A thermodynamic theory for the membrane electroporation of curved membranes
such as those of lipid vesicles and cylindrical membrane tubes has been de
veloped. The theory covers in particular the observation that electric pore
formation and shape deformation of vesicles and cells are dependent on the
salt concentration of the suspending solvent. It is shown that transmembra
ne salt gradients can appreciably modify the electrostatic part of Helfrich
's spontaneous curvature, elastic bending rigidity and Gaussian curvature m
odulus of charged membranes. The Gibbs reaction energy of membrane electrop
oration can be explicitely expressed in terms of salt gradient-dependent co
ntributions of bending, the ionic double layers and electric surface potent
ials and dielectric polarisation of aqueous pores. In order to cover the va
rious physical contribution to the chemical process of electroporation-rese
aling, we have introduced a generalised chemophysical potential covering al
l generalised forces and generalised displacements in terms of a transforme
d Gibbs energy formalism. Comparison with, and analysis of, the data of ele
ctrooptical relaxation kinetic studies show that the Gibbs reaction energy
terms can be directly determined from turbidity dichroism (Planck's conserv
ative dichroism). The approach also quantifies the electroporative cross-me
mbrane material exchange such as electrolyte release, electrohaemolysis of
red blood cells or uptake of drugs and dyes and finally gene DNA by membran
e electroporation. (C) 2000 Elsevier Science B.V. All rights reserved.