Divalent ions and the surface potential of charged phospholipid membranes.

Phospholipid bilayer membranes were bathed in a decimolar solution of monovalent ions, and the conductance produced by neutral carriers of these monovalent cations and anions was used to assess the electric potential at the surface of the membrane. When the bilayers were formed from a neutral lipid, phosphatidylethanolamine, the addition of alkaline earth cations produced no detectable surface potential, indicating that little or no binding occurs to the polar head group with these ions. When the bilayers were formed from a negatively charged lipid, phosphatidylserine, the addition of Sr and Ba decreased the magnitude of the surface potential as predicted by the theory of the diffuse double layer. In particular, the potential decreased 27 mv for a 10-fold increase in concentration in the millimolar-decimolar range. A 10-fold increase in the Ca or Mg concentration also produced a 27 mv decrease in potential in this region, which was again due to screening, but it was necessary to invoke some specific binding to account for the observation that these cations were effective at a lower concentration than Ba or Sr. It is suggested that the ability of the alkaline earth cations to shift the conductance-voltage curves of a nerve along the voltage axis by 20-26 mv for a 10-fold increase in concentration may be due to essentially a screening rather than a binding phenomenon.