Theory and algorithms to compute Helfrich bending forces: a review.

Cell membranes are vital to shield a cell's interior from the environment. At the same time they determine to a large extent the cell's mechanical resistance to external forces. In recent years there has been considerable interest in the accurate computational modeling of such membranes, driven mainly by the amazing variety of shapes that red blood cells and model systems such as vesicles can assume in external flows. Given that the typical height of a membrane is only a few nanometers while the surface of the cell extends over many micrometers, physical modeling approaches mostly consider the interface as a two-dimensional elastic continuum. Here we review recent modeling efforts focusing on one of the computationally most intricate components, namely the membrane's bending resistance. We start with a short background on the most widely used bending model due to Helfrich. While the Helfrich bending energy by itself is an extremely simple model equation, the computation of the resulting forces is far from trivial. At the heart of these difficulties lies the fact that the forces involve second order derivatives of the local surface curvature which by itself is the second derivative of the membrane geometry. We systematically derive and compare the different routes to obtain bending forces from the Helfrich energy, namely the variational approach and the thin-shell theory. While both routes lead to mathematically identical expressions, so-called linear bending models are shown to reproduce only the leading order term while higher orders differ. The main part of the review contains a description of various computational strategies which we classify into three categories: the force, the strong and the weak formulation. We finally give some examples for the application of these strategies in actual simulations.