In Electrophorus electroplaques, the agonist-induced postsynaptic conductance depends on membrane potential. During steady exposure to agonists, after a voltage step the conductance relaxes on a millisecond time scale, exponentially approaching a new equilibrium value. The relaxation rate constant k is an instantaneous function of voltage, insensitive to the past or present conductance.Two components sum to form k. A concentration-sensitive component increases linearly with agonist concentration and decreases during desensitization or exposure to curare. Thus this component reflects the average frequency at which acetylcholine receptors are opening.The voltage-sensitive component, obtained by extrapolating k to zero agonist concentration, increases at more positive potentials. For acetylcholine, the voltage-sensitive component equals the rate constant for the exponential decay of postsynaptic currents; it thus seems to be the closing rate for active receptors. The voltage-sensitive component has the relative amplitudes acetylcholine < carbamoylcholine < decamethonium, and for each agonist equals the closing rate determined from "noise" measurements at neuromuscular junctions.The kinetic data explain several aspects of the steady-state conductance induced by agonists, but shed no light on apparent cooperative effects.At a nicotinic synapse, when an agonist reaches the postsynaptic membrane, the acetylcholine receptors produce an increase in membrane conductance. In the presence of a steady agonist concentration, the receptors form an equilibrium population. At a given time, some receptors are in an "open" state (or possibly in one of two such states); other receptors are in one or more "closed" states (1, 2).It is not yet certain which molecular events (binding of agonist, conformational change in the receptor protein, etc.) govern the transitions between the active and inactive states. In order to gain more information on these rate-limiting steps, we have applied two relaxation techniques to the acetylcholine receptors of eel electroplaques. "Voltage-jump" experiments are based on the fact that, for a steady agonist concentration, the equilibrium between "open" and "closed" receptors also depends upon the potential across the postsynaptic membrane (2-4). The proportion of open receptors increases as the potential becomes more negative. Thus, when one rapidly changes the membrane potential from one level to another, the agonist-induced conductance relaxes to a new value, with a time course which reflects the kinetics of the transition is between "open" and "closed" receptors.The second experiment derives from the observation that, at the frog neuromuscular junction, endplate currents ordinarily last much longer than the acetylcholine concentration in the synaptic cleft (5-8). Thus the declining phase of the endplate current is the response of receptors to an instantaneous removal of acetylcholine. We report here that the declining phase of eel postsynaptic currents has precisely the same significance.We...