Using fluorescent, anionic dyes such as carboxyfluorescein
as model solutes, it is shown that the forces allowing such solutes
to be retained within sealed lipid vesicles, against a large concentration
gradient, can be primarily electrostatic in nature. At temperatures
distant from that of the ordered-fluid lipid phase transition
a small number of the anionic dye molecules trapped within lipid
vesicles are capable of traversing the lipid bilayer and establishing
an electrical diffusion potential across the membrane. Further
solute movement can then only occur with the concomitant permeation
of ions which restore electrical balance. A significant flux of
dye can be triggered by (a) increasing the permeability of the
membrane to ions (for example by the addition of ionophores such
as gramicidin, or by allowing the lipid to approach a phase transition)
or by (b) adding lipophilic counterions such as tetraphenylborate
or dinitrophenol to the system.