The motor nuclei of the glossopharyngeal-vagal and the accessorius nerves in the rat

Retrograde cobalt labeling was performed by incubating the rootlets of cranial nerves IX, X and XI, or the central stumps of the same nerves, in a cobaltic lysine complex solution, and the distribution of efferent neurons sending their axons into these nerves was investigated in serial sections of the medulla and the cervical spinal cord in young rats. The following neuron groups were identified. The inferior salivatory nucleus lies in the dorsal part of the tegmentum at the rostral part of facial nucleus. It consists of a group of medium-sized and a group of small neurons. Their axons make a hair-pin loop at the midline and join the glossopharyngeal nerve. The dorsal motor nucleus of the vagus situates in the dorsomedial part of the tegmentum. Its rostral tip coincides with the first appearance of sensory fibres of the glossopharyngeal nerve, the caudal end extends into the pyramidal decussation. The constituting cells have globular or fusiform perikarya and they are the smallest known efferent neurons. The ambiguous nucleus is in the ventrolateral part of the tegmentum. The rostral tip lies dorsal to the facial nucleus, and the caudal tip extends to the level of the pyramidal decussation. The rostral one third of the ambiguous nucleus is composed of tightly-packed medium sized neurons, while larger neurons are arranged more diffusely in the caudal two thirds. The long dendrites are predominantly oriented in the dorsoventral direction. The dorsally-oriented axons take a ventral bend anywhere between the ambiguous nucleus and dorsal motor nucleus of the vagus. The motoneurons of the accessorius nerve are arranged in a medial, a lateral and a weak ventral cell column. The medial column begins at the caudal aspect of the pyramidal decussation and terminates in C2 spinal cord segment. The lateral and ventral columns begin in C2 segment and extend into C6 segment. The neurons have large polygonal perikarya and characteristic cross-shaped dendritic arborizations. The axons follow a dorsally-arched pathway between the ventral and dorsal horns. The accessorius motoneurons have no positional relation to any of the vagal efferent neurons. It is concluded that the topography and neuronal morphology of accessorius motoneurons do not warrant the designation of a bulbar accessorius nucleus and a bulbar accessorius nerve.     

Swallowing reflex and brain stem neurons activated by superior laryngeal nerve stimulation in the mouse

The purpose of the present study was to identify vagal subnuclei that participate in reflex swallowing in response to electrical stimulation of the left superior laryngeal nerve (SLN). SLN stimulation at 10 Hz evoked primary peristalsis, including oropharyngeal and esophageal peristalsis, and LES relaxation. It also induced c-fos expression in interneurons in the interstitial (SolI), intermediate (SolIM), central (SolCe), dorsomedial (SolDM) and commissural (SolC) solitary subnuclei. Neurons in parvicellular reticular nucleus (PCRt) and area postrema (AP) and motoneurons in the semicompact (NAsc), loose (NAl), and compact (NAc) formations of the nucleus ambiguus and both rostral (DMVr) and caudal (DMVc) parts of the dorsal motor nucleus of vagus were also activated. The activated neurons represent all neurons concerned with afferent SLN-mediated reflexes, including the swallowing-related neurons. SLN stimulation at 5 Hz elicited oropharyngeal and LES but not esophageal responses and evoked c-fos expression in neurons in SolI, SolIM, SolDM, PCRt, AP, NAsc, NAl, and DMVc but not in SolCe, NAc, or DMVr. These data are consistent with the role of SolI, SolIM, SolDM, NAsc, NAl, and DMVc circuit in oropharyngeal peristalsis and LES relaxation and SolCe, NAc, DMVc, and DMVr in esophageal peristalsis and LES responses.     

Efferent fibres to the aortic bodies: a search for their cell bodies in the brainstem of the cat and rabbit

An attempt has been made to determine where in the lower brainstem the cell bodies of nonsympathetic efferent fibres in the aortic nerve of the cat and rabbit are located. Horseradish peroxidase (HRP) was placed on the central end of the right cut aortic nerve of anaesthetized animals and, after an appropriate time, sections of the brainstem encompassing the rostral and caudal limits of the dorsal vagal motor nucleus and nucleus ambiguus were examined microscopically for retrogradely transported HRP. Cell bodies labelled by exogenous HRP were not found in any of the cats or rabbits exposed to HRP although reaction product, due to an endogenous response, was observed. Appropriate control experiments were performed to show that the sensitivity of the technique for demonstrating HRP in our hand was adequate. We conclude that the cell bodies of efferent fibres, of non sympathetic origin, in the aortic nerve are likely to be located outside the central nervous system.     

Possible mechanisms of the anterior limbic cortex influence on the neuron activity of the vago-solitary complex

In ananesthetized cats, neurons of the nucleus of the tractus solitarius (NTS) and the dorsal motor nucleus of the vagus nerve (DMNV) revealed phasic excitatory responses to separate single vagal and cortical stimuli. Stimulation of the anterior limbic cortex combined with vagal stimulation resulted in inhibitory or excitatory modification of the vagal induced responses of the NTS and DMNV neurons. The data obtained suggest that complete inhibitory effects are related to general cortical mechanisms of control of the functional state of the brain stem visceral neurons. Selective inhibition of the vagal induced responses by limbic cortex stimulation is due to particular cortical mechanisms of the visceral sensory transmission control via the NTS neurons.     

Vestibular efferent neurons forming projections to the saccule in the guinea pig

A comparative analysis was made of the distribution of vestibular efferent neurons projecting to the saccule and efferent cells sending out axons to the auditory nerve ("cochlear efferent neurons") in the guinea pig, using retrograde horseradish peroxidase axonal transport techniques. Saccular efferent neurons were discovered bilaterally in the subependymal granular layer at the base of the fourth cerebral ventricle and laterally to the facial nerve genu ispsilaterally in the parvocellular reticular nucleus, as well as nuclei of the superior olivary complex: the lateral olivary nucleus and lateral nucleus of the trapezoid body. Cochlear efferent neurons are located ipsilaterally in the pontine reticular caudal nucleus, in the anteroventral cochlear nucleus, and in the lateral and medial olivary nuclei. Neurons were found contralaterally in the medial nucleus of the trapezoid body. It thus emerged that location zones of vestibular saccular efferent neurons and those of cochlear efferent units partially overlapped. The possible involvement of saccular vestibular efferent neurons in the mechanisms of auditory perception is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 22, No. 5, pp. 657–665, September–October, 1990.     

Prenatal development of the human nucleus ambiguus during the embryonic and early fetal periods

The ontogenetic development of the nucleus ambiguus was studied in a series of human embryos and fetuses ranging from 3 to 12.5 weeks of menstrual age (4 to 66 mm crown-rump length). They were prepared by Nissl and silver methods. Nucleus ambiguus neuroblasts, whose neurites extend towards and into the IXth and rostral Xth nerve roots, appear in the medial motor column of 4-6-week-old embryos (4.25-11 mm). These cells then migrate laterally (6.5 weeks, 14 mm) to a position near the dorsal motor nucleus of X. At 7 weeks (15 mm), nucleus ambiguus cells begin their migration, which progresses rostrocaudally, into their definitive ventrolateral position. The basic pattern of organization of the nucleus is established in its rostral region at 8 weeks (22.2-24 mm) and extends into its caudal region by 9 weeks (32 mm), when its nearly adult organization is evident. Cells having the characteristics of mature neurons first appear rostrally in the nucleus during the 8.5-9-week period (24.5-32 mm), gradually increase in number, and constitute the entire nucleus at 12.5 weeks (65.5 mm). Definitive neuronal subgroups first appear at 10 weeks (37.5 mm) in the large rostral nuclear region. These features suggest that the human nucleus ambiguus develops along a rostrocaudal temporospatial gradient. Evidence indicates that function of nucleus ambiguus neurons, manifested by fetal reflex swallowing, occurs after the cells migrate into their definitive position, establish the definitive nuclear pattern, and exhibit mature characteristics.     

刺激家兔腹部迷走神经外周端降压效应中枢机制的研究

应用电解损毁和脑室内注射药物的方法研究了刺激家兔腹部迷走神经外周端所致降压效应的中枢机制。结果表明:1.电刺激延脑闩部尾侧1.5—2mm、中线旁开0.25mm、深1—2mm 处主要引起降压反应。2.电解损毁该部位可以使刺激腹部迷走神经外周端所引起的降压效应显著减弱(n=20,P<0.001),但对刺激减压神经所致降压反应无影响。3.在延脑闩部水平电解损毁减压神经纤维在孤束核的主要投射区可以使刺激减压神经所致降压反应显著减弱,而对刺激腹部迷走神经外周端所致降压反应无影响。4.第四脑室注射5,6-双羟色胺的动物较之注射人工脑脊液的动物颈、胸髓5-羟色胺含量明显降低、动物动脉压增高、心率明显增快、刺激减压神经所致降压反应未见减弱,而刺激腹部迷走神经外周端所致降压反应却明显减小。因此,我们认为家兔腹部迷走神经外周端所致降压效应依赖于延脑闩下部的中缝隐核及连合核等结构,而与减压神经的投射部位无关。延脑中缝核至脊髓的下行性5-HT能神经纤维抑制脊髓交感节前神经元的活动,是这个降压效应的中枢机制之一。     

Musings on the wanderer: what's new in our understanding of vago-vagal reflex? IV. Current concepts of vagal efferent projections to the gut

Vagal efferents, consisting of distinct lower motor and preganglionic parasympathetic fibers, constitute the motor limb of vagally mediated reflexes. Arising from the nucleus ambiguus, vagal lower motor neurons (LMN) mediate reflexes involving striated muscles of the orad gut. LMNs provide cholinergic innervation to motor end plates that are inhibited by myenteric nitrergic neurons. Preganglionic neurons from the dorsal motor nucleus implement parasympathetic motor and secretory functions. Cholinergic preganglionic neurons form parallel inhibitory and excitatory vagal pathways to smooth muscle viscera and stimulate postganglionic neurons via nicotinic and muscarinic receptors. In turn, the postganglionic inhibitory neurons release ATP, VIP, and NO, whereas the excitatory neurons release ACh and substance P. Vagal motor effects are dependent on the viscera's intrinsic motor activity and the interaction between the inhibitory and excitatory vagal influences. These interactions help to explain the physiology of esophageal peristalsis, gastric motility, lower esophageal sphincter, and pyloric sphincter. Vagal secretory pathways are predominantly excitatory and involve ACh and VIP as the postganglionic excitatory neurotransmitters. Vagal effects on secretory functions are exerted either directly or via release of local mediators or circulating hormones.     

On the anatomical organization of the vagal nuclei

The neurons of origin of the right vagus and its components in both the monkey (Macaca fascicularis) and albino rats were localized by the retrograde transport of horseradish peroxidase (HRP) applied to the stomach wall, the vagal trunk and its recurrent laryngeal branch. An attempt was also made to localize the neurons forming the superior laryngeal nerve and those supplying the thoracic organs by a combination of operative procedures. The results showed that the stomach was innervated by neurons distributed throughout the entire rostrocaudal extent of the dorsal motor nucleus (DMN) on both sides of the brain stem. Neurons scattered throughout the entire extent of the DMN and nucleus ambiguus (NA) supplied the thoracic viscera. There did not appear to be any topographic arrangement in the DMN neurons supplying the abdominal and thoracic viscera as reported by other workers, and there was no clear evidence of crossing of vagal fibers in the monkey brain stem, though such crossing was seen in the rat brain stem. Both the superior and inferior ganglia of the vagus nerve were labeled following application of HRP to the vagal trunk. Neurons in the caudal part of the NA gave rise to fibers in the ipsilateral recurrent laryngeal nerve, at least on the right side. The neurons giving rise to the superior laryngeal nerve could not be delineated in this study. In all the experimental procedures described, the hypoglossal nucleus was labeled only after applying HRP to the hypoglossal nerve.     

Arrangement and connections of mesencephalic trigeminal neurons in the rat

The morphology of the mesencephalic trigeminal nucleus was examined microscopically in serial frozen sections. The nucleus extends over a length of about 4.5 mm, and its cell number was calculated to range from 1,000 to 1,600. 60% of the cells were located in the caudal third of the nucleus. Clustering of large unipolar cells was seen throughout the nucleus. Small spindle-shaped multipolar cells were found in the pontine part of the nucleus. The efferent connections of the mesencephalic trigeminal neurons were investigated by means of iontophoretically delivered Phaseolus vulgaris leuco-agglutinin or horseradish peroxidase after electrophysiological identification of mesencephalic trigeminal neurons. All projections were found ipsilateral to the injection site; they were confined to the trigeminal motor nucleus, especially to its lateral part, and to the dorsolateral reticular formation. The latter projection area included the supratrigeminal nucleus, the nucleus of Probst, and the parvocellular reticular zone. There were no direct projections to the facial or hypoglossal motor nuclei. It is concluded that proprioceptive input from one side is mediated polysynaptically to the bilateral oral final common-path neurons, with the exception of the ipsilateral trigeminal motoneurons.     

Establishment of vagal sensorimotor circuits during fetal development in rats

The differentiation of vagal motor neurons and their emerging central relationship with vagal sensory afferents was examined in fetal rats. To identify peripherally projecting sensory and motor neurons, 1,1′-dioctadecyl 3,3,3′,3′-tetramethylindocarbocyanine perchloarate (DiI) was inserted into the proximal gut or cervical vagus nerve in fixed preparations. At embryonic day (E) 12, labeled vagal sensory neurons are present in the nodose ganglia and a few sensory axons project into the dorsolateral medulla. Central sensory processes become increasingly prevalent between E13 and E14 but remain restricted to the solitary tract. Vagal motor neurons are first labeled at E13, clustered within a region corresponding to the nucleus ambiguus (NA). Additional motor neurons appear to be migrating toward the NA from the germinal zone of the fourth ventricle. Motor neurons in the dorsal motor nucleus of the vagus (DMV) first project to the gut at E14 and have processes that remain in physical contact with the ventricular zone through E16. Sensory axons emerge from the solitary tract at E15 and project medially through the region of the nucleus of the solitary tract (NST) to end in the ventricular zone. A possible substrate for direct vagovagal, sensorimotor interaction appears at E16, when vagal sensory fibers arborize within the DMV and DMV dendrites extend into the NST. By E18, the vagal nuclei appear remarkably mature. These data suggest specific and discrete targeting of vagal sensory afferents and motor neuron dendrites in fetal rats and define an orderly sequence of developmental events that precedes the establishment of vagal sensorimotor circuits. © 1993 John Wiley & Sons, Inc.     

The quantitative morphological characteristics of the spinal neurons in the kitten developing under conditions of limited afferent input]

Golgi preparations of cervical part of the spinal cord were used to study sparely and densely branching neurons of lamina VII, sparely and densely branching neurons of lamina VIII and large densely branching motor neurons of lamina IX in 30-day kittens developing under the conditions of limited reticulospinal input. Limitation was induced by electrocoagulation of bulbar magnocellular nucleus and caudal reticular pontine nucleus performed at the 7th day. Computerized morphometry revealed that partial deafferentation affects the geometry in all studied cell types except densely branching neurons of lamina VIII. Adaptive nature of structural reorganization of spinal neurons and possible relationship between morphological properties and functional profile of cells are discussed.     

Transneuronal labeling of neurons in the adult rat central nervous system following inoculation of pseudorabies virus into the colon

Transneuronal tracing with pseudorabies virus (PRV) was used to identify sites in the central nervous system involved in the neural control of colon function. PRV-immunoreactive (IR) cells were primarily localized to the caudal lumbosacral (L6-S1) and caudal thoracic-rostral lumbar (T13-L1) spinal segments with the distribution varying according to survival time (72-96 h). In the lumbosacral spinal cord at all time points examined, significantly (PА.005) greater numbers of PRV-IR cells were present in the region of the sacral parasympathetic nucleus (SPN) of the S1 spinal segment compared to that of the L6 segment. These studies also revealed morphologically distinct cell types with a differential distribution (probably interneurons and preganglionic parasympathetic neurons) in the region of the SPN in the L6-S1 spinal segments following colon inoculation. PRV-labeled neurons were located at various levels of the neuraxis and at many sites had a distribution similar to that following injection of virus to other urogenital organs. However, some unique sites in the dorsal motor nucleus of the vagus, nucleus of the solitary tract, nucleus ambiguus and area postrema were also identified. To determine if labeling in these caudal medullary sites was mediated by spinal or vagal pathways, the colon was inoculated with PRV in animals with a complete spinal cord (T8) transection (5-7 days prior). Following spinal transection, PRV-infected cells were detected in the same caudal medullary regions; however, labeling in other regions (e.g., Barrington's nucleus) was eliminated or significantly reduced. These studies have yielded several novel observations concerning the central neural control of colonic function: (1) the preganglionic efferent and primary afferent innervation of the colon arises primarily from the S1 spinal segment; (2) the distribution of PRV-infected neurons in the central nervous system following colon inoculation was similar to that following PRV inoculation of other urogenital organs; (3) Barrington's nucleus, which has been identified previously as the pontine micturition center, may have a role in colonic function; and (4) PRV infection in Barrington's nucleus following colon inoculation is mediated by bulbospinal pathways whereas labeling in caudal medullary regions is mediated, at least in part, by vagal pathways.     

Early ontogeny of the vagus nerve: an analysis of the medulla oblongata and cervical spinal cord of the postnatal rat.

Retrograde transport of cholera toxin conjugated with horseradish peroxidase in the postnatal rat has revealed remarkable features of dendritic fields of vagal motor neurons in the medulla oblongata and cervical spinal cord during the period of early development (0-10 days). At birth, vagal motor neurons in the dorsal motor nucleus of the vagus, nucleus ambiguus, nucleus retroambigualis, nucleus dorsomedials and the spinal nucleus of the accessory nerve are small with relatively few, unbranched processes. The span of the dendritic tree is much smaller than that found in adult animals. By the postnatal Day 2 there are marked changes in the soma as well as in the dendritic tree of these neurons. There is dispersion of the cell bodies within the neuropil as well as an expansion of the total area of the brain stem occupied by these motor neurons and their dendritic processes which show extensive growth and branching. By postnatal Day 3 the most extensive proliferation of these neurons is seen and appears to represent the peak of dendritic growth of vagal motor neurons such that the area occupied by the dendritic tree of a single neuron is three times that seen in an adult rat. This proliferation gradually decreased during the subsequent seven days of early development (i.e. Days 4-10) so that by Day 10 the dendritic span of vagal motor neurons was reduced to about twice the adult size. This growth progressively decreased from Days 10 to 30 at which time adult levels were reached. Ultrastructural examination of these horseradish peroxidase labeled dendrites showed a positive correlation between the number of dendritic processes and the number of axo-dendritic synapses. This was accompanied by an increase in the number of identifiable synaptic junctions. These morphological complexities observed during the period of early development of vagal motor neurons indicate that the vagus nerve undergoes dramatic changes during the period of early development including the establishment of numerous synaptic contacts between vagal afferents and efferents in the brainstem. A number of these changes occur in developing dendritic fields of vagal motor neurons during the first three days of neonatal life. It is reasonable to assume that developmental abnormalities during this "critical period" could produce significant functional changes in the pattern of respiration as well as in the control of airway smooth muscle.     

Chemical profile of vagal preganglionic motor cells innervating the airways in ferrets: the absence of noncholinergic neurons.

In ferrets, we investigated the presence of choline acetyltransferase (ChAT), vasoactive intestinal peptide (VIP), and markers for nitric oxide synthase (NOS) in preganglionic parasympathetic neurons innervating extrathoracic trachea and intrapulmonary airways. Cholera toxin beta-subunit, a retrograde axonal transganglionic tracer, was used to identify airway-related vagal preganglionic neurons. Double-labeling immunohistochemistry and confocal microscopy were employed to characterize the chemical nature of identified airway-related vagal preganglionic neurons at a single cell level. Physiological experiments were performed to determine whether activation of the VIP and ChAT coexpressing vagal preganglionic neurons plays a role in relaxation of precontracted airway smooth muscle tone after muscarinic receptor blockade. The results showed that 1) all identified vagal preganglionic neurons innervating extrathoracic and intrapulmonary airways are acetylcholine-producing cells, 2) cholinergic neurons innervating the airways coexpress ChAT and VIP but do not contain NOS, and 3) chemical stimulation of the rostral nucleus ambiguus had no significant effect on precontracted airway smooth muscle tone after muscarinic receptor blockade. These studies indicate that vagal preganglionic neurons are cholinergic in nature and coexpress VIP but do not contain NOS; their stimulation increases cholinergic outflow, without activation of inhibitory nonadrenergic, noncholinergic ganglionic neurons, stimulation of which induces airway smooth muscle relaxation. Furthermore, these studies do not support the possibility of direct inhibitory innervation of airway smooth muscle by vagal preganglionic fibers that contain VIP.     

Efferent neurons of the lateral-line system and the VIII cranial nerve in the brainstem of anurans

Summary The octavo-lateral efferent system of several anuran species was studied by means of retrograde transport of horseradish peroxidase. This system is organized similarly in all larval anurans and in all adult aglossids. All have two groups of efferent neurons in the nucleus reticularis medialis between the VIIIth and the IXth motor nucleus. The caudal group consists of efferent neurons that supply the posterior lateral-line nerve (NLLp) and a considerably smaller group of neurons supplying both the NLLp and the anterior lateral-line nerve (NLLa). The rostral group is composed of efferent neurons supplying the NLLa, neurons projecting to the inner ear and neurons supplying both the inner ear and the NLLa. Efferent neurons of the VIIIth cranial nerve exhibit a rostrocaudal cytoarchitectonic differentiation. Caudal perikarya, which are rounder in shape than those of the rostral part, have a dendritic projection to the superior olive. It is suggested that this differentiation reflects a functional differentiation of acoustic and vestibular efferent neurons.Labeled neurons were ipsilateral to the site of application of HRP. None were found in the vestibular nuclei or in the cerebellum.Efferent axons projecting to neuromasts of the NLLa leave the medulla with the VIIth nerve, axons projecting to neuromasts of the NLLp exit via the IXth nerve. Cell counts and the observation of axonal branching revealed that efferent units of both the lateral-line and the VIIIth-nerve system supply more than one receptor organ. In contrast to the lateral-line system, dendrites of efferent neurons of the VIIIth nerve project dorsally onto its nuclei, and afferents of the VIIIth nerve project onto efferent neurons. These structures most probably represent a feedback loop between the afferent and efferent systems of the VIIIth cranial nerve.     

Brainstem neurons projecting to the ampullae of anterior,lateral, and posterior semicircular canals in the guinea pig

Neurons projecting to the ampullae of anterior, lateral, and posterior semicircular canals were identified in the guinea pig brainstem using horseradish peroxidase labeling techniques. Two groups of neurons forming bilateral connections were found, one located dorsally and the other ventrally to facial nerve trajectories. The dorsal group of vestibular efferent neurons projecting to all three canals was detected in the subependymal granular layer of the floor of the 4th ventricle lateral to the facial nerve genu and in the abducent nerve nucleus. Efferent neurons belonging to the ventral group were unevenly distributed through different areas of the parvocellularis nucleus and the rostral portion of the pontine caudal reticular nucleus. The morphological characteristics and distribution pattern of these cells are taken as confirmation of their heterogeneity of neuronal and functional organization in the vestibular efferent system of semicircular canal ampullae.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 20, No. 4, 1988, pp. 526–532.     

Possible mechanisms of amygdala participation in the regulation of gastric motor activity

Mechanisms of the amygdala central nucleus (CNA) influence on gastric motor reflex activity were studied in electrophysiological and neuroanatomical experiments in Wistar rats. In the anaesthetized animals, electrical stimulation of the CNA affected spontaneous gastric motility and caused inhibitory as well as excitatory changes of vagus-induced gastric relaxation. The most significant and mainly inhibitory effects were observed under the stimulation of the medial CNA. Microinjection of the anterograde tracer Phaseolus vulgaris-leucoagglutimn (PHA-L) into the different divisions of the CNA revealed direct projections from its dorso-medial portion to the gastric related area of the dorsal vagal complex. Electrical stimulation of this amygdaloid area was found to change activity of the bulbar gastric related neurons. Inhibitory and excitatory changes of their vagus-induced responses under the amigdala stimulation were manifested as a general modulation of all phases of the reaction or a selective modulation of some of them. These mechanisms may underlie the amygdalo-fugal modulation of gastric motor reflex activity.     

Afferent and efferent connections of the area postrema demonstrated by the horseradish peroxidase method

The previous autoradiographic study of the efferent connections of the arca postrema have been completed by using horseradish peroxidase (H.R.P.) as a marking agent. These two labelling techniques have made it possible to determine two categories of structures connected with A.P.: those with afferent connections (Nucleus ambiguus), or efferent connections (mesencephalic nucleus of V, Locus coeruleus, griseum centrale and inferior and superior colliculi) and those having both types of connections (Nucleus tractus solitarii (N.F.S.), Dorsal vagal nucleus, nucleus intercalatus, hypoglossal nucleus). The afferent and efferent pathways of the A.P. identified with H.R.P. coincide with the catecholaminergic pathways demonstrated in the rat by Lindvall and Bj?rklund (1974).