On the Origin of the Bioelectrical Potential Generated by the Freshwater Clam Mantle

Abstract
Mantles from freshwater clams develop potential differences (PD''s) between the 2 surfaces when they are bathed in vitro with artificial saline solutions. The magnitude and polarity of the PD is dependent on [Ca2+] in the solution bathing the mantle''s shell surface. When the solutions are gassed with 5% CO2 in oxygen, the PD is in the range 25 to 50 mv, shell side positive. It decreases of [Ca2+] in the shell solution is elevated. The concentration dependence is logarithmic with a slope of about -27 mv per 10-fold change in [Ca2+], slightly less than predicted by the Nernst equation for a membrane acting as a Ca electrode. Analysis of the electrical behavior both in intact mantles and in isolated epithelia indicates that most of the PD develops across the external membranes of epithelial cells on the shell side. There is no evidence that an active Ca transport system is involved in electro-genesis, and a model based on Ca diffusion across a selectively permeable membrane can explain existent data. If CO2 is absent, the mantle PD is very small (2-10 mv), but still sensitive to change in external [Ca2+]. it is proposed that CO2 alters intracellular pH, thereby changing the equilibrium between a large store of nonionized calcium and [Ca2+] in the cells. A role for carbonic anhydrase in the CO2 effect is suggested by the action of a specific inhibitor of this enzyme. The diffusion model predicts that increasing ionized Ca should increase the PD as is actually observed. Some implications of this system for the physiology of Ca movement in vivo are discussed.