Motivated by the still ongoing debate about the various possible meanings of the term surface tension of bilayer membranes, we present here a detailed discussion that explains the differences between the "intrinsic", "renormalized", and "mechanical" tensions. We use analytical considerations and computer simulations to show that the membrane spectrum of thermal fluctuations is governed by the mechanical and not the intrinsic tension. Our study highlights the fact that the commonly used quadratic approximation of Helfrich effective Hamiltonian is not rotationally invariant. We demonstrate that this non-physical feature leads to a calculated mechanical tension that differs dramatically from the correct mechanical tension. Specifically, our results suggest that the mechanical and intrinsic tensions vanish simultaneously, which contradicts recent theoretical predictions derived for the approximated Hamiltonian.
Considering two rigid conical inclusions embedded in a membrane subject to lateral tension, we study the membrane-mediated interaction between these inclusions that originates from the hat-shaped membrane deformations associated with the cones. At non-vanishing lateral tensions, the interaction is found to depend on the orientation of the cones with respect to the membrane plane. The interaction of inclusions of equal orientation is repulsive at all distances between them, while the inclusions of opposite orientation repel each other at small separations, but attract each other at larger ones.…
We evaluate the effective Hamiltonian governing, at the optically resolved scale, the elastic properties of micro-manipulated membranes. We identify floppy, entropic-tense and stretched-tense regimes, representing different behaviors of the effective area-elasticity of the membrane. The corresponding effective tension depends on the microscopic parameters (total area, bending rigidity) and on the optically visible area, which is controlled by the imposed external constraints. We successfully compare our predictions with recent data on micropipette experiments.
By using the grand canonical Monte Carlo simulations on spherical surfaces with two fixed vertices separated by the distance L, we find that the second-order phase transition changes to the first-order one when L is sufficiently large. We find that string tension σ\not= 0 in the smooth phase while σ\to 0 in the wrinkled phase.
We study the nucleation of a single pore in a fluctuating lipid membrane, specifically taking into account the membrane fluctuations, as well as the shape fluctuations of the pore. For large enough pores, the nucleation free energy is well-described by shifts in the effective membrane surface tension and the pore line tension. Using our framework, we derive the stability criteria for the various pore formation regimes. In addition to the well-known large-tension regime from the classical nucleation theory of pores, we also find a low-tension regime in which the effective line and surface tensi…
Tension in lipid membranes is often controlled externally, by pulling on the boundary of the membrane or changing osmotic pressure across a curved membrane. But modifications of the tension can also be induced in an internal fashion, for instance as a byproduct of changing a membranes electric potential or, as observed experimentally, by activity of membrane proteins. Here we develop a theory which demonstrate how such internal contributions to the tension are moderated through elastic stretching of the membrane when the membrane is initially in a low tension floppy state.
Solid sheets and fluid membranes exhibit buckling under lateral compression. Here, it is revealed that fluid membranes have anisotropic buckling surface tension contrary to solid sheets. Surprisingly, the surface tension perpendicular to the buckling direction shows stronger dependence than that parallel to it. Our theoretical predictions are supported by numerical simulations of a meshless membrane model. This anisotropic tension can be used to measure the membrane bending rigidity. It is also found phase synchronization occurs between multilayered buckled membranes.
Adsorption of proteins onto membranes can alter the local membrane curvature. This phenomenon has been observed in biological processes such as endocytosis, tubulation and vesiculation. However, it is not clear how the local surface properties of the membrane, such as membrane tension, change in response to protein adsorption. In this paper, we show that the classical elastic model of lipid membranes cannot account for simultaneous changes in shape and membrane tension due to protein adsorption in a local region, and a viscous-elastic formulation is necessary to fully describe the system. Ther…
Recent progress in the fusion of biological membranes is reviewed to highlight the central role played by the line tension, which permits exquisite control of the process.
In recent years it has been argued that the tension parameter driving the fluctuations of fluid membranes, differs from the imposed lateral stress, the 'frame tension'. In particular, stress-free membranes were predicted to have a residual fluctuation tension. In the present paper, this argument is reconsidered and shown to be inherently inconsistent -- in the sense that a linearized theory, the Monge model, is used to predict a nonlinear effect. Furthermore, numerical simulations of one-dimensional stiff membranes are presented which clearly demonstrate, first, that the internal 'intrinsic' s…
We use computer simulations and a simple free energy model to study the response of a bilayer membrane to the application of a negative (compressive) mechanical tension. Such a tension destabilizes the long wavelength undulation modes of giant vesicles, but it can be sustained when small membranes and vesicles are considered. Our negative tension simulation results reveal two regimes - (i) a weak negative tension regime characterized by stretching-dominated elasticity, and (ii) a strong negative tension regime featuring bending-dominated elastic behavior. This resembles the findings of the cla…
We propose a mechanism for mechanical regulation at the membrane of living cells, based on the exchange of membrane area between the cell membrane and a membrane reservoir. The reservoir is composed of invaginated membrane microdomains which are liable to flatten upon increase of membrane strain, effectively controlling membrane tension. We show that the domain shape transition is first order, allowing for coexistence between flat and invaginated domains. During coexistence, the membrane tension is controlled by the domains elasticity and by the kinetics of the shape transition. We show that t…
In this paper, we study the boundary effect on the surface (or frame) tension of elastic membrane surface models. The frame tension generally depends only on the projected area of the boundary over which the surface spans. However, from a spin model analogy, the frame tension is expected to be dependent also on the boundary shape at the continuous transition point. We confirm this expectation using the following fixed-connectivity and tethered surface models: the surface model of Helfrich and Polyakov and a surface model with deficit angle term. We also discuss the reason why this expectation …
Branching of bilayer membranes appear in the inverted hexagonal phase as well as in metastable states of the lamellar phase such as membrane fusion intermediates. A method for estimating the line tension of the branching junction is proposed for molecular simulations. The line tension is calculated from the pressure tensor of equiangularly branched membranes. The simulation results agree very well with the theoretical prediction of Hamm and Kozlov's tilt model. The transition between the lamellar and inverted hexagonal phases is also investigated using the tilt model.
We identify a class of composite membranes: fluid bilayers coupled to an elastic meshwork, that are such that the meshwork's energy is a function $F_\mathrm{el}[A_ξ]$ \textit{not} of the real microscopic membrane area $A$, but of a \textit{smoothed} membrane's area $A_ξ$, which corresponds to the area of the membrane coarse-grained at the mesh size $ξ$. We show that the meshwork modifies the membrane tension $σ$ both below and above the scale $ξ$, inducing a tension-jump $Δσ=dF_\mathrm{el}/dA_ξ$. The predictions of our model account for the fluctuation spectrum of red blood cells membranes cou…
Membranes containing a wide variety of ternary mixtures of high chain-melting temperature lipids, low chain-melting temperature lipids, and cholesterol undergo lateral phase separartion into coexisting liquid phases at a miscibility transition. When membranes are prepared from a ternary lipid mixture at a critical composition, they pass through a miscibility critical point at the transition temperature. Since the critical temperature is typically on the order of room temperature, membranes provide an unusual opportunity in which to perform a quantitative study of biophysical systems that exhib…
We discuss the fluctuation-induced force, a finite-temperature analog of the Casimir force, between two inclusions embedded in a fluid membrane under tension. We suggest a method to calculate this Casimir interaction in the most general case, where membrane fluctuations are governed by the combined action of surface tension, bending modulus, and the Gaussian rigidity. We find that the surface tension strongly modifies the power law in the separation dependence of the Casimir interaction. This results in a strong suppression of the Casimir force at separations beyond a characteristic length, wh…
We study the elastic response of bilayer membranes with fixed projected area to both stretching and shape deformations. A surface tension is associated to each of these deformations. By using model amphiphilic membranes and computer simulations, we are able to observe both the types of deformation, and thus, both the surface tensions, related to each type of deformation, are measured for the same system. These surface tensions are found to assume different values in the same bilayer membrane: in particular they vanish for different values of the projected area. We introduce a simple theory whi…
Up-down asymmetric inclusions impose a local, spontaneous curvature to an elastic membrane. When several of them are inserted in a same membrane, they feel effective forces mediated by the membrane, both of elastic and entropic nature. Following an approach initiated by Dommersnes and Fournier in the vanishing tension case [Eur. Phys. J. B 12, 9 (1999)], and also using a pseudo-analytical micellization theory, we derive the statistical mechanics of asymmetric inclusion assemblies when they are also subject to an additional short-range, attractive interaction. Our main conclusion is that generi…
How long a fluid membrane vesicle stressed with a steady ramp of micropipette last before rupture? Or conversely, how high the surface tension should be to rupture a membrane? To answer these challenging questions we have developed a theoretical framework that allows description and reproduction of Dynamic Tension Spectroscopy (DTS) observations. The kinetics of the membrane rupture under ramps of surface tension is described as a combination of initial pore formation followed by Brownian process of the pore radius crossing the time-dependent energy barrier. We present the formalism and derive…
Three types of surface tensions can be defined for lipid membranes: the internal tension, $σ$, conjugated to the real membrane area in the Hamiltonian, the mechanical frame tension, $τ$, conjugated to the projected area, and the "fluctuation tension", $r$, obtained from the fluctuation spectrum of the membrane height. We investigate these surface tensions by means of a Monge gauge lattice Monte Carlo simulation involving the exact, nonlinear, Helfrich Hamiltonian and a measure correction for excess entropy of the Monge gauge. Our results for the relation between $σ$ and $τ$ agrees well with th…
We present a model for pore stabilization in membranes without surface tension. Whereas an isolated pore is always unstable (since it either shrinks tending to re-seal or grows without bound til to membrane disintegration), it is shown that excluded volume interactions in a system of many pores can stabilize individual pores of a given size in a certain range of model parameters. For such a multipore membrane system, the distribution of pore size and associated pore lifetime are calculated within the mean field approximation. We predict that, above certain temperature when the effective line t…
It has been known for long that the fluctuation surface tension of membranes $r$, computed from the height fluctuation spectrum, is not equal to the bare surface tension $σ$ introduced in the Helfrich theory. In this work we relate these two surface tensions both analytically and numerically and compare them to the Laplace tension $γ$, and the mechanical frame tension $τ$. Using one-loop renormalisation calculations, we obtain, in addition to the effective bending modulus $κ_{\rm eff}$, a new expression for the effective surface tension $σ_{\rm eff}=σ- εk_{\rm B}T/(2a_p)$ where $a_p$ the proje…
We report a new and facile method for measuring edge tensions of lipid membranes. The approach is based on electroporation of giant unilamellar vesicles and analysis of the pore closure dynamics. We applied this method to evaluate the edge tension in membranes with four different compositions: egg phosphatidylcholine (EggPC), dioleoylphosphatidylcholine (DOPC), and mixtures of the latter with cholesterol and dioleoylphosphatidylethanolamine (DOPE). Our data confirm previous results for EggPC and DOPC. The addition of 17 mol % cholesterol to the DOPC membrane causes an increase in the membrane …
The effect of AC electric fields on the elasticity of supported lipid bilayers has been investigated at the microscopic level using grazing incidence synchrotron x-ray scattering. A strong decrease in the membrane tension up to 1mN/m and a dramatic increase of its effective rigidity up to 300kBT are observed for local electric potentials seen by the membrane < 1V. The experimental results were analyzed using detailed electrokinetic modeling and non-linear Poisson-Boltzmann theory. Based on a modeling of the electromagnetic stress which provides an accurate description of bilayer separation vs …
The free energy of inserting a protein into a membrane is determined by considering the variation in the spectrum of thermal fluctuations in response to the presence of a rigid inclusion. Both numerically and through a simple analytical approximation, we find that the primary effect of fluctuations is to reduce the effective surface tension, hampering the insertion at low surface tension. Our results, which should also be relevant for membrane pores, suggest (in contrast to classical nucleation theory) that a finite surface tension is necessary to facilitate the opening of a pore.
We present measurements of the fluctuation spectrum of giant vesicles containing bacteriorhodopsin (BR) pumps using video-microscopy. When the pumps are activated, we observe a significant increase of the fluctuations in the low wavevector region, which we interpret as due to a lowering of the effective tension of the membrane.
Two different tensions can be defined for a fluid membrane: the internal tension, $γ$, conjugated to the real membrane area in the Hamiltonian, and the frame tension, $τ$, conjugated to the projected (or frame) area. According to the standard statistical description of a membrane, the fluctuation spectrum is governed by $γ$. However, using rotational invariance arguments, several studies argued that fluctuation spectrum must be governed by the frame tension $τ$ instead. These studies disagree on the origin of the result obtained with the standard description yet: either a miscounting of config…
In clathrin-mediated endocytosis (CME), clathrin and various adaptor proteins coat a patch of the plasma membrane, which is reshaped to form a budded vesicle. Experimental studies have demonstrated that elevated membrane tension can inhibit bud formation by a clathrin coat. In this study, we investigate the impact of membrane tension on the mechanics of membrane budding by simulating clathrin coats that either grow in area or progressively induce greater curvature. At low membrane tension, progressively increasing the area of a curvature-generating coat causes the membrane to smoothly evolve f…
In the framework of a phase field model of a single cell crawling on a substrate, we investigate how the properties of the cell membrane affect the shape and motility of the cell. Since the membrane influences the cell dynamics on multiple levels and provides a nontrivial feedback, we consider the following fundamental interactions: (i) the reduction of the actin polymerization rate by membrane tension; (ii) area conservation of the cell's two-dimensional cross-section vs. conservation of its circumference (i.e. membrane inextensibility); and (iii) the contribution from the membrane's bending …
We investigate the phenomenon of protein induced tubulation of lipid bilayer membranes within a continuum framework using Monte Carlo simulations coupled with the Widom insertion technique to compute excess chemical potentials. Tubular morphologies are spontaneously formed when the density and the curvature-field strength of the membrane bound proteins exceed their respective thresholds and this transition is marked by a sharp drop in the excess chemical potential. We find that the planar to tubular transition can be described by a micellar model and that the corresponding free energy barrier …
The function of membrane-embedded proteins such as ion channels depends crucially on their conformation. We demonstrate how conformational changes in asymmetric membrane proteins may be inferred from measurements of their diffusion. Such proteins cause local deformations in the membrane, which induce an extra hydrodynamic drag on the protein. Using membrane tension to control the magnitude of the deformations and hence the drag, measurements of diffusivity can be used to infer--- via an elastic model of the protein--- how conformation is changed by tension. Motivated by recent experimental res…
We calculate the fluctuation spectrum of the shape of a lipid vesicle or cell exposed to a nonthermal source of noise. In particular we take into account constraints on the membrane area and the volume of fluid that it encapsulates when obtaining expressions for the dependency of the membrane tension on the noise. We then investigate three possible origins of the non-thermal noise taken from the literature: A direct force, which models an external medium pushing on the membrane. A curvature force, which models a fluctuating spontaneous curvature, and a permeation force coming from an active tr…
We study the force generation by a set of parallel actin filaments growing against an elastic membrane. The elastic membrane tries to stay flat and any deformation from this flat state, either caused by thermal fluctuations or due to protrusive polymerization force exerted by the filaments, costs energy. We study two lattice models to describe the membrane dynamics. In one case, the energy cost is assumed to be proportional to the absolute magnitude of the height gradient (gradient model) and in the other case it is proportional to the square of the height gradient (Gaussian model). For the gr…
We consider a one-dimensional elastic membrane, which is pushed by growing filaments. The filaments tend to grow by creating local protrusions in the membrane and this process has surface energy and bending energy costs. Although it is expected that with increasing surface tension and bending rigidity, it should become more difficult to create a protrusion, we find that for a fixed bending rigidity, as the surface tension increases, protrusions are more easily formed. This effect also gives rise to nontrivial dependence of membrane velocity on the surface tension, characterized by a dip and a …
The effect of asymmetry in the transbilayer lipid distribution on the dynamics of phase separation in fluid vesicles is investigated numerically for the first time. This asymmetry is shown to set a spontaneous curvature for the domains that alter the morphology and dynamics considerably. For moderate tension, the domains are capped and the spontaneous curvature leads to anomalously slow dynamics, as compared to the case of symmetric bilayers. In contrast, in the limiting cases of high and low tensions, the dynamics proceeds towards full phase separation.
The elastic properties of a self-assembled bilayer membrane are studied using the self-consistent field theory, applied to a model system composed of flexible amphiphilic chains dissolved in hydrophilic polymeric solvents. Examining the free energy of bilayer membranes with different geometries allows us to calculate their bending modulus, Gaussian modulus, two fourth-order membrane moduli, and the line tension. The dependence of these parameters on the microscopic characteristics of the amphiphilic chain, characterized by the volume fraction of the hydrophilic component, is systematically stu…
Membrane tubes (spicules) arise in cells, or artificial membranes, in the nonlinear deformation regime due to, e.g. the growth of microtubules, actin filaments or sickle hemoglobin fibers towards a membrane. We calculate the axial force exerted by the cylindrical membrane tube, and its average radius, by taking into account steric interactions between the fluctuating membrane and the enclosed rod. The force required to confine a fluctuating membrane near the surface of the enclosed rod diverges as the separation approaches zero. This results in a smooth crossover of the axial force between a s…
Lipid membranes constitute very particular materials: on the one hand, they break very easily under microscopical stretching; on the other hand, they are extremely flexible, presenting deformations even at small scales. Consequently, a piece of membrane has an area excess relative to its optically resolvable area, also called projected area. From a mechanical point of view, we can thus identify three tensions associated to lipid membranes: the mechanical effective tension $τ$, associated to an increase in the projected area and to the flattening of the fluctuations; the tension $σ$, associated…
Atomically thin sheets, such as graphene, are widely used in nanotechnology. Recently they have also been used in applications including kirigami and self-folding origami, where it becomes important to understand how they respond to external loads. Motivated by this, we investigate how isotropic sheets respond to uniaxial tension by employing the self-consistent screening analysis method and molecular dynamics simulations. Previously, it was shown that for freely suspended sheets thermal fluctuations effectively renormalize elastic constants, which become scale-dependent beyond a characteristi…
The buckling of elastic bodies is a common phenomenon in the mechanics of solids. Wrinkling of membranes can often be interpreted as buckling under constraints that prohibit large amplitude deformation. We present a combination of analytic calculations, experiments, and simulations to understand wrinkling patterns generated in a bilayer membrane. The model membrane is composed of a flexible spherical shell that is under tension and that is circumscribed by a stiff, essentially incompressible strip with bending modulus B. When the tension is reduced sufficiently to a value σ, the strip forms wr…
A Monte Carlo (MC) study is performed to evaluate the surface tension $γ$ of spherical membranes that may be regarded as the models of the lipid layers. We use the canonical surface model defined on the self-avoiding triangulated lattices. The surface tension $γ$ is calculated by keeping the total surface area $A$ constant during the MC simulations. In the evaluation of $γ$, we use $A$ instead of the projected area $A_p$, which is unknown due to the fluctuation of the spherical surface without boundary. The pressure difference ${\itΔ}p $ between the inner and the outer sides of the surface is …
The adhesion bonds connecting a lipid bilayer to an underlying surface may undergo a condensation transition resulting from an interplay between a short range attractive potential between them, and a long range fluctuation-induced potential of mean force. Here, we use computer simulations of a coarse-grained molecular model of supported lipid bilayers to study this transition in confined membranes, and in membranes subjected to a non-vanishing surface tension. Our results show that confinement may alter significantly the condensation transition of the adhesion bonds, whereas the application of…
We investigate the effect of an applied electric potential on the mechanics of a coarse grained POPC bilayer under tension. The size and duration of our simulations allow for a detailed and accurate study of the fluctuations. Effects on the fluctuation spectrum, tension, bending rigidity, and bilayer thickness are investigated in detail. In particular, the least square fitting technique is used to calculate the fluctuation spectra. The simulations confirm a recently proposed theory that the effect of an applied electric potential on the membrane will be moderated by the elastic properties of t…
We consider the influence of electrostatic forces on driven translocation dynamics of a flexible polyelectrolyte being pulled through a nanopore by an external force on the head monomer. To this end, we augment the iso-flux tension propagation (IFTP) theory with electrostatics for a negatively charged biopolymer pulled through a nanopore embedded in a similarly charged anionic membrane. We show that for the realistic case such as a single-stranded DNA, the translocation dynamics at low salt where screening is weak and at finite negative membrane charge is unexpectedly accelerated despite the l…
We review recent computer simulation studies of undulating lipid bilayers. Theoretical interpretations of such fluctuating membranes are most commonly based on generalized Helfrich-type elastic models, with additional contributions of local "protrusions" and/or density fluctuations. Such models provide an excellent basis for describing the fluctuations of tensionless bilayers in the fluid phase at a quantitative level. However, this description is found to fail for membranes in the gel phase and for membranes subject to high tensions. The fluctuations of tilted gel membranes show a signature o…
In lieu of abstract, first paragraph reads: Galimzyanov et al. [1] find that line tension between thick liquid-ordered ($L_{o}$) and thinner liquid-disordered ($L_{d}$) registered lipid bilayer phases is minimised by an asymmetric "slip region", length $L\!\sim\!5\,\textrm{nm}$ (Fig. 1). They claim that line tensions alone explain domain registration, without "direct" (area-dependent) inter-leaflet interaction [2,3]. We show this is unfounded, without direct interaction their results would predict \textit{antiregistration}, dependent on composition. To find equilibrium from line energies, line…
Motivated by a long-standing debate concerning the nature and interrelations of surface-tension variables in fluid membranes, we reformulate the thermodynamics of a membrane vesicle as a generic two-dimensional finite system enclosing a three-dimensional volume. The formulation is shown to require two tension variables, conjugate to the intensive constraints of area per molecule and volume-to-area ratio. We obtain the relation between these two variables in various scenarios, as well as their correspondence to other definitions of tension variables for membranes. Several controversies related …
We report on the modeling of the dynamics of confined lipid membranes. We derive a thin film model in the lubrication limit which describes an inextensible liquid membrane with bending rigidity confined between two adhesive walls. The resulting equations share similarities with the Swift-Hohenberg model. However, inextensibility is enforced by a time-dependent nonlocal tension. Depending on the excess membrane area available in the system, three different dynamical regimes, denoted as A, B and C, are found from the numerical solution of the model. In regime A, membranes with small excess area …
In practice, physical membranes are exposed to a certain amount of external strain (tension or compression), due to the environment where they are placed. As a result, the behavior of the phonon modes of the membrane is modified. We show that anharmonic effects in stiff two-dimensional membranes are highly suppressed under the application of tension. For this, we consider the anharmonic coupling between bending and stretching modes in the self-consistent screening approximation (SCSA), and compare the obtained height-height correlation function in the SCSA to the corresponding harmonic propaga…
Recent measurements of the short-wavelength (~ 1--100 nm) fluctuations in stacks of lipid membranes have revealed two distinct relaxations: a fast one (decay rate of ~ 0.1 ns^{-1}), which fits the known baroclinic mode of bulk lamellar phases, and a slower one (~ 1--10 μs^{-1}) of unknown origin. We show that the latter is accounted for by an overdamped capillary mode, depending on the surface tension of the stack and its anisotropic viscosity. We thereby demonstrate how the dynamic surface tension of membrane stacks could be extracted from such measurements.
The astronomically observed value of the cosmological constant is small but non-zero. This raises two questions together known as the cosmological constant problem a) why is lambda so nearly zero? b) why is lambda not EXACTLY zero? Sorkin has proposed that b) can be naturally explained as a one by square root N fluctuation by invoking discreteness of spacetime at the Planck scale due to quantum gravity. In this paper we shed light on these questions by developing an analogy between the cosmological constant and the surface tension of membranes. The ``cosmological constant problem'' has a natur…
Recent research has shown that motile cells can adapt their mode of propulsion to the mechanical properties of the environment in which they find themselves--crawling in some environments while swimming in others. The latter can involve movement by blebbing or other cyclic shape changes, and both highlysimplified and more realistic models of these modes have been studied previously. Herein we study swimming that is driven by membrane tension gradients that arise from flows in the actin cortex underlying the membrane, and does not involve imposed cyclic shape changes. Such gradients can lead to…
We study the shapes of elastic membranes under the simultaneous exertion of tensile and compressive forces when the translational symmetry along the tension direction is broken. We predict a multitude of novel morphological phases in various regimes of a 2-dimensional parameter space $(ε,ν)$ that defines the relevant mechanical and geometrical conditions. Theses parameters are, respectively, the ratio between compression and tension, and the wavelength contrast along the tension direction. In particular, our theory associates the repetitive increase of pattern periodicity, recently observed on…
We study the effects of thermal fluctuations on symmetric tensionless heterogeneous (two-component) fluid membranes in a simple minimal model. Close to the critical point $T_c$ of the associated miscibility phase transition of the composition and for sufficiently strong curvature-composition interactions, mediated through a composition-dependent bending modulus, thermal fluctuations lead to enhancement of the effective bending modulus. Thus, the membrane conformation fluctuations will be {\em suppressed} near $T_c$, in comparison with a pure fluid membrane, for which thermal fluctuations are k…
