Hydrogen bonds were restrained for all systems using LINCS 47 allowing integration steps of 2 fs. Temperature and pressure were kept constant at 310 kelvin and 1 bar using the vrescaling thermostat 48 (τ t = 1 ps) and the Parrinello-Rahman barostat (τ p = 12 ps). The c-rescale barostat 51 (τ p = 4 ps) was used for equilibration. Semi-isotropic pressure coupling was used for the membrane systems. Compressability was set to 4.5 × 10 -5 .