RESUMEN
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, which is introduced in the theory either as a Lagrange multiplier to conserve the total membrane area or as an external constraint. In this work we relate these two surface tensions both analytically and numerically. They are also compared to the Laplace tension γ, and the mechanical frame tension τ. Using the Helfrich model and one-loop renormalisation calculations, we obtain, in addition to the effective bending modulus κeff, a new expression for the effective surface tension σeff = σ - εkBT/(2ap) where kBT is the thermal energy, ap the projected cut-off area, and ε = 3 or 1 according to the allowed configurations that keep either the projected area or the total area constant. Moreover we show that the crumpling transition for an infinite planar membrane occurs for σeff = 0, and also that it coincides with vanishing Laplace and frame tensions. Using extensive Monte Carlo (MC) simulations, triangulated membranes of vesicles made of N = 100-2500 vertices are simulated within the Helfrich theory. As compared to alternative numerical models, no local constraint is applied and the shape is only controlled by the constant volume, the spontaneous curvature and σ. It is shown that the numerical fluctuation surface tension r is equal to σeff both with radial MC moves (ε = 3) and with corrected MC moves locally normal to the fluctuating membrane (ε = 1). For finite vesicles of typical size R, two different regimes are defined: a tension regime for [small sigma, Greek, circumflex]eff = σeffR2/κeff > 0 and a bending one for -1 < [small sigma, Greek, circumflex]eff < 0. A shape transition from a quasi-spherical shape imposed by the large surface energy, to more deformed shapes only controlled by the bending energy, is observed numerically at [small sigma, Greek, circumflex]eff ≃ 0. We propose that the buckling transition, observed for planar supported membranes in the literature, occurs for [small sigma, Greek, circumflex]eff ≃ -1, the associated negative frame tension playing the role of a compressive force. Hence, a precise control of the value of σeff in simulations cannot but enhance our understanding of shape transitions of vesicles and cells.
RESUMEN
A model of lipid bilayers made of a mixture of two lipids with different average compositions on both leaflets, is developed. A Landau Hamiltonian describing the lipid-lipid interactions on each leaflet, with two lipidic fields ψ 1 and ψ 2, is coupled to a Helfrich one, accounting for the membrane elasticity, via both a local spontaneous curvature, which varies as C 0 + C 1(ψ 1 - ψ 2/2), and a bending modulus equal to κ 0 + κ 1(ψ 1 + ψ 2)/2. This model allows us to define curved patches as membrane domains where the asymmetry in composition, ψ 1 - ψ 2, is large, and thick and stiff patches where ψ 1 + ψ 2 is large. These thick patches are good candidates for being lipidic rafts, as observed in cell membranes, which are composed primarily of saturated lipids forming a liquid-ordered domain and are known to be thick and flat nano-domains. The lipid-lipid structure factors and correlation functions are computed for globally spherical membranes and planar ones and for a whole set of parameters including the surface tension and the coupling in the two leaflet compositions. Phase diagrams are established, within a Gaussian approximation, showing the occurrence of two types of Structure Disordered phases, with correlations between either curved or thick patches, and an Ordered phase, corresponding to the divergence of the structure factor at a finite wave vector. The varying bending modulus plays a central role for curved membranes, where the driving force κ 1 C 0 (2) is balanced by the line tension, to form raft domains of size ranging from 10 to 100 nm. For planar membranes, raft domains emerge via the cross-correlation with curved domains. A global picture emerges from curvature-induced mechanisms, described in the literature for planar membranes, to coupled curvature- and bending-induced mechanisms in curved membranes forming a closed vesicle.
