Bilayer elasticity at the nanoscale: the need for new terms
Anne-Florence Bitbol, Doru Constantin, Jean-Baptiste Fournier

TL;DR
This paper emphasizes the importance of including gradient and Laplacian terms of area per lipid in continuum elastic models to accurately describe nanoscale membrane deformations, supported by reanalysis of experimental and numerical data.
Contribution
It introduces new terms involving the gradient and Laplacian of area per lipid into membrane elasticity models, highlighting their significance at the nanoscale.
Findings
The gradient term amplitude is estimated at 13-60 mN/m.
The model explains variations in numerical data.
The gradient term aligns with interfacial tension estimates.
Abstract
Continuum elastic models that account for membrane thickness variations are especially useful in the description of nanoscale deformations due to the presence of membrane proteins with hydrophobic mismatch. We show that terms involving the gradient and the Laplacian of the area per lipid are significant and must be retained in the effective Hamiltonian of the membrane. We reanalyze recent numerical data, as well as experimental data on gramicidin channels, in light of our model. This analysis yields consistent results for the term stemming from the gradient of the area per molecule. The order of magnitude we find for the associated amplitude, namely 13-60 mN/m, is in good agreement with the 25 mN/m contribution of the interfacial tension between water and the hydrophobic part of the membrane. The presence of this term explains a systematic variation in previously published numerical…
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