Neuronal architecture of the nucleus of the solitary tract in the hamster

This study provides a scheme for subdividing the nucleus of the solitary tract of the hamster on the basis of cytoarchitectonic criteria, cell measurements, and neuronal cell types identified with the Golgi method. Reduced silver-stained sections revealed the feltlike neuropil that characterizes the nucleus of the solitary tract and were used to define the boundaries of the nuclear complex. Adjacent sections stained for Nissl substance revealed ten subdivisions, each with a characteristic neuronal architecture based on cell sizes, shapes, and packing density. Some subdivisions, e.g., the ventral and medial subnuclei, were identified at all rostrocaudal levels of the nuclear complex, while other subdivisions, e.g., the caudally located dorsolateral and ventrolateral subnuclei, were restricted to particular levels. Golgi preparations were counterstained for Nissl substance, thus allowing dendro- and cytoarchitecture to be compared directly. This material permitted the identification of a number of functionally relevant features of the neuronal constituents of the subdivisions. This approach, employing three cytological methods, has permitted the assembly of a detailed atlas of the nucleus of the solitary tract. The subdivisions of the present atlas have been compared with their likely counterparts identified in previous investigations of the mammalian nucleus of the solitary tract. In order to relate cytoarchitecture with primary afferent termination sites and to define the gustatory-recipient subdivisions, the differential relationships of the subdivisions with lingual afferent projections in the hamster are also described. The present parcellation scheme is intended to facilitate anatomical and physiological investigations of the types of circuits that compose the medullary gustatory and general visceral sensory systems.     

Direct cortical projections to the parabrachial nucleus in the cat

Direct projections from the cerebral cortex to the parabrachial nucleus in the cat were examined by the horseradish peroxidase (HRP)method. When HRP was injected into the parabrachial nucleus, retrogradely labeled neuronal cell bodies were seen, bilaterally with an ipsilateral predominance, mainly in the orbital gyrus, the lateral bank of the presylvian sulcus, and a restricted region in the infralimbic cortex on the medial surface of the frontal lobe (stereotaxic coordinates; Fr: 22, L: 1, H: -1); all labeled neurons were in deep pyramidal cell layer. After injecting HRP conjugated to wheat germ agglutinin (WGA-HRP) into the cortical regions where retrogradely labeled neurons were found after injecting HRP into the parabrachial nucleus, anterogradely labeled cortical fibers were traced to the parabrachial nucleus. Corticoparabrachial fibers originating from the orbital gyrus and the lateral bank of the presylvian sulcus ran ipsilaterally through the internal capsule and the cerebral peduncle down to the lower brainstem, whereas those from the infralimbic cortex coursed down ipsilaterally through the medial forebrain bundle. These cortical fibers to the parabrachial nucleus were distributed bilaterally with an ipsilateral predominance. Cortical fiber terminals in the parabrachial nucleus were topographically arranged: Corticoparabrachial fibers from the lateral bank of the presylvian sulcus ended most massively in the dorsal part of the lateral parabrachial nucleus. Corticoparabrachial fibers from the orbital gyrus ended most heavily in the medial parabrachial nucleus and less heavily in the lateral parabrachial nucleus. Corticoparabrachial fibers from the infralimbic cortex ended mostly in the parabrachial regions surrounding the brachium conjunctivum.     

Sodium deprivation and salt intake activate separate neuronal subpopulations in the nucleus of the solitary tract and the parabrachial complex

Salt intake is an established response to sodium deficiency, but the brain circuits that regulate this behavior remain poorly understood. We studied the activation of neurons in the nucleus of the solitary tract (NTS) and their efferent target nuclei in the pontine parabrachial complex (PB) in rats during sodium deprivation and after salt intake. After 8-day dietary sodium deprivation, immunoreactivity for c-Fos (a neuronal activity marker) increased markedly within the aldosterone-sensitive neurons of the NTS, which express the enzyme 11-beta-hydroxysteroid dehydrogenase type 2 (HSD2). In the PB, c-Fos labeling increased specifically within two sites that relay signals from the HSD2 neurons to the forebrain--the pre-locus coeruleus and the innermost region of the external lateral parabrachial nucleus. Then, 1-2 hours after sodium-deprived rats ingested salt (a hypertonic 3% solution of NaCl), c-Fos immunoreactivity within the HSD2 neurons was virtually eliminated, despite a large increase in c-Fos activation in the surrounding NTS (including the A2 noradrenergic neurons) and area postrema. Also after salt intake, c-Fos activation increased within pontine nuclei that relay gustatory (caudal medial PB) and viscerosensory (rostral lateral PB) information from the NTS to the forebrain. Thus, sodium deficiency and salt intake stimulate separate subpopulations of neurons in the NTS, which then transmit this information to the forebrain via largely separate relay nuclei in the PB complex. These findings offer new perspectives on the roles of sensory information from the brainstem in the regulation of sodium appetite.     

Optic afferents to the parabrachial nucleus

Following intraocular injection of cholera toxin subunit B (CTB), optic afferents to the dorsal pontine region were observed in Mongolian gerbils, Chilean degus, and laboratory rats. CTB-positive optic axons emerge at the caudal pole of the superior colliculus, descend through the periaqueductal gray, and innervate the lateral parabrachial nucleus. This projection appears to be a continuation of the retinal pathway that innervates the dorsal raphe nucleus in these same species.     

电刺激大鼠杏仁中央核对脑桥臂旁核味觉神经元的影响

利用电生理学方法观察了电刺激杏仁中央核对脑桥臂旁核味觉神经元的影响.结果表明:电刺激杏仁中央核抑制大部分臂旁核味觉神经元的活动,并且提高臂旁核味觉神经元对五种基本味觉刺激反应的特异性.电刺激杏仁中央核对臂旁核的抑制作用以对盐酸和奎宁刺激的反应尤为明显(P＜0.01),并且对这两种厌味刺激反应的抑制作用是基本一致的.本研究的结果提示,杏仁中央核可能通过抑制脑干味觉神经元对厌味刺激的反应,从而参与对摄食行为的调控.     

电刺激大鼠杏仁中央核对脑桥臂旁核味觉神经元的影响(英文)

利用电生理学方法观察了电刺激杏仁中央核对脑桥臂旁核味觉神经元的影响。结果表明 :电刺激杏仁中央核抑制大部分臂旁核味觉神经元的活动 ,并且提高臂旁核味觉神经元对五种基本味觉刺激反应的特异性。电刺激杏仁中央核对臂旁核的抑制作用以对盐酸和奎宁刺激的反应尤为明显 (P <0 .0 1) ,并且对这两种厌味刺激反应的抑制作用是基本一致的。本研究的结果提示 ,杏仁中央核可能通过抑制脑干味觉神经元对厌味刺激的反应 ,从而参与对摄食行为的调控     

A subpopulation of neurons in the rat rostral nucleus of the solitary tract that project to the parabrachial nucleus express glutamate-like immunoreactivity

In rodents, gustatory information is transmitted from second order neurons in the rostral nucleus of the solitary tract (rNST) to the parabrachial nucleus (PBN) in the pons. The chemical nature of this projection is unknown. Therefore, the goal of the current study was to determine if rNST neurons that project to the PBN express glutamate-like immunoreactivity. Projection neurons were retrogradely labeled following stereotaxic injection of rhodamine-filled latex microspheres into the right PBN of seven rats while glutamate-immunoreactive (GLU-IR) structures were visualized in the same tissue using an immunoperoxidase procedure. The number of single- and double-labeled neurons located in the right (ipsilateral) and left rNST, in each of the nuclear subdivisions as well as their position along the rostral-caudal axis of the rNST was determined. GLU-IR cell bodies were located throughout the rNST. Although the rostral central subdivision contained the highest percentage (33.8%) of GLU-IR perikarya, immunolabeled neurons were most concentrated (number/area of subdivision) within the medial subnucleus. The rostral third of the rNST contained the fewest (20. 5%) and lowest density of GLU-IR cell bodies. The highest percentage of rNST neurons retrogradely labeled from the PBN were located ipsilateral (85.4%) to the pontine injection site, in the middle third of the nucleus (44.2%) and within the rostral central subdivision (52.4%). Overall, 18% of the labeled rNST projection neurons were GLU-IR. The distribution of double-labeled neurons mirrored that of the projection neurons with the largest number located in the ipsilateral rNST (84.5%), middle third of the nucleus (40.5%) and rostral central subdivision (64.7%). These results indicate that glutamate may be a main component of the ascending pathway from the rNST to the PBN. In addition, since GLU-IR neurons were located throughout the rNST and most were not retrogradely-labeled, the current results suggest that glutamate may be an important neurotrans-mitter within the medulla.     

Electrophysiological identification of neurons in the parabrachial nucleus projecting directly to the hypothalamus in the rat

Experiments were done in urethane anesthetized rats to identify single units in the region of the parabrachial nucleus (PBN) projecting directly to ‘cardiovascular’ responsive sites in either the paraventricular nucleus of the hypothalamus (PVH) or the supraoptic commissure and nucleus (SOC-SON) region. Fifty-five single units were antidromically activated in the ipsilateral PBN by electrical stimulation of either the PVH (n = 27) or SOC-SON region (n = 28) with latencies corresponding to conduction velocities of 0.3–5.1 m/s. The axons of PBN units projecting to the PVH conducted at significantly slower velocities (0.5 ± 0.04m/s) than those projecting to the SOC-SON region (1.6 ± 0.25m/s). These data suggest that aacending fibers from the PBN to the PVH are unmyelinated, whereas those to the SOC-SON region are primarily a little myelinated. In addition, since the PBN is known to receive cardiovascular and visceral afferent inputs, it is suggested that these neurons likely function in relaying this afferent information to hypothalamic areas involved in autonomic regulation.     

代谢型谷氨酸受体在大鼠脑干臂旁核中的分布

采用免疫组织化学方法观察了大鼠脑干臂旁核中代谢型谷氨酸受体亚型的分布情况。结果表明：代谢型谷氨酸受体１α亚型免疫阳性神经元仅分布于小脑上脚内侧极处,外内亚核、外外亚核、内外亚核及腰区中只有弱免疫阳性的神经毡;上外亚核、外外亚核及外内亚核中有浓密至中等强度的代谢型谷氨酸受体５亚型免疫阳性神经毡的分布;此外,内外亚核中尚有少量代谢型谷氨酸受体７亚型弱免疫阳性神经元。而代谢型谷氨酸受体２／３亚型在臂旁核所有亚核中均未见分布。本研究结果提示不同亚型的代谢型谷氨酸受体可能参与臂旁核对不同生理过程的调控。     

The organization of the efferent projections of the parabrachial nucleus to the forebrain in the rat: A retrograde fluorescent double-labeling study

The organization of the efferent projections of the parabrachial nucleus (PBN) to the forebrain has been investigated in the rat by means of combined injections of two fluorescent retrograde tracers: red fluorescent Evans Blue and a blue fluorescent mixture of 4′,6′-diamidino-2-phenylindol 2 HCl and primuline. First, the distributions of retrogradely labeled neurons in the PBN after bilateral injections of tracers in the central nucleus of the amygdala (CNA) was examined. The CNA on one side of the brain was injected with one of the tracers and the CNA on the opposite side of the brain was injected with the other tracer. Next, the distributions of labeled neurons were examined after bilateral ventral medial thalamus (VMT) injections. Finally, the retrograde labeling of the PBN was studied after combined ipsilateral injections of one tracer in the CNA and the other tracer in the VMT. After the various injections, characteristic distributions of populations of labeled neurons within the PBN were seen. Double-labeled neurons were present only after bilateral VMT injections. From this it was concluded that the PBN projections to the VMT in the rat are bilateral. Based on the relative distributions of populations of retrogradely labeled neurons in the PBN, it was suggested that the PBN projects primarily taste information to the VMT and mainly visceral information to the CNA. This transfer of information to the forebrain is discussed.     

Maturation of neuron types in nucleus of solitary tract associated with functional convergence during development of taste circuits

Late fetal through postnatal development in sheep is a period of increasing convergence of afferent taste fibers onto second-order neurons in the nucleus of the solitary tract (NST). To learn whether neuron morphology alters in concert with convergence and neurophysiological development in NST, three-dimensional neuron reconstructions were made of cells in a functionally defined region of gustatory NST from Golgi preparations of the brainstem. Elongate, multipolar, and ovoid neurons were studied in fetuses from 85 days of gestation through the perinatal period (term = 147 days of gestation), to postnatal stages. Somal size and form, and dendritic complexity and extent, increased markedly from 85 to about 110 days of gestation in both of the proposed NST projection neurons, elongate and multipolar. From 130 days of gestation to postnatal ages, growth of dendrites of elongate neurons plateaued or declined, whereas dendrites of multipolar neurons apparently continued to increase in size and extent. In addition, spine density decreased on elongate neurons but remained stable on multipolar neurons. Morphological variables of ovoid cells, proposed interneurons in NST, did not alter over this later period. The data suggest that multipolar, not elongate or ovoid, neurons are logical candidates to receive the increasing afferent fiber input onto NST cells during late gestation. Also, neural activity from taste afferent fibers is more likely to have a role in altering NST neuron morphology at later, rather than earlier, developmental periods. © 1994 Wiley-Liss, Inc.     

Transfer of information about taste from the nucleus of the solitary tract to the parabrachial nucleus of the pons

In the study of the neural code for gustation, the relative sensitivity of a cell to a variety of taste stimuli is defined as its response profile. To study the construction of response profiles from incoming signals, electrophysiological responses to NaCl, HCl, quinine-HCl, sucrose and Na saccharin were recorded simultaneously in pairs of single cells: one in the nucleus of the solitary tract (NTS) and the other in the parabrachial nucleus of the pons (PbN), respectively the first and second synapses in the central pathway for gustation. Of 37 units recorded in the NTS and 32 in the PbN, 12 (32%) pairs showed evidence of functional connectivity. Although PbN responses were significantly larger than those in the NTS in general, no amplification of NTS activity was apparent among those units that were functionally connected. Analysis of NTS–PbN connectivity patterns suggests that PbN units receive input from NTS units with response profiles that are both similar and different from their own pattern of sensitivities. Further analysis suggests that the stimulus-selectivity of the response profile of a PbN unit may be determined by stimulus-selective input from NTS cells that show similar response profiles. However, input from NTS cells with response profiles different from their own appears to be non-stimulus-selective. Analysis of the organization of response profiles in the two structures suggests that the cells in both the NTS and PbN cannot be easily distinguished by their patterns of sensitivity to taste stimuli.     

Connections of the parabrachial nucleus with the nucleus of the solitary tract and the medullary reticular formation in the rat.

We examined the subnuclear organization of projections to the parabrachial nucleus (PB) from the nucleus of the solitary tract (NTS), area postrema, and medullary reticular formation in the rat by using the anterograde and retrograde transport of wheat germ agglutinin-horseradish peroxidase conjugate and anterograde tracing with Phaseolus vulgaris-leucoagglutinin. Different functional regions of the NTS/area postrema complex and medullary reticular formation were found to innervate largely nonoverlapping zones in the PB. The general visceral part of the NTS, including the medial, parvicellular, intermediate, and commissural NTS subnuclei and the core of the area postrema, projects to restricted terminal zones in the inner portion of the external lateral PB, the central and dorsal lateral PB subnuclei, and the "waist" area. The dorsomedial NTS subnucleus and the rim of the area postrema specifically innervate the outer portion of the external lateral PB subnucleus. In addition, the medial NTS innervates the caudal lateral part of the external medial PB subnucleus. The respiratory part of the NTS, comprising the ventrolateral, intermediate, and caudal commissural subnuclei, is reciprocally connected with the K?lliker-Fuse nucleus, and with the far lateral parts of the dorsal and central lateral PB subnuclei. There is also a patchy projection to the caudal lateral part of the external medial PB subnucleus from the ventrolateral NTS. The rostral, gustatory part of the NTS projects mainly to the caudal medial parts of the PB complex, including the "waist" area, as well as more rostrally to parts of the medial, external medial, ventral, and central lateral PB subnuclei. The connections of different portions of the medullary reticular formation with the PB complex reflect the same patterns of organization, but are reciprocal. The periambiguus region is reciprocally connected with the same PB subnuclei as the ventrolateral NTS; the rostral ventrolateral reticular nucleus with the same PB subnuclei as both the ventrolateral (respiratory) and medial (general visceral) NTS; and the parvicellular reticular area, adjacent to the rostral NTS, with parts of the central and ventral lateral and the medial PB subnuclei that also receive rostral (gustatory) NTS input. In addition, the rostral ventrolateral reticular nucleus and the parvicellular reticular formation have more extensive connections with parts of the rostral PB and the subjacent reticular formation that receive little if any NTS input. The PB contains a series of topographically complex terminal domains reflecting the functional organization of its afferent sources in the NTS and medullary reticular formation.     

Intramedullary connections of the rostral nucleus of the solitary tract in the hamster

The rostral nucleus of the solitary tract (NST) figures prominently in the gustatory system, giving rise to ascending taste pathways that are well documented. Less is known of the local connections of the rostral NST with sites in the medulla. This study defines the intramedullary connections of the rostral NST in the hamster. Small iontophoretic injections of horseradish peroxidase (HRP), confined to the rostral NST, resulted in Golgi-like filling of axons that exited the NST or that interconnected cytoarchitectonic subdivisions within the NST complex. The NST efferent axons terminated sparsely in the trigeminal, facial and hypoglossal motor nuclei, but axons and endings were heavily distributed in the parvicellular reticular formation ventral to the NST. HRP injections centered in this part of the reticular formation resulted in heavy projections to the orofacial motor nuclei. Intranuclear connections, labelled after NST injections, linked NST subdivisions that receive primary afferent taste inputs to subdivisions involved in (1) projections to the preoromotor reticular formation, (2) projections to swallowing motor neurons, (3) activation of preganglionic parasympathetic neurons, and (4) general viscerosensation. In general, the connections defined in the present study provide anatomical details about the substrate for gustatory-motor and gustatory-visceral interactions.     

Substance P-containing trigeminal sensory neurons project to the nucleus of the solitary tract

Intense substance P-like immunoreactivity (SPLI) was identified in fiber bundles coursing between the spinal nucleus of the trigeminal nerve and the ventrolateral nucleus of the solitary tract at the level of the area postrema. These bundles were apparent only when tissue was stained for substance P immunoreactivity and were not visible in preparations treated with antisera to somatostatin or neurotensin. Following unilateral section of the trigeminal nerve, the SPLI-containing fiber bundles were absent ipsilateral to the nerve section. The fibers were absent bilaterally in rats which were previously injected with capsaicin. Unilateral removal of the nodose ganglion did not diminish the intensity or apparent number of SPLI fibers. These data indicate the presence of a trigeminosolitary projection which is composed of primary trigeminal sensory neurons containing substance P. The results provide an anatomical route by which substance P of trigeminal origin may modulate vagal or glossopharyngeal sensory information.     

Central afferents to the nucleus of the solitary tract in rats and mice

The nucleus of the solitary tract (NTS) regulates life-sustaining functions ranging from appetite and digestion to heart rate and breathing. It is also the brain's primary sensory nucleus for visceral sensations relevant to symptoms in medical and psychiatric disorders. To better understand which neurons may exert top-down control over the NTS, here we provide a brain-wide map of all neurons that project axons directly to the caudal, viscerosensory NTS, focusing on a medial subregion with aldosterone-sensitive HSD2 neurons. Injecting an axonal tracer (cholera toxin b) into the NTS produces a similar pattern of retrograde labeling in rats and mice. The paraventricular hypothalamic nucleus (PVH), lateral hypothalamic area, and central nucleus of the amygdala (CeA) contain the densest concentrations of NTS-projecting neurons. PVH afferents are glutamatergic (express Slc17a6/Vglut2) and are distinct from neuroendocrine PVH neurons. CeA afferents are GABAergic (express Slc32a1/Vgat) and are distributed largely in the medial CeA subdivision. Other retrogradely labeled neurons are located in a variety of brain regions, including the cerebral cortex (insular and infralimbic areas), bed nucleus of the stria terminalis, periaqueductal gray, Barrington's nucleus, Kölliker-Fuse nucleus, hindbrain reticular formation, and rostral NTS. Similar patterns of retrograde labeling result from tracer injections into different NTS subdivisions, with dual retrograde tracing revealing that many afferent neurons project axon collaterals to both the lateral and medial NTS subdivisions. This information provides a roadmap for studying descending axonal projections that may influence visceromotor systems and visceral “mind–body” symptoms.     

Localization of parabrachial nucleus neurons projecting to the thalamus or the amygdala in the cat using horseradish peroxidase.

Topographical localization of parabrachial nucleus (PBN) neurons projecting directly to the thalamus or the amygdala was examined in the cat by the horseradish peroxidase (HRP) method. After HRP injection in the central nucleus of the amygdala, PBN neurons labeled with the enzyme were seen ipsilaterally in the ventral portion of the lateral PBN as well as in the medial PBN. When the HRP injections were centered on the parvocellular portion of the posteromedial ventral nucleus of the thalamus (VPMpc), HRP-labeled neurons were observed ipsilaterally in the dorsal portion of the lateral PBN as well as in the medial PBN. Within the medial PBN, the distribution of neurons projecting to the amygdala overlapped that of neurons projecting to VPMpc; the cell bodies of the former neurons, however, tended to be more elongated than the latter, and the mean of the average soma diameters of the former was significantly larger than the latter. On the other hand, in the lateral PBN no significant differences were noted between the means of the average soma diameters of neurons projecting to VPMpc and those projecting to the amygdala. The PBN neurons in the cat were presumed to transmit gustatory and general visceral information ipsilaterally to the thalamic taste region and the limbic areas in the basal forebrain.     

Role of lateral parabrachial nucleus in the inhibition of water intake produced by right atrial stretch

Rats with either bilateral electrolytic or sham lesions of the ventrolateral portion of the lateral parabrachial nucleus (VLLPBN) were implanted with latex balloons that lay at the right superior vena cava/atrial junction (RSVC/AJ). Water intake in response to isoproterenol was measured both with and without inflation of the balloon. Water intake of the sham-lesioned rats was significantly depressed by balloon inflation during the first hour of the experiment. In contrast, water intake in the VLLPBN-lesioned rats was unaffected by balloon inflation. These results suggest that the VLLPBN is involved in the processing of afferent input from stretch-activated RSVC/AJ receptors.     

Conditioned place preference but not rewarding self-stimulation after electrical activation of the external lateral parabrachial nucleus

The objective of this experiment was to examine the rewarding effect of electrical stimulation of the external lateral parabrachial nucleus (LPBe) and of the lateral hypothalamus (LH) in concurrent Conditioned Place Preference (cCPP) and Brain Self-Stimulation Rewarding tasks. As expected, LH-stimulated animals readily learned cCPP tasks and developed self-stimulation behaviours following the rate–frequency procedure. As previously demonstrated, stimulation of the parabrachial complex generated rewarding or aversive behaviours in cCPP procedures. However, stimulation of this subnucleus induced consistent cCPP behaviours but not brain self-stimulation in rewarding LPBe animals. These results are analysed in the context of the different natural and artificial rewarding effects found in the LPBe nucleus.     

The pontine parabrachial region mediates some of the descending inhibitory effects of stimulating the anterior pretectal nucleus

Electrical stimulation of the anterior pretectal nucleus (APtN) elicits antinociception by inhibiting the responses of spinal multireceptive neurones to noxious stimuli. This descending inhibition is mediated, in part, by activating cells in the ventrolateral medulla. Neuronal tract tracing has previously shown that the APtN also projects directly to the pontine parabrachial region (PPR). The PPR, investigated by Katayama et al. (Brain Res., 296 (1984) 263-283), corresponds to the cholinergic cell group Ch5 of Mesulam et al. (Neuroscience, 10 (1983) 1185-1201). In this study, the pathway from APtN to PPR was investigated using urethane anaesthetised rats. Electrical stimulation (single square wave 0.2 ms pulses, 1-10 V, 5 Hz) of the APtN potently excites 40% of the cells recorded in the PPR. In the reverse experiment, stimulation of the PPR at the same parameters excited 36% of the cells recorded in the APtN. The contribution of this pathway to the spinal inhibitory effects of APtN stimulation was then examined. Unanaesthetised animals received electrical stimulation to the APtN (35 microA r.m.s., 15 s) and the increase in tail-flick latencies was measured. Bilateral electrolytic lesions of the PPR caused a 67% reduction of the antinociceptive effect of APtN stimulation. In urethane anaesthetised rats, microinjection of tetracaine into the PPR blocked the inhibition of multireceptive dorsal horn neurones caused by APtN stimulation (20 s train of 50 microA square wave 0.1 ms pulses, 100 Hz). In conclusion, these experiments strongly sugget that the PPR may be an important part of a descending antinociceptive pathway originating in the APtN.