The Ionic Permeability Changes during Acetylcholine-Induced Responses of Aplysia Ganglion Cells

ACh-induced depolarization (D response) in D cells markedly decreases as the external Na⁺ is reduced. However, when Na⁺ is completely replaced with Mg⁺⁺, the D response remains unchanged. When Na⁺ is replaced with Tris(hydroxymethyl)aminomethane, the D response completely disappears, except for a slight decrease in membrane resistance. ACh-induced hyperpolarization (H response) in H cells is markedly depressed as the external Cl^- is reduced. Frequently, the reversal of the H response; i.e., depolarization, is observed during perfusion with Cl^--free media. In cells which show both D and H responses superimposed, it was possible to separate these responses from each other by perfusing the cells with either Na⁺-free or Cl^--free Ringer's solution. High [K⁺]₀ often caused a marked hyperpolarization in either D or H cells. This is due to the primary effect of high [K⁺]₀ on the presynaptic inhibitory fibers. The removal of this inhibitory afferent interference by applying Nembutal readily disclosed the predicted K⁺ depolarization. In perfusates containing normal [Na⁺]₀, the effects of Ca⁺⁺ and Mg⁺⁺ on the activities of postsynaptic membrane were minimal, supporting the current theory that the effects of these ions on the synaptic transmission are mainly presynaptic. The possible mechanism of the hyperpolarization produced by simultaneous perfusion with both high [K⁺]₀ and ACh in certain H cells is explained quantitatively under the assumption that ACh induces exclusively an increase in Cl^- permeability of the H membrane.