title: Minimal model of plasma membrane heterogeneity requires coupling cortical actin to criticality.
abstract: We present a minimal model of plasma membrane heterogeneity that combines criticality with connectivity to cortical cytoskeleton. The development of this model was motivated by recent observations of micron-sized critical fluctuations in plasma membrane vesicles that are detached from their cortical cytoskeleton. We incorporate criticality using a conserved order parameter Ising model coupled to a simple actin cytoskeleton interacting through point-like pinning sites. Using this minimal model, we recapitulate several experimental observations of plasma membrane raft heterogeneity. Small (r ∼ 20 nm) and dynamic fluctuations at physiological temperatures arise from criticality. Including connectivity to the cortical cytoskeleton disrupts large fluctuations, prevents macroscopic phase separation at low temperatures (T ≤ 22°C), and provides a template for long-lived fluctuations at physiological temperature (T = 37°C). Cytoskeleton-stabilized fluctuations produce significant barriers to the diffusion of some membrane components in a manner that is weakly dependent on the number of pinning sites and strongly dependent on criticality. More generally, we demonstrate that critical fluctuations provide a physical mechanism for organizing and spatially segregating membrane components by providing channels for interaction over large distances.
authors.affiliation: Department of Physics, Cornell University, Ithaca, New York, USA.
authors.forename: Benjamin B
authors.initials: BB
authors.lastname: Machta
authors.forename: Stefanos
authors.initials: S
authors.lastname: Papanikolaou
authors.forename: James P
authors.initials: JP
authors.lastname: Sethna
authors.forename: Sarah L
authors.initials: SL
authors.lastname: Veatch
mesh_terms: D000199:Actins / Q000737:chemistry* / Q000378:metabolism; D002462:Cell Membrane / Q000737:chemistry*; D003599:Cytoskeleton / Q000378:metabolism; D004058:Diffusion; D008954:Models, Biological; D013696:Temperature; D013997:Time Factors
chemical_list: D000199:Actins