Electromechanical effects on tether formation from lipid membranes: A theoretical analysis

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Abstract
The material properties of biomembranes can be measured by forming a tether, a thin bilayer tube that extends from the membrane surface. Recent experiments have demonstrated that the force required to maintain a tether is sensitive to the transmembrane potential. As a first approach towards understanding this phenomenon, a thermodynamic analysis of the influence of electrical fields on tether formation from an aspirated lipid vesicle is developed. The analysis considers contributions from Maxwell stresses as well as two forms of electromechanical coupling: coupling between the electric field and curvature strain (flexoelectric coupling) and between the electric field and areal strain (piezoelectric coupling). Predictions of equilibrium tether conformations are obtained numerically. For expected values of the dimensionless coupling parameters, flexoelectric coupling alters the force required to form a tether of a given length, while piezoelectric coupling and Maxwell forces do not greatly change the force versus tether length behavior. The results of this analysis indicate that tether experiments have the potential to characterize electromechanical coupling in both synthetic and cellular membranes.