Transients in toad skin: Short circuit current and ionic fluxes related to inner sodium substitution by monovalent cations

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Abstract
When the Na electrochemical potential difference across the skin (ΔμNa) is altered by perturbing the transmembrane electrical potential difference or the external Na concentration, effects on transport and associated oxygen consumption can be described by the formalism of linear nonequilibrium thermodynamics (Vieira, Caplan & Essig, 1972,J. Gen. Physiol. 59:77; Danisi & Lacaz-Vieira, 1974,J. Gen. Physiol. 64:372; Procópio and Lacaz-Vieira, 1977,J. Membrane Biol. 35:219). We now show that with modifications of ΔμNa by substitution of Li or choline for Na in the inner bathing solution, this formalism is no longer applicable. Inner Na by K substitution ((Na×K) i ) causes profound alterations in short-circuit current (SCC),J Na in , K efflux (J K eff ) and PD. SCC drops transiently after (Na×K) i in Cl and in SO4 media, increasing subsequently. In Cl medium, following the initial transient, there is a late decline in SCC toward a steady state. The rate of SCC decline in Cl medium is more pronounced than that observed in SO4 medium. (Na×K) i causes a transient increase inJ Na in with a peak synchronous to the minimum in SCC, both in Cl and in SO4 media. This was interpreted as due to depolarization of the inner membrane. In SO4 medium, following the peak observed after (Na×K) i J Na in drops, to increase again toward a steady state in which SCC andJ Na in are not statistically different, resembling the control condition before (Na×K) i . In Cl medium, however, theJ Na in steady state is approximately 100% higher than SCC. This difference is due to an important K efflux (J K eff ), which builds up progressively after the substitution. The apparent K permeability [J K eff /(K i )] is of comparable magnitude in Cl and in SO4 media before (Na×K) i , the apparent K permeability increases one order of magnitude as compared to the control condition before the ionic substitution. In Cl medium, the high levels ofJ Na in and ofJ K eff observed in the steady state after (Na×K) i were interpreted as being a consequence of cell swelling. SCC and PD follow very different temporal patterns after (Na×K) i which are characterized by transients in SCC and a simple fall in PD. Reasons for these differences are discussed.