Clustering on Membranes: Fluctuations and More

Clustering of extracellular ligands and proteins on the plasma membrane is required to perform specific cellular functions, such as signaling and endocy- tosis. Attractive forces that originate in perturbations of the membrane’s phys- ical properties contribute to this clustering, in addition to direct protein–protein interactions. However, these membrane-mediated forces have not all been equally considered, despite their importance. In this review, we describe how line tension, lipid depletion, and membrane curvature contribute to mem- brane-mediated clustering. Additional attractive forces that arise from protein- induced perturbation of a membrane’s fluctuations are also described. This review aims to provide a survey of the current understanding of membrane- mediated clustering and how this supports precise biological functions.