Elementary excitations of biomembranes: Differential geometry of undulations in elastic surfaces

Biomembrane undulations are elementary excitations in the elastic surfaces of cells and vesicles. As such they can provide surprising insights into the mechanical processes that shape and stabilize biomembranes. We explain how naturally these undulations can be described by classical differential geometry. In particular, we apply the analytical formalism of differential-geometric calculus to the surfaces generated by a cell membrane and underlying cytoskeleton. After a short derivation of the energy due to a membraneʹs elasticity, we show how undulations arise as elementary excitations originating from the second derivative of an energy functional. Furthermore, we expound the efficiency of classical differential-geometric formalism to understand the effect of differential operators that characterize processes involved in membrane physics. As an introduction to concepts the paper is self-contained and rarely exceeds calculus level.