Dual skin functions in amphibian osmoregulation

Abstract

August Krogh's studies of the frog identified the respiratory function of the skin in 1904 and the osmoregulatory function of the skin in 1937. It is the thesis of my review that the osmoregulatory function of the skin has evolved for meeting quite different demands. In freshwater the body fluid homeostasis is challenged by loss of ions to the environment. This is compensated for by active ion uptake energized by the sodium-pump ATPase and the V-type proton pump ATPase. I conclude that Krogh's astonishing observation of cutaneous chloride uptake from μM concentrations of NaCl is compatible with the free energy changes of ATP hydrolysis catalyzed by the sodium‑potassium pump ATPase and the V-type proton pump ATPase operating in series, and in parallel with experimentally verified vanishingly small leak fluxes. On land the frog is challenged by evaporative water loss through the highly water permeable skin, similar to the water permeable conducting airways of terrestrial vertebrates including man. The epithelia serving respiratory gas exchanges are heterocellular and have molecular, structural and functional properties in common. The cutaneous surface liquid of amphibians evolved for protecting the skin epithelium from desiccation like the airway surface liquid of the lung. Published studies of ion transport mechanisms of acinar cells and the two types of epithelial cells, lead to the hypothesis that subepithelial gland secretion, evaporative water loss, and ion reabsorption by the epithelium regulate composition and volume of the cutaneous surface liquid.