Additionally, the bilayer area compressibility modulus (KA), the P2 lipid tail order parameter and lipid diffusion were calculated. KA was calculated from the amplitude of the box area fluctuations or KA = kT⟨A⟩/(N⟨(A -A0) 2 ⟩), where A is the box area, A0 is the equilibrium area, N is the number of lipids pear leaflet, and kT is the Boltzmann constant and temperature in kelvin. The P2 lipid tail order parameter was calculated from the angle between the normal of the bilayer surface and the vector along each bond in the lipid tails (θ), as P2 = ½ (3 cos 2 θ -1) and results for different tail bonds and leaflets were averaged. Lipid lateral diffusion coefficients were calculated with diffusion-GLS 195 . Diffusion-GLS uses a generalized least squares estimator to provide an optimal trade-off between potential systematic errors due to non-diffusive dynamics at short time lags, and statistical errors due to increasing uncertainties in the MSD values at longer time lags. Prior to analysis, the trajectories of the lipid centers of mass were unwrapped using an updated unwrapping scheme 196 which preserves the diffusive character of the wrapped trajectory 197 . For each property the average over the last 800 ns was reported with SE, evaluated with block averaging.