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We investigated the effect of charge on the membrane morphology of giantunilamellar vesicles (GUVs) composed of various mixtures containing chargedlipids. We observed the membrane morphologies by fluorescent and confocal lasermicroscopy in lipid mixtures consisting of a neutral unsaturated lipid[dioleoylphosphatidylcholine (DOPC)], a neutral saturated lipid[dipalmitoylphosphatidylcholine (DPPC)], a charged unsaturated lipid[dioleoylphosphatidylglycerol (DOPG$^{\scriptsize{(-)}}$)], a charged saturatedlipid [dipalmitoylphosphatidylglycerol (DPPG$^{\scriptsize{(-)}}$)], andcholesterol (Chol). In binary mixtures of neutral DOPC/DPPC and chargedDOPC/DPPG$^{\scriptsize{(-)}}$, spherical vesicles were formed. On the otherhand, pore formation was often observed with GUVs consisting ofDOPG$^{\scriptsize{(-)}}$ and DPPC. In a DPPC/DPPG$^{\scriptsize{(-)}}$/Cholternary mixture, pore-formed vesicles were also frequently observed. Thepercentage of pore-formed vesicles increased with the DPPG$^{\scriptsize{(-)}}$concentration. Moreover, when the head group charges of charged lipids werescreened by the addition of salt, pore-formed vesicles were suppressed in boththe binary and ternary charged lipid mixtures. We discuss the mechanisms ofpore formation in charged lipid mixtures and the relationship between phaseseparation and the membrane morphology. Finally, we reproduce the results seenin experimental systems by using coarse-grained molecular dynamics simulations.

Coupling between pore formation and phase separation in charged lipid membranes