Swimming behavior in the marine mollusc Tritonia diomedea is episodic, consisting of a series of alternating dorsal and ventral flexions initiated by a brief sensory stimulus. The swim motor pattern is generated by a network formed of four groups of premotor interneurons: cerebral cell 2 (C2), dorsal swim interneurons (DSIs), and two types of ventral swim interneurons (VSI-A and VSI-B). The initiation and maintenance of swimming depends on the establishment of a long-lasting ramp depolarization in both the premotor, pattern-generating interneurons, and the motor neurons (i.e., flexion neurons). Voltage clamp was used to measure the membrane current responsible for the ramp depolarization. In all cell classes the current had two components: a tonic inward current, which decayed as the swim progressed, and phasic inward current waves, which provided the synaptic drive during each swim burst. The ramp current in the flexion neurons and in C2 was generated largely by activity within the interneuronal pattern-generating network (PGN). The ramp current could be mimicked by driving activity in the pattern-generating interneurons. In VSI-B, the tonic component of the ramp current was independent of activity within the PGN and appeared to be derived from the long-lasting effect of an extrinsic input. The phasic components of the ramp, however, were dependent on PGN activity. The phasic inward current waves were blocked when pattern generation was prevented. In addition, phasic inward currents similar to those occurring during swimming could be produced by driving the C2. The tonic component of the ramp current in a DSI was dependent both on extrinsic inputs and PGN activity. Extrinsic inputs appeared to control the first 10-15 s of the tonic current. At longer times, activity within the DSI population itself maintained the ramp current. When one DSI was driven in a quiescent preparation, all other DSIs were inhibited, yet the DSIs are known to be coupled by monosynaptic, reciprocal excitatory synapses. This effect could be explained by the action of an unidentified inhibitory interneuron (I-neuron), which was excited by DSIs and in turn inhibited all other DSIs. The DSIs were therefore coupled reciprocally by both monosynaptic excitation and polysynaptic inhibition. Activity in C2 switched the DSI-DSI interaction from inhibition to excitation by inhibiting the I-neuron.(ABSTRACT TRUNCATED AT 400 WORDS)

The nucleus tractus solitarius (NTS) contains neurons that are part of the central neuronal network controlling rhythmic breathing movements in mammals. Nerve terminals within the NTS show immunoreactivity to thyrotropin-releasing hormone (TRH), a neuropeptide that has potent stimulatory effects on respiration. By means of a brainstem slice preparation in vitro, TRH induced rhythmic bursting in neurons in the respiratory division of the NTS. The frequency of bursting was voltage-dependent and could be reset by short depolarizing current pulses. In the presence of tetrodotoxin, TRH produced rhythmic oscillations in membrane potential whose frequency was also voltage-dependent. These observations suggest that TRH modulates the membrane excitability of NTS neurons and allows them to express endogenous bursting activity.

1. An in vitro brain stem slice preparation from adult guinea pigs was used to determine the properties of neurons located in the ventral part of the nucleus tractus solitarius (NTS), an area associated with the dorsal respiratory group. Based upon their morphology and their repetitive firing properties, three classes of ventral NTS neurons, termed types I, II, and III, were observed. 2. Type I neurons were multipolar with pyramidal-shaped cell bodies. These neurons responded to prolonged depolarizations from a resting level of -50 mV with a discrete, high-frequency burst of spikes, which rapidly adapted to a low steady-state level. When depolarized from levels more negative than -65 mV, the initial burst was diminished. 3. Type II neurons were multipolar with fusiform-shaped cell bodies. Type II neurons responded to depolarizations from -50 mV with an initial high spike frequency, which gradually adapted to a steady-state level. When depolarized from levels more negative than -60 mV, these neurons displayed a delay between the onset of the stimulus and the first spike. This delay has been termed "delayed excitation." The expression of delayed excitation was modulated by both the size and duration of hyperpolarizing prepulses that preceded depolarization. 4. Type III neurons were multipolar with spherical shaped-cell bodies. In response to depolarizations from -50 mV, these neurons displayed high-frequency firing with little adaptation. The repetitive firing properties of type III neurons were not modulated by hyperpolarization. 5. Bulbospinal neurons in the ventral NTS were identified using retrograde transport of rhodamine-labeled latex beads injected into the region of the phrenic motor nucleus at spinal cord levels C4 through C6. Only type I and type II neurons were labeled in the ventral NTS (0.2-1.0 mm rostral to the obex). Both contralateral and ipsilateral projections were observed. Contralaterally, type I and II neurons were evenly distributed. Ipsilaterally, however, type II neurons accounted for two-thirds of the labeled neurons. 6. Type I and II neurons had similar input resistances and time constants: 97.0 +/- 17.6 M omega and 14.4 +/- 2.2 ms (n = 5) for type I and 107.0 +/- 11.2 M omega and 13.7 +/- 1.6 ms for type II (n = 5).(ABSTRACT TRUNCATED AT 400 WORDS)

1. The ventral part of the nucleus tractus solitarius in guinea pigs comprises the dorsal respiratory group and is composed of three classes of neurons. These have been termed types I, II, and III. Each cell type possesses a unique set of repetitive firing properties. An in vitro brain stem slice preparation was used to study the ionic basis for these repetitive firing properties. 2. Three different membrane currents were shown to contribute to the repetitive firing properties. These were: a slow calcium current (ICa), an early, transient potassium current (IKA), and a calcium-activated potassium current (IKC). Type I and II neurons displayed physiologically significant amounts of these currents; type III neurons did not. 3. During depolarization from potential levels between -50 and -60 mV, the repetitive firing properties of type I and II neurons were determined primarily by ICa and IKC. IKA was inactivated in this potential range. The expression of IKC was greater in type I neurons than in type II neurons, and as a result, type I neurons exhibited a self-terminating burst of spike activity early in depolarization, whereas type II neurons displayed a gradual decline in spike frequency throughout depolarization. 4. The properties of IKA in type I and II neurons were studied using the single-electrode voltage-clamp technique. The kinetics of IKA in type I neurons was approximately twice as slow as those of type II neurons. In addition, the voltage dependence of activation and the removal of inactivation for IKA in type I neurons were shifted by about -10 mV with respect to type II neurons. 5. Depolarization of type I neurons from membrane potential levels where inactivation of IKA was removed caused a decrease in the frequency of the initial burst of spikes. This decrease in spike frequency was result of the coactivation of IKA with ICa. 6. Depolarization of type II neurons from membrane potentials where inactivation of IKA was removed caused a long delay between the onset of depolarization and the beginning of spike activity. The delay in excitation was modulated by both the magnitude and duration of the prestimulus hyperpolarization. This modulation of delayed excitation paralleled the time and voltage dependence for the removal of IKA inactivation in type II neurons.

SUMMARY1. Two different methods are described which allow the reversal potential (Er) for the channels opened by L-glutamate at the voltage-clamped, crayfish neuromuscular junction to be measured accurately. In both cases the value of Er was found to be about + 6 mV.2. Reversal potentials were also measured in solutions where Na+ was replaced by K+, Ca2+, or Mg2+; or in which Cl-was replaced by isethionate.3. In solutions where Na+ was partially replaced by K+, the measured reversal potentials were compared to theoretical values predicted by both the constant-field and equivalent-circuit equations. The experimental values were more accurately described by the constant-field equation.4. Permeability ratios (PX/PNa) for K+, Ca2+, Mg2+, and Cl-were calculated using the constant-field equation. K+ and Na+ were equally permeant while Ca2+ and Mg2+ were about half as permeant as the monovalent cations. Cl-was impermeant.5. The results of these experiments indicate that the L-glutamate activated channel is non-selective for cations. Furthermore, the value of the permeability ratios for the physiological cations tested are very similar to those obtained for the acetylcholine activated channel in vertebrate skeletal muscle.