with the formation of the ripple structure is observed close to the main phase transition.
Can the presence of molecular-tilt order significantly affect the shapes of lipid bilayer membranes, particularly membrane shapes with narrow necks? Motivated by the propensity for tilt order and the common occurrence of narrow necks in the intermediate stages of biological processes such as endocytosis and vesicle trafficking, we examine how tilt order inhibits the formation of necks in the equilibrium shapes of vesicles. For vesicles with a spherical topology, point defects in the molecular order with a total strength of $+2$ are required. We study axisymmetric shapes and suppose that there is a unit-strength defect at each pole of the vesicle. The model is further simplified by the assumption of tilt isotropy: invariance of the energy with respect to rotations of the molecules about the local membrane normal. This isotropy condition leads to a minimal coupling of tilt order and curvature, giving a high energetic cost to regions with Gaussian curvature and tilt order. Minimizing the elastic free energy with constraints of fixed area and fixed enclosed volume determines the allowed shapes. Using numerical calculations, we find several branches of solutions and identify them with the branches previously known for fluid membranes. We find that tilt order changes the relative energy of the branches, suppressing thin necks by making them costly, leading to elongated prolate vesicles as a generic family of tilt-ordered membrane shapes.
Vesicles prepared in water from a series of diblock copolymers and termed polymersomes are physically characterized. With increasing molecular weight $bar{M}_n$, the hydrophobic core thickness $d$ for the self-assembled bilayers of polyethyleneoxide - polybutadiene (PEO-PBD) increases up to 20 $nm$ - considerably greater than any previously studied lipid system. The mechanical responses of these membranes, specifically, the area elastic modulus $K_a$ and maximal areal strain $alpha_c$ are measured by micromanipulation. As expected for interface-dominated elasticity, $K_a$ ($simeq$ 100 $pN/nm$) is found to be independent of $bar{M}_n$. Related mean-field ideas also predict a limiting value for $alpha_c$ which is universal and about 10-fold above that typical of lipids. Experiments indeed show $alpha_c$ generally increases with $bar{M}_n$, coming close to the theoretical limit before stress relaxation is opposed by what might be chain entanglements at the highest $bar{M}_n$. The results highlight the interfacial limits of self-assemblies at the nano-scale.
Theoretical studies of nearly spherical vesicles and microemulsion droplets, that present typical examples for thermally-excited systems that are subject to constraints, are reviewed. We consider the shape fluctuations of such systems constrained by fixed area $A$ and fixed volume $V$, whose geometry is presented in terms of scalar spherical harmonics. These constraints can be incorporated in the theory in different ways. After an introductory review of the two approaches: with an exactly fixed by delta-function membrane area $A$ [Seifert, Z. Phys. B, 97, 299, (1995)] or approximatively by means of a Lagrange multiplier $sigma$ conjugated to $A$ [Milner and Safran, Phys. Rev. A, 36, 4371 (1987)], we discuss the determined role of the stretching effects, that has been announced in the framework of a model containing stretching energy term, expressed via the membrane vesicle tension [Bivas and Tonchev, Phys.Rev.E, 100, 022416 (2019)]. Since the fluctuation spectrum for the used Hamiltonian is not exactly solvable an approximating method based on the Bogoliubov inequalities for the free energy has been developed. The area constraint in the last approach appears as a self-consistent equation for the membrane tension. In the general case this equation is intractable analytically. However, much insight into the physics behind can be obtained either imposing some restrictions on the values of the model parameters, or studying limiting cases, in which the self-consistent equation is solved. Implications for the equivalence of ensembles have been discussed as well.
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