Fluctuations and conformational stability of a membrane patch with curvature inducing inclusions

Membranes with curvature inducing inclusions display a range of cooperative phenomena, which can be linked to biomembrane function, e.g. membrane tubulation, vesiculation, softening and spontaneous tension. We investigate how these phenomena are related for a fluctuating, framed membrane through analysis of a descretized membrane model by Monte Carlo simulation techniques. The membrane model is based on a dynamically triangulated surface equipped with non-interacting, up-down symmetry breaking inclusions where only terms coupled linearly to mean-curvature are maintained. We show that the lateral configurational entropy plays a key role for the mechanical properties of the semi-flexible membrane, e.g. a pronounced softening at intermediate inclusion coverages of the membrane and generation of membrane tension. Tensionless framed membranes will remain quasi-flat up to some threshold coverage, where a shape instability occurs with formation of pearling or tubular membranes, which below full coverage is associated with segregation of inclusions between the curved and flat membrane geometries. For inclusions with preference for highly curved membranes the instability appears at dilute inclusion coverages and is accompanied by strong configurational fluctuations.