Inking in Aplysia californica. III. Two different synaptic conductance mechanisms for triggering central program for inking.

1. In the previous two papers we described the motor elements which mediate all-or-none inking behavior in Aplysia (4) and the contribution of specific membrane properties of the ink-gland motor cells to the central program for inking (5). In the present paper we show that the central program for inking (a characteristic accelerating burst of action potentials in the motor cells) can be triggered in two different ways: 1) by means of a suprathreshold stimulus delivered to a single site on the animal; and 2) by means of two subthreshold stimuli closely spaced in time, delivered either to the same or to different sites on the animal. 2. The single suprathreshold stimulus triggers the central program primarily by means of a complex increased-conductance EPSP. The two subthreshold stimuli trigger the program by means of a complex decreased-conductance EPSP which is capable of augmenting subsequent afferent synaptic input. 3. The synaptic augmentation produced by the decreased-conductance EPSP is attributable to two properties of the EPSP: 1) it increases the input resistance of the ink-gland motor cells so that the synaptic current from afferent input onto the cells produces larger, more effective EPSP's; and 2) it increases the electrotonic coupling among the motor cells, thereby producing increased positive feedback among them. Thus, the decreased-conductance EPSP provides a novel mechanism for spatial and temporal summation.