Stochastic resonance for adhesion of membranes with active stickers.

The behavior of two membranes that interact by active adhesion molecules or stickers is studied theoretically using mean-field theory and Monte Carlo simulations. The stickers are anchored in one of the membranes and undergo conformational transitions between on and off states. In their on states, the stickers can bind to ligands that are anchored in the other membrane. The transitions between the on and off states arise from the coupling of the stickers to some active, energy-releasing process, which keeps the system out of equilibrium. As one varies the transition rates of this active process, the membrane separation undergoes a stochastic resonance: this separation is maximal at intermediate rates of the sticker transitions and considerably smaller both at high and at low transition rates. This implies that the effective, fluctuation-induced repulsion between the membranes contains a rate-dependent contribution that arises from the switching of the active stickers.

Indirect interactions of membrane-adsorbed cylinders.

Biological and biomimetic membranes often contain aggregates of embedded or adsorbed macromolecules. In this paper, the indirect interactions of cylindrical objects adhering to a planar membrane are considered theoretically. The adhesion of the cylinders causes a local perturbation of the equilibrium membrane shape, which leads to membrane-mediated interactions. For a planar membrane under lateral tension, the interaction is repulsive for a pair of cylinders adhering to the same side of the membrane, and attractive for cylinders adhering at opposite membrane sides. For a membrane in an external harmonic potential, the interaction of adsorbed cylinders is always attractive and increases if forces perpendicular to the membrane act on the cylinders.

Adhesion of membranes with competing specific and generic interactions.

Biomimetic membranes in contact with a planar substrate or a second membrane are studied theoretically. The membranes contain specific adhesion molecules (stickers) which are attracted by the second surface. In the absence of stickers, the trans-interaction between the membrane and the second surface is assumed to be repulsive at short separations. It is shown that the interplay of specific attractive and generic repulsive interactions can lead to the formation of a potential barrier. This barrier induces a line tension between bound and unbound membrane segments which results in lateral phase separation during adhesion. The mechanism for adhesion-induced phase separation is rather general, as is demonstrated by considering two distinct cases involving: i) stickers with a linear attractive potential, and ii) stickers with a short-ranged square-well potential. In both cases, membrane fluctuations reduce the potential barrier and, therefore, decrease the tendency of phase separation.

Adhesion-induced phase behavior of multicomponent membranes.

Biomimetic membranes that contain several molecular components are studied theoretically. In contact with another surface, such as a solid substrate or another membrane, some of these intramembrane components are attracted by the second surface and, thus, act as local stickers. The cooperative behavior of these systems is characterized by the interplay of (i) attractive binding energies, (ii) entropic contributions arising from the shape fluctuations of the membranes, and (iii) the entropy of mixing of the stickers. A systematic study of this interplay, which starts from the corresponding partition functions, reveals that there are several distinct mechanisms for adhesion-induced phase separation within the membranes. The first of these mechanisms is effective for flexible stickers with attractive cis interactions (within the same membrane) and arises from the renormalization of these interactions by the confined membrane fluctuations. A second, purely entropic mechanism is found for rigid stickers without attractive cis interactions and arises from a fluctuation-induced line tension. Finally, a third mechanism is present if the membrane contains both stickers and repellers, i.e., nonadhesive molecules that protrude from the membrane surface. This third mechanism is based on an effective potential barrier and becomes less effective if the shape fluctuations of the membrane become more pronounced.

Unbinding transitions and phase separation of multicomponent membranes.

Multicomponent membranes in contact with another surface or wall are studied by a variety of theoretical methods and Monte Carlo simulations. The membranes contain adhesion molecules which are attracted to the wall and, thus, act as local stickers. It is shown that this system undergoes lateral phase separation leading to discontinuous unbinding transitions if the adhesion molecules are larger than the nonadhesive membrane components. This process is driven by an effective line tension which depends on the size of the stickers and arises from the interplay of shape fluctuations and sticker clusters.