Shape Fluctuations of a Droplet Containing a Polymer

We consider the problem of an ideal polymer confined in a droplet. When the droplet radius is smaller than the (unconfined) polymer radius of gyration, the polymer entropy will depend on the droplet shape. We compute the resulting surface free energy. Using parameters appropriate for polymers confined in microemulsions, we find that the polymer and bending surface energies are comparable for the lowest modes. Finally, we argue that chain self-avoidance will decrease the strength of the polymer contribution to the surface energy.Comment: Revtex, 12 pages, one figur

Fluctuation-Induced Interactions between Rods on Membranes and Interfaces

We consider the interaction between two rods embedded in a fluctuating surface which is governed by either surface tension or rigidity. The modification of fluctuations by the rods leads to an attractive long-range interaction that falls off as $1/R^4$ with their separation. The orientational dependence of the resulting interaction is non-trivial and may lead to interesting patterns of rod-like objects on such surfaces.Comment: Revtex, 10 pages, one figur

Budding and vesiculation induced by conical membrane inclusions

Conical inclusions in a lipid bilayer generate an overall spontaneous curvature of the membrane that depends on concentration and geometry of the inclusions. Examples are integral and attached membrane proteins, viruses, and lipid domains. We propose an analytical model to study budding and vesiculation of the lipid bilayer membrane, which is based on the membrane bending energy and the translational entropy of the inclusions. If the inclusions are placed on a membrane with similar curvature radius, their repulsive membrane-mediated interaction is screened. Therefore, for high inclusion density the inclusions aggregate, induce bud formation, and finally vesiculation. Already with the bending energy alone our model allows the prediction of bud radii. However, in case the inclusions induce a single large vesicle to split into two smaller vesicles, bending energy alone predicts that the smaller vesicles have different sizes whereas the translational entropy favors the formation of equal-sized vesicles. Our results agree well with those of recent computer simulations.Comment: 11 pages, 12 figure

Lateral diffusion of a protein on a fluctuating membrane

Measurements of lateral diffusion of proteins in a membrane typically assume that the movement of the protein occurs in a flat plane. Real membranes, however, are subject to thermal fluctuations, leading to movement of an inclusion into the third dimension. We calculate the magnitude of this effect by projecting real three-dimensional diffusion onto an effective one on a flat plane. We consider both a protein that is free to diffuse in the membrane and one that also couples to the local curvature. For a freely diffusing inclusion the measured projected diffusion constant is up to 15% smaller than the actual value. Coupling to the curvature enhances diffusion significantly up to a factor of two.Comment: 6 pages, 4 figure

Interactions between proteins bound to biomembranes

We study a physical model for the interaction between general inclusions bound to fluid membranes that possess finite tension, as well as the usual bending rigidity. We are motivated by an interest in proteins bound to cell membranes that apply forces to these membranes, due to either entropic or direct chemical interactions. We find an exact analytic solution for the repulsive interaction between two similar circularly symmetric inclusions. This repulsion extends over length scales of order tens of nanometers, and contrasts with the membrane-mediated contact attraction for similar inclusions on tensionless membranes. For non circularly symmetric inclusions we study the small, algebraically long-ranged, attractive contribution to the force that arises. We discuss the relevance of our results to biological phenomena, such as the budding of caveolae from cell membranes and the striations that are observed on their coats.Comment: 22 pages, 2 figure

Fluctuation-Induced Interactions between Rods on a Membrane

We consider the interaction between two rods embedded in a fluctuating surface. The modification of fluctuations by the rods leads to an attractive long-range interaction between them. We consider fluctuations governed by either surface tension (films) or bending rigidity (membranes). In both cases the interaction falls off with the separation of the rods as $1/R^4$. The orientational part of the interaction is proportional to $\cos^2\left[ \theta_1+\theta_2 \right]$ in the former case, and to $\cos^2\left[ 2\left(\theta_1+\theta_2\right) \right]$ in the latter, where $\theta_1$ and $\theta_2$ are angles between the rods and the line joining them. These interactions are somewhat reminiscent of dipolar forces and will tend to align collections of such rods into chains.Comment: REVTEX, 14 pages, with 2 Postscript figure

A New Phase of Tethered Membranes: Tubules

We show that fluctuating tethered membranes with {\it any} intrinsic anisotropy unavoidably exhibit a new phase between the previously predicted ``flat'' and ``crumpled'' phases, in high spatial dimensions $d$ where the crumpled phase exists. In this new "tubule" phase, the membrane is crumpled in one direction but extended nearly straight in the other. Its average thickness is $R_G\sim L^{\nu_t}$ with $L$ the intrinsic size of the membrane. This phase is more likely to persist down to $d=3$ than the crumpled phase. In Flory theory, the universal exponent $\nu_t=3/4$, which we conjecture is an exact result. We study the elasticity and fluctuations of the tubule state, and the transitions into it.Comment: 4 pages, self-unpacking uuencoded compressed postscript file with figures already inside text; unpacking instructions are at the top of file. To appear in Phys. Rev. Lett. November (1995

Segregation of receptor-ligand complexes in cell adhesion zones: Phase diagrams and role of thermal membrane roughness

The adhesion zone of immune cells, the 'immunological synapse', exhibits characteristic domains of receptor-ligand complexes. The domain formation is likely caused by a length difference of the receptor-ligand complexes, and has been investigated in experiments in which T cells adhere to supported membranes with anchored ligands. For supported membranes with two types of anchored ligands, MHCp and ICAM1, that bind to the receptors TCR and LFA1 in the cell membrane, the coexistence of domains of TCR-MHCp and LFA1-ICAM1 complexes in the cell adhesion zone has been observed for a wide range of ligand concentrations and affinities. For supported membranes with long and short ligands that bind to the same cell receptor CD2, in contrast, domain coexistence has been observed for a rather narrow ratio of ligand concentrations. In this article, we determine detailed phase diagrams for cells adhering to supported membranes with a statistical-physical model of cell adhesion. We find a characteristic difference between the adhesion scenarios in which two types of ligands in a supported membrane bind (i) to the same cell receptor or (ii) to two different cell receptors, which helps to explain the experimental observations. Our phase diagrams fully include thermal shape fluctuations of the cell membranes on nanometer scales, which lead to a critical point for the domain formation and to a cooperative binding of the receptors and ligands.Comment: 23 pages, 6 figure