Area Compressibility Moduli of the Monolayer Leaflets of Asymmetric Bilayers from Simulations

Extraction from simulations of the area compressibility moduli of the monolayers in a bilayer is considered theoretically. A statistical mechanical derivation shows that the bilayer modulus is the sum of the two monolayer moduli, as is often supposed but contrary to a recent study. Seemingly plausible assumptions regarding fluctuations are tested rigorously. Prospects for future research are discussed.