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We investigated the phase separation of dioleoylphosphatidylserine (DOPS) anddipalmitoylphosphatidylcholine (DPPC) in giant unilamellar vesicles inhypotonic solution using fluorescence and confocal laser scanning microscopy.Although phase separation in charged lipid membranes is generally suppressed bythe electrostatic repulsion between the charged headgroups, osmotic stress canpromote the formation of charged lipid domains. Interestingly, we observedthree-phase coexistence even in DOPS/DPPC binary lipid mixtures. The threephases were DPPC-rich, dissociated DOPS-rich, and nondissociated DOPS-richphases. The two forms of DOPS were found to coexist owing to the ionization ofthe DOPS headgroup, such that the system could be regarded as quasi-ternary.The three formed phases with differently ionized DOPS domains were successfullyidentified experimentally by monitoring the adsorption of positively chargedparticles. In addition, coarse-grained molecular dynamics simulations confirmedthe stability of the three-phase coexistence. Attraction mediated by hydrogenbonding between protonated DOPS molecules and reduction of the electrostaticinteractions at the domain boundaries stabilized the three-phase coexistence.

Three-phase coexistence in binary charged lipid membranes in hypotonic solution