Evaluation of kinetic and energetic parameters of active sodium transport

Active sodium transport is classically analyzed in terms of an equivalent circuit, comprising an active conductance Ka and an electromotive force of sodium transport ENa. Although ENa is commonly considered the driving force of transport, model experiments have suggested that ENa is a composite parameter, incorporating both kinetic and energetic factors. An alternative approach considers both transport and the associated oxidative metabolism in terms of a nonequilibrium thermodynamic (NET) formulation, involving phenomenological coefficients and the affinity A, presumed to represent kinetic and energetic factors, respectively. Model experiments testing the NET formulation suggest that the affinity is indeed an energetic parameter. Calculated values of A in untreated frog skins and toad bladders range from about 20 to 80 kcal per mole of O2 consumption. Assuming a P/O ratio of 3, this range corresponds to about 3--13 kcal per mole of ATP utilization, values compatible with reported direct measurements. Although brief perturbations of transepithelial electrical potential deltapsi resulted in linear current-voltage relationship, indicating constancy of ENa and Ka, 15-min perturbations of deltapsi resulted in nonlinearity, indicating changes in ENa and Ka; perturbations of deltapsi enhancing active transport were associated with decrease of ENa and increase of Ka; slowing of active transport produced the converse effects.