Membrane-mediated interactions between circular particles in the strongly curved regime

Particles binding to a fluid lipid membrane can induce bilayer deformations, for instance when these particles are curved. Since the energy of two overlapping warp fields depends on the mutual distance between the two particles creating them, they will experience forces mediated by the curvature of the membrane. If the deformations are sufficiently weak, the associated differential equations for the membrane shape are linear, and the resulting interactions are understood very well; but very little is known for stronger curvature imprint, owing to the highly nonlinear nature of the problem. Here we numerically calculate the magnitude of such membrane-mediated interactions in the case of two axisymmetric particles over a wide range of curvature imprints, deep into the nonlinear regime. We show that over an intermediate distance range the sign of the force reverses beyond a sufficiently strong deformation. These findings are quantitatively confirmed by a simple analytical close-distance expansion. The sign flip can be traced to a change in magnitude between the two principal curvatures midway between the two particles, which can only occur at sufficient particle tilt, a condition which is by construction ruled out in the linearized description. We also show these large perturbation results to agree with coarse-grained molecular dynamics simulations and suggest that a favorable comparison is indeed more likely to hold in the strongly deformed regime.