Distribution and medullary projection of respiratory neurons in the dorsolateral pons of the rat.

The dorsolateral pons around the parabrachial nucleus including the Kölliker-Fuse nucleus is closely linked with the medullary respiratory center and plays an important role in respiratory control. We aimed to elucidate the firing properties, detailed distributions, and medullary projections of pontine respiratory neurons in pentobarbitone-anesthetized, paralyzed, and artificially ventilated rats with intact vagi. A total of 235 respiratory neurons were recorded from the dorsolateral pons in and around the Kölliker-Fuse nucleus. Six types of firing patterns were identified: inspiratory, expiratory-inspiratory phase spanning, inspiratory-expiratory phase spanning, decrementing expiratory, augmenting expiratory, and whole-phase expiratory patterns. Of these, the inspiratory neurons and the expiratory-inspiratory phase spanning neurons, which constituted the largest population (61%), were characterized most carefully by changing lung inflation levels, since under some conditions both showed similar firing patterns. Many (58%) of the 133 respiratory neurons examined were antidromically activated by electrical stimulation of the medulla. They were activated from the ventrolateral medulla around the ventral respiratory group and the Bötzinger complex and from the dorsomedial medulla around the nucleus tractus solitarii and the hypoglossal nucleus. The projections to the dorsomedial medulla were bilateral in many cases, and those to the ventrolateral medulla were unilateral. Of these medullary projections, two specific projections could be characterized in detail. First, many expiratory-inspiratory phase spanning neurons projected to the hypoglossal nucleus, suggesting that these pontine neurons are important premotor neurons of the hypoglossal motoneurons. This projection explains well the hypoglossal inspiratory activity, which is often dissociated from the phrenic inspiratory activity. Second, most whole-phase expiratory neurons that were distributed medially to the KF nucleus sent their axons toward the spinal cord via the midline medulla. These findings provide a new insight into the pontine control of medullary and spinal respiratory function.

GABA, in some cases together with glycine, is used as the inhibitory transmitter by pump cells in the Hering-Breuer reflex pathway of the rat.

The Hering-Breuer reflex is one of the fundamental respiratory reflexes and is mediated by second-order relay neurons of the slowly adapting lung stretch receptors. These neurons, which are called pump cells, are located in the nucleus tractus solitarii and include a population of inhibitory neurons. We aimed to determine which transmitter, GABA or glycine, the inhibitory pump cells use. In addition, we examined whether or not second-order relay neurons of the rapidly-adapting lung stretch receptors (RAR-cells), whose excitatory or inhibitory nature is not known, use these inhibitory neurotransmitters. In Nembutal-anesthetized, neuromuscularly blocked and artificially ventilated rats, we labeled pump cells (n=33) and RAR-cells (n=26) with Neurobiotin and processed the tissues for detection of mRNA encoding either glutamic acid decarboxylase isoform 67 (GAD67) or glycine transporter 2 (GLYT2) using in situ hybridization. The pump cells were located in the interstitial nucleus and its vicinity and the RAR-cells in the commissural subnucleus. The majority (64%) of the pump cells examined for GAD67 mRNA and many (26%) of the pump cells examined for GLYT2 mRNA expressed respective mRNAs. Of the eight pump cells in which both mRNAs were double-detected, three expressed both mRNAs and one expressed GAD67 mRNA but not GLYT2 mRNA, the other four expressing neither mRNAs. On the other hand, RAR-cells expressed neither GAD67 mRNA nor GLYT2 mRNA. The results suggest that the inhibitory pump cells are basically GABAergic and some of them may corelease GABA and glycine, and that RAR-cells are neither GABAergic nor glycinergic. These findings expand our understanding of the networks of lung receptor-mediated reflexes including the Hering-Breuer reflex.

Morphology of pulmonary rapidly adapting receptor relay neurons in the rat.

The term rapidly adapting pulmonary stretch receptor (RAR) refers to one of the major pulmonary sensory receptors that responds to inflation and deflation of the lungs as well as to irritant stimuli with rapidly adapting irregular discharges. The functional role and central pathways are largely unknown. The aim of this study was to elucidate morphological characteristics of second-order neurons (RAR cells) activated by vagal afferent fibers originating from RARs. A mixture of horseradish peroxidase (HRP) and Neurobiotin was injected intracellularly into physiologically identified RAR cells in Nembutal-anesthetized, immobilized, and artificially ventilated Wister rats. Direct visualization of individual RAR cells (n = 12), including their somata, dendritic arborizations, and fine axonal branches with terminal boutons, was possible for the first time. Their somata were located in the commissural or medial subdivision of the nucleus of the solitary tract, caudal to the level of the area postrema. The RAR cells had, in addition to dendrites extending into the NTS area, one or two long dendrites extending laterally and/or ventrolaterally into the medullary reticular formation. The stem axons issuing from the RAR cells first coursed ventrolaterally toward the reticular formation in the vicinity of the ambiguus nucleus and then bifurcated into ascending and descending axons: three RAR cells possessed only ascending axons. Some of the ascending axons could be traced as far as the level of the facial nucleus and some of the descending axons beyond the spinomedullary junction. These ascending and/or descending axons gave off extensive axon collaterals distributing boutons within and in the vicinity of the ambiguus nucleus. These results, showing an anatomical substrate for the network implicated in RAR-evoked reflexes, provide useful clues for study of the RAR system.

Identification of deflation-sensitive inspiratory neurons in the dorsal respiratory group of the rat.

It has been well established that inspiratory neurons of the dorsal respiratory group (DRG) are classified into two types based on whether they receive inputs from slowly-adapting pulmonary stretch receptors (SARs) or not. Inspiratory neurons with SAR inputs are called Ibeta and the others are called Ialpha neurons. In this study, we identified a novel group of inspiratory neurons in Nembutal-anesthetized, paralyzed, and artificially-ventilated rats. (1) These DRG inspiratory neurons were activated characteristically by lung deflation. (2) They were orthodromically activated by electrical stimulation of the vagus nerve at a low intensity just above the threshold for afferents from SARs or rapidly-adapting pulmonary stretch receptors (RARs). (3) The orthodromic latencies (ranged from 1.9 to 2.5 ms) indicated that they receive direct inputs from low-threshold vagal afferents. (4) Unlike Ibeta neurons, they hardly responded to lung inflation and never exhibited tonic firing in response to maintained lung inflation. (5) The majority (92%) of them were antidromically activated by electrical stimulation of the cervical spinal cord. These deflation-sensitive inspiratory neurons clearly form a distinct group, and their firing pattern is consistent with the hypothesis that they receive inputs from RAR afferents as well as the central inspiratory drive. The results indicating that DRG inspiratory neurons are classified into at least three groups provide new insights into the organization and role of the DRG.

Lung inflation inhibits rapidly adapting receptor relay neurons in the rat.

Pulmonary slowly adapting receptors (SARs) and rapidly adapting receptors (RARs) are important components of various respiratory reflexes. In anesthetized, paralyzed and artificially ventilated rats, we found an inhibitory linkage from the former to the latter system at the level of their second-order relay neurons (P cells and RAR cells, respectively). Lung inflation which activates RARs as well as SARs suppressed RAR cell activity evoked by electrical stimulation of the vagus nerve. Intracellular recordings from RAR cells showed IPSPs locked to electrical stimulation of the ipsilateral and contralateral vagus nerves at an intensity just above the threshold for P cell activation. Activation of P cells with glutamate suppressed RAR cell firing. Since P cells project to the area of RAR cells, taken together, these results strongly suggest that P cells synaptically inhibit RAR cells.

Excitatory and inhibitory synaptic inputs shape the discharge pattern of pump neurons of the nucleus tractus solitarii in the rat.

The second-order relay neurons of the slowly-adapting pulmonary stretch receptors (SARs) are called pump neurons (P cells) and are located in the nucleus tractus solitarii (NTS). We have shown recently that P cells do not act merely as simple relay neurons of SAR afferents but also receive rhythmic inputs from the central respiratory system. This study aimed to analyze two aspects of the respiratory inputs to P cells: (1) suppression of P cell firing at early inspiration (eI suppression) and (2) facilitation of P cell firing at around the period from late inspiration to early expiration (IE facilitation). This study employed extracellular recordings combined with iontophoretic applications of neuroactive drugs to single P cells, in Nembutal-anesthetized, paralyzed, and artificially ventilated rats. The results showed that several excitatory and inhibitory neurotransmitters were involved in these synaptic events. First, the glycine antagonist strychnine and the GABA(A) antagonist bicuculline were applied to identify the neurotransmitters acting in eI suppression. Strychnine greatly diminished eI suppression, but bicuculline had little effect. This suggested that eI suppression was elicited by inspiratory neurons that were glycinergic and had a decrementing firing pattern. Second, on the other hand bicuculline markedly enhanced IE facilitation as well as the baseline frequency of P cell firing. The enhancement of IE facilitation was distinctive even when the effects of increased baseline firing on this enhancement were taken into account. Third, IE facilitation was diminished by applications of the NMDA glutamate receptor antagonists D-2-amino-5-phosphonovaleric acid (APV) and dizocilpine (MK-801). These results suggested that glutamatergic synapses on P cells from some unidentified respiratory neurons form excitatory inputs for IE facilitation and GABA(A) receptor-mediated processes control the strength of IE facilitation, possibly at the presynaptic level. Finally, iontophoretic application of the non-NMDA glutamate receptor antagonist, 6-cyano-7-nitroquinoxaline-2, 3-dione disodium (CNQX), almost completely abolished P cell firing in response to both lung inflation and electrical stimulation of the vagus nerve. This confirmed the previous report that glutamate is the primary neurotransmitter at the synapses between SAR afferents and P cells. We concluded that complicated synaptic inputs involving glycinergic and GABAergic inhibitions, and non-NMDA and NMDA glutamate receptor-mediated excitations form the basic pattern of P cell firing.

Electrophysiological and pharmacological analysis of synaptic inputs to pulmonary rapidly adapting receptor relay neurons in the rat.

The information from pulmonary rapidly adapting stretch receptors (RARs) to the central nervous system (CNS) is relayed in the nucleus tractus solitarii (NTS). The second-order neurons in the NTS referred to as RAR cells have recently been shown to receive rhythmic inputs from the central respiratory system in addition to the main inputs from RAR afferents. The present study analyzed these synaptic inputs by intracellular recordings from RAR cells, and by extracellular recordings combined with local applications of neuroactive drugs to RAR cells, in Nembutal-anesthetized, paralyzed, and artificially ventilated rats. The intracellular analysis identified both excitatory postsynaptic potentials (EPSPs) elicited presumably by RAR afferents and inhibitory postsynaptic potentials (IPSPs) synchronous with central inspiratory activity. This inhibitory input, called I suppression, was the origin of respiratory modulation of RAR cell firing, and its time course suggested that some unidentified inspiratory neurons with an augmenting firing pattern were the source of the inhibition. The pharmacological analysis suggested the types of neurotransmitters used in these synaptic events. First, glutamate was shown to be the primary neurotransmitter at the synapse between RAR afferents and RAR cells. Iontophoretic applications of the non-NMDA glutamate antagonist, CNQX, abolished RAR cell firing almost completely in response to lung inflation and deflation and to electrical stimulation of the vagus nerve. Second, glycinergic inputs which inhibited RAR cells in the inspiratory phase were revealed by applications of the glycine antagonist, strychnine. That is, the I suppression was greatly diminished by applications of strychnine. Third, although applications of the GABA(A) receptor antagonist, bicuculline, had little effect on I suppression, bicuculline markedly increased the baseline firing of RAR cells. These results imply that the information path from RARs to the CNS is regulated at the level of RAR cells by phasically-acting glycinergic inhibition in the inspiratory phase and tonically-acting GABAergic inhibition; the results also provide new insights into the neuronal mechanisms of RAR-induced reflexes.

Activity of rat pump neurons is modulated with central respiratory rhythm.

The firing properties of the second-order neurons of the slowly-adapting pulmonary stretch receptors called pump neurons (P-cells), were studied in Nembutal-anesthetized, paralyzed, and artificially-ventilated rats. Extracellular recording was made from single P-cells which were monosynaptically activated by electrical stimulation of the vagal afferents and fired in phase with lung inflations. In the majority of the P-cells examined (49/52), the firing was suppressed to various extents in the inspiratory phase: in effect, their firing was accentuated at the time of the inspiratory off-switch. This issue has never been reported in cat P-cells, on which relatively rich data have been accumulated. The present results suggest that rat P-cells do not only relay information from the lung stretch receptors but also integrate the central respiratory inputs.

Pontine projections of pulmonary slowly adapting receptor relay neurons in the cat.

Pontine projections of second-order neurons activated by vagal afferents originating from pulmonary slowly adapting receptors were studied electrophysiologically in Nembutal-anesthetized, paralyzed and artificially ventilated cats. Extracellular recordings from these neurons (referred to as P-cells) were made in the nucleus tractus solitarii. Antidromic mapping of the brain stem showed that many P-cells examined projected their axons to the ipsilateral pons as well as the medulla. Axonal arborizations were found in the parabrachial nucleus, i.e. in the area corresponding to the pontine respiratory group. In some P-cells, axonal arborizations were also found in the A5 area. The present results suggest that the P-cells are involved in pontine pneumotaxic mechanisms.

Inspiratory inhibition of pulmonary rapidly adapting receptor relay neurons in the rat.

Second-order relay neurons (referred to as rapidly adapting pulmonary stretch receptors (RARs)-cells) activated by vagal afferents originating from RARs were identified electrophysiologically in Nembutal-anesthetized, paralyzed and artificially ventilated rats. Their location in the commissural subnucleus of the nucleus tractus solitarii (NTS) and firing properties in response to lung inflations and deflations were studied in detail for the first time in the rat. Importantly, in all the RAR-cells examined, their firing was suppressed to various extent during a period from inspiration to early expiration. This means that the information from the RARs to the central nervous system is gated by the central respiratory system at the level of RAR-cells in the NTS.

Convergence of central respiratory and locomotor rhythms onto single neurons of the lateral reticular nucleus.

We have analyzed the behavior of neurons of the lateral reticular nucleus (LRN) during fictive respiration and locomotion and found that some LRN neurons have both central respiratory and locomotor. rhythms. Experiments were conducted on decerebrate, decerebellate, immobilized, and artificially ventilated cats, with the spinal cord transected at the lower thoracic cord. Fictive respiration and fictive forelimb locomotion were ascertained by monitoring activities from the phrenic nerve and forelimb extensor and flexor nerves, respectively. Fictive locomotion was evoked by electrical stimulation of the mesencephalic locomotor region (MLR) or sometimes occurred spontaneously. During fictive locomotion many LRN neurons fired in certain phases of the locomotion cycle; i.e., with respect to the nerve discharge of the ipsilateral forelimb they fired in either the extensor, flexor, extensor-flexor, or flexor-extensor phase. Firing of some LRN neurons was modulated synchronously with central respiratory rhythm. Neurons with inspiratory activity and those with expiratory activity were both found. More than half of these respiration-related LRN neurons had locomotor rhythm as well. The majority of the three types of LRN neurons, i.e., neurons with only locomotor rhythm, those with only respiratory rhythm, and those with both respiratory and locomotor rhythms, were antidromically activated by electrical stimulation of the ipsilateral inferior cerebellar peduncle. Electrical stimulation of the upper cervical cord showed that these LRN neurons, not only locomotion-related but also respiration-related neurons, received short latency inputs from the spinal cord. The LRN neurons studied were distributed widely in the LRN, relatively densely in the caudal two-thirds of the nucleus. No particular differences were detected between the three types of LRN neurons with respect to their location in the nucleus. These results indicate that the information about central respiratory and locomotor rhythms that is necessary for cerebellar control of the coordination between respiration and locomotion converges, at least partly, at the level of the LRN.

Contralateral projections of barosensitive neurons of the nucleus tractus solitarii.

Axonal projections of single barosensitive neurons of the nucleus tractus solitarii (NTS) which fired in synchrony with heartbeat, were studied in Nembutal-anesthetized, paralyzed, and artificially ventilated cats. The majority of them were orthodromically activated by electrical stimulation of the ipsilateral cervical vagal nerve. Antidromic mapping by electrical stimulation of the medulla could identify the axonal projections in 14 of the 25 barosensitive NTS neurons examined. Their stem axons crossed the midline to the contralateral side and ascended rostrally. The contralateral axons of some neurons issued collaterals in the rostral ventrolateral medulla (RVLM) medial to and ventromedial to the facial nucleus. The contralateral axons of the other neurons ascended toward the pons without any sign of issuing collaterals within the medulla.

Pump neurons of the nucleus of the solitary tract project widely to the medulla.

Axonal projections of pump neurons (P-cells) of the nucleus of the solitary tract were studied in Nembutal-anesthetized, paralyzed, and artificially ventilated cats. Extracellular recordings were made from single P-cells which were monosynaptically activated by electrical stimulation of the ipsilateral vagal afferents and distributed in close proximity to the solitary tract. Antidromic stimulation of the medulla showed that all P-cells examined projected their axons to the ipsilateral ventrolateral medulla. Axonal arborizations were found in respiration-related areas and their vicinity between the level of the retrofacial nucleus and the level a few millimeters caudal to the obex. Some of these P-cells sent axon collaterals to the contralateral nucleus of the solitary tract (NTS) as well. This study is the first to show wide spread medullary projections of P-cell axons.

Activity of bulbar respiratory neurons during fictive coughing and swallowing in the decerebrate cat.

1. The behaviour of medullary respiratory neurons was studied during fictive coughing and swallowing evoked by electrical stimulation of the superior laryngeal nerve (SLN) in decerebrate, paralysed and artificially ventilated cats. Fictive coughing, swallowing and respiration were monitored by recording activities of the phrenic, hypoglossal and abdominal nerves. 2. Extracellular recordings were made from respiratory neurons in the ventral respiratory group (VRG) and in the Bötzinger complex (BOT). The neuronal types analysed included decrementing inspiratory neurons (I-DEC), augmenting expiratory neurons (E-AUG) and decrementing expiratory neurons (E-DEC) from the BOT area, and augmenting inspiratory neurons (I-AUG) and augmenting expiratory neurons (E-AUG) from the VRG area. 3. During fictive coughing, all the inspiratory and expiratory neurons were active during the inspiratory and expiratory phases of coughing, respectively. The firing of both I-DEC and I-AUG neurons was increased and prolonged in association with the augmented inspiratory activity of the phrenic nerve. The activity of E-AUG neurons of the VRG did not parallel the abdominal nerve activity, suggesting the existence of additional neurons which participate in the generation of abdominal nerve activity during fictive coughing. 4. During fictive swallowing, half of I-DEC neurons fired transiently at the onset of hypoglossal bursts associated with swallowing; the firing was suppressed during the rest of the hypoglossal bursts. Other I-DEC neurons were silent during hypoglossal bursts. Some I-AUG neurons fired during the initial half of hypoglossal bursts, and others were silent. The brief phrenic activity accompanying the swallowing might have originated from this activity in I-AUG neurons. The discharges of all E-AUG neurons (BOT and VRG) and the majority of E-DEC BOT neurons were suppressed during swallowing. 5. We conclude that these five types of respiratory neurons of the BOT and VRG are involved in the generation of the spatiotemporally organized activity of coughing and swallowing, and that at least a part of the neuronal network for respiration is shared by networks for these non-respiratory activities.

Location and axonal projection of one type of swallowing interneurons in cat medulla.

Extracellular recordings were made from a type of relay neurons of the superior laryngeal nerve (SLN) afferents in the vicinity of the retrofacial nucleus (RFN) in either pentobarbitone-anesthetized or unanesthetized and decerebrate cats, which were paralyzed and artificially ventilated. A total of 26 neurons that could be activated both orthodromically by electrical stimulation of the SLN and antidromically by stimulation of the brainstem were analyzed. All 26 neurons were activated from the ipsilateral SLN and 13 were activated from the contralateral SLN with mean latencies of 7.7 ms and 11.4 ms, respectively. The majority of these neurons were located in the parvocellular reticular formation dorsomedial to the RFN and to the rostral part of the nucleus ambiguus (AMB). Antidromic stimulation of the medulla showed that 22 of the 26 neurons projected to the hypoglossal nucleus (HYP) and 19 neurons tested projected to the AMB. Of these, 15 neurons projected to both the HYP and AMB and two projected to the lateral reticular nucleus as well. Seventeen neurons were tested for their behavior during fictive swallowing which was elicited by continual electrical stimulation of the SLN and monitored by the activity of the hypoglossal nerve. Twelve neurons showed brief (100-200 ms) burst firing at the onset of swallowing; the firing of the other 5 neurons were suppressed during swallowing. Both the swallowing-active and swallowing-inactive neurons projected to the HYP and AMB. Thus, the SLN relay neurons in the vicinity of the RFN might participate in the early stage of SLN-induced swallowing by integrating inputs from SLN afferents.

Location and axonal projection of early-onset decrementing expiratory neurons in the cat.

The location and axonal projection of early-onset decrementing expiratory (eE-DEC) neurons were studied in Nembutal-anesthetized, paralyzed, and artificially ventilated cats. The eE-DEC neurons started firing around the peak of phrenic activity and attained maximum firing at the transition from inspiration to expiration. The eE-DEC neurons were distributed in the dorsomedial border of the dense assembly of respiratory neurons of the ventral respiratory group and the Bötzinger complex. Antidromic microstimulation showed that the eE-DEC neurons projected mainly to the ipsilateral reticular formation medial and dorsal to the nucleus ambiguus at the level of the obex but rarely to the medullary respiration-related areas. Participation of these eE-DEC neurons in the inspiratory off-switch may not be possible.

Projections from the commissural subnucleus of the solitary tract onto catecholamine cell groups of the ventrolateral medulla.

Efferent projections of the commissural subnucleus of the solitary tract (COM) to the ventrolateral medulla were studied in the cat using anterograde labeling with biocytin combined with dopamine beta-hydroxylase immunohistochemistry. COM neurons were observed to send their axons densely to the areas of distribution of respiration-related neurons in the ventrolateral medulla, e.g. ventral respiratory group, Bötzinger complex. Axon terminals from COM neurons were further found in the areas of distribution of catecholamine neurons (C1 and A1 cell groups), that were distributed in the close vicinity of the reported respiration-related areas in the ventrolateral medulla. Putative synaptic contacts of axon terminals from COM neurons with catecholamine neurons were often observed in the C1 area.

Projections from the commissural subnucleus of the nucleus of the solitary tract: an anterograde tracing study in the cat.

The commissural subnucleus (COM) of the nucleus of the solitary tract (NTS) is known to receive primary afferents from the lungs and other viscera innervated by the vagus nerve, and thus to participate in central autonomic and respiratory control. The aim of the present study was to identify the areas of terminal arborizations of COM neurons in order to examine brainstem sites which may be involved in reflex responses mediated by these neurons. The projections were studied in cats, using biocytin as an anterograde tracer. Labeled fibers and terminal boutons were visualized by horseradish-peroxidase histochemistry, 2-3 days after microinjection of the tracers into the COM 1-2 mm caudal to the obex. Labeled axons were examined in the brainstem from the rostral pons to the caudal medulla and were found bilaterally, with an ipsilateral predominance, mainly in the following regions: (1) The dorsolateral rostral pons. Terminal boutons were observed in the lateral and medial parabrachial nuclei, Kölliker-Fuse nucleus, and around the mesencephalic trigeminal tract. This area corresponds to the pontine respiratory group also known as the "pneumotaxic center." (2) The pontine area dorsolateral to the superior olivary nucleus. This region contains the A5 noradrenergic cell group; (3) Near the ventral surface, below the facial nucleus. This area overlaps with the 'retrotrapezoid nucleus.' (4) Respiration-related areas of the medulla, including the dorsal and ventral respiratory groups, and the Bötzinger complex. (5) The dorsal motor nucleus of the vagus. These results suggest that the COM is involved in reflex arcs, which have both respiratory functions and autonomic functions. The pathway to the dorsolateral pons, which has been identified in our recent electrophysiological study is likely to play a role in mediating respiratory responses from pulmonary rapidly adapting receptors. Other pathways may represent additional projections from second-order neurons receiving input from this group of lung receptors, or projections from as yet unidentified neurons that relay information from different afferents terminating in the COM.

Possible inspiratory off-switch neurones in the ventrolateral medulla of the cat.

The location and axonal projection of a type of respiratory neurones (termed bIE neurones), which show burst firing at the time of phase transition from inspiration to expiration, were studied in Nembutal-anaesthetized, paralysed and artificially ventilated cats. The bIE neurones showed maximum firing at the sharp decline of inspiratory activity. All of the bIE neurones tested projected to medullary respiration-related areas of the ventral respiratory group (VRG), the dorsal respiratory group (DRG), and the Bötzinger complex (BOT). The bIE neurones were distributed in the dorso-medial border of the main assembly of respiratory neurones of the VRG and BOT. The possibility that these bIE neurones participate in inspiratory termination is discussed.

Behavior of inhibitory and excitatory propriobulbar respiratory neurons during fictive vomiting.

The behavior of propriobulbar respiratory neurons was studied during fictive vomiting in decerebrate, paralyzed, artificially ventilated cats. Fictive vomiting was identified by a characteristic series of synchronous phrenic and abdominal nerve bursts, induced by electrical stimulation of abdominal vagal afferents and/or i.v. infusion of emetic drugs. Data were obtained from inspiratory neurons having decrementing (I-DEC) or constant (I-CON) discharge patterns and expiratory decrementing (E-DEC) neurons located in the Bötzinger complex and adjacent rostral ventral respiratory group. These neurons are known to make excitatory (I-CON) and inhibitory (I-DEC, E-DEC) connections with a variety of medullary respiratory neurons. During fictive vomiting: 8 of 14 I-DEC neurons fired in phase with synchronous bursts of phrenic and abdominal nerve discharge; the other 6 were silent. Of 12 I-CON neurons, 5 fired in phase with phrenic and abdominal bursts; 7 were silent. All (6) E-DEC neurons were either silent or fired weakly between bursts of phrenic and abdominal discharges. The possible roles of I-DEC and I-CON neurons in actively reorganizing the behavior of other respiratory neurons during fictive vomiting are discussed. In particular, the firing of many I-DEC neurons was found to be appropriate to inhibit inspiratory, and two types of expiratory, bulbospinal neurons during fictive vomiting.