No Arabic abstract
Lipid bilayer membranes undergo rapid bending undulations with wavelengths from tens of nanometers to tens of microns due to thermal fluctuations. Here, we probe such undulations and the membranes mechanics by measuring the time-varying orientation of single gold nanorods (GNRs) adhered to the membrane, using high-speed dark field microscopy. In a lipid vesicle, such measurements allow the determination of the membranes viscosity, bending rigidity and tension as well as the friction coefficient for sliding of the monolayers over one another. The in-plane rotation of the GNR is hindered by undulations in a membrane tension dependent manner, consistent with simulations. The motion of single GNRs adhered to the plasma membrane of living cultured cells similarly reveals that membranes complex physics and coupling to the cells actomyosin cortex.
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.
Inverse bicontinuous cubic phases are ubiquitous in lipid-water mixtures and consist of a lipid bilayer forming a cubic minimal surface, thereby dividing space into two cubic networks of water channels. For small hydrocarbon chain lengths, the monolayers can be modeled as parallel surfaces to a minimal midsurface. The bending energy of the cubic phases is determined by the distribution of Gaussian curvature over the minimal midsurfaces which we calculate for seven different structures (G, D, P, I-WP, C(P), S and F-RD). We show that the free-energy densities of the structures G, D and P are considerably lower than those of the other investigated structures due to their narrow distribution of Gaussian curvature. The Bonnet transformation between G, D, and P implies that these phases coexist along a triple line, which also includes an excess water phase. Our model includes thermal membrane undulations. Our qualitative predictions remain unchanged when higher order terms in the curvature energy are included. Calculated phase diagrams agree well with the experimental results for 2:1 lauric acid/dilauroyl phosphatidylcholine and water.
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.
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 study pore nucleation in a model membrane system, a freestanding polymer film. Nucleated pores smaller than a critical size close, while pores larger than the critical size grow. Holes of varying size were purposefully prepared in liquid polymer films, and their evolution in time was monitored using optical and atomic force microscopy to extract a critical radius. The critical radius scales linearly with film thickness for a homopolymer film. The results agree with a simple model which takes into account the energy cost due to surface area at the edge of the pore. The energy cost at the edge of the pore is experimentally varied by using a lamellar-forming diblock copolymer membrane. The underlying molecular architecture causes increased frustration at the pore edge resulting in an enhanced cost of pore formation.