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We present a minimal model of plasma membrane heterogeneity that combinescriticality with connectivity to cortical cytoskeleton. Our model is motivatedby recent observations of micron-sized critical fluctuations in plasma membranevesicles that are detached from their cortical cytoskeleton. We incorporatecriticality using a conserved order parameter Ising model coupled to a simpleactin cytoskeleton interacting through point-like pinning sites. Using thisminimal model, we recapitulate several experimental observations of plasmamembrane raft heterogeneity. Small (r~20nm) and dynamic fluctuations atphysiological temperatures arise from criticality. Including connectivity tocortical cytoskeleton disrupts large fluctuations, prevents macroscopic phaseseparation at low temperatures (T<=22{\deg}C), and provides a template for longlived fluctuations at physiological temperature (T=37{\deg}C).Cytoskeleton-stabilized fluctuations produce significant barriers to thediffusion of some membrane components in a manner that is weakly dependent onthe number of pinning sites and strongly dependent on criticality. Moregenerally, we demonstrate that critical fluctuations provide a physicalmechanism to organize and spatially segregate membrane components by providingchannels for interaction over large distances.

A minimal model of plasma membrane heterogeneity requires coupling cortical actin to criticality