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We find that the density-threshold affinity for M4 weakens with chain rigidity, which suggests that flexible chains may help relax packing defects caused by the conical protein shape. For any site, PE headgroups have the strongest affinity of all phospholipid headgroups, but anionic lipids still yield moderately high affinities for the M4 sites as expected. We observe cooperative effects between anionic headgroups and saturated chains at the M4 site in the inner leaflet. We also analyze affinities for individual anionic headgroups. When
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