Distribution and hypothalamic projection of tyrosine-hydroxylase containing neurons of the nucleus of the solitary tract in the pigeon.

The avian nucleus of the solitary tract has an extensive subnuclear organization. Several subnuclear cell groups can be distinguished on the basis of cytoarchitectonic criteria. In general, the subnuclei of the medial division of the nucleus of the solitary tract receive gastrointestinal afferents, whereas the subnuclei of the lateral division of the nucleus of the solitary tract receive cardiopulmonary afferents. Forebrain afferents to the nucleus of the solitary tract are segregated to medial and lateral subnuclei, which are located at the periphery of the nucleus. These peripheral subnuclei of the nucleus of the solitary tract are also the source of ascending axonal projections to the forebrain. In this study, the tyrosine hydroxylase (initial enzyme for catecholamine synthesis) content of the anteromedial hypothalamic projecting neurons of the nucleus of the solitary tract is determined by use of a combined retrograde fluorescent dye-immunofluorescence method. Fast Blue implanted into the anteromedial hypothalamus (in the region of the nucleus periventricularis magnocellularis) resulted in the retrograde labeling of neurons in the caudal two-thirds of the nucleus of the solitary tract. At levels rostral to the obex, dye-labeled cells were mostly observed in the dorsally located subnuclei medialis superficialis pars posterior and lateralis dorsalis pars posterior and in the ventrally located subnucleus medialis ventralis pars posterior. More centrally located subnuclei contained few labeled cells, if any. For example, subnucleus medialis intermedius pars posterior only had a few retrogradely labeled cells, whereas the centrally located subnucleus medialis dorsalis pars posterior was almost devoid of labeled cells. At levels caudal to the obex, many retrogradely labeled neurons of the nucleus of the solitary tract were observed. Neurons immunoreactively labeled for tyrosine hydroxylase were mostly found within subnuclei, which contain anteromedial hypothalamic projection neurons. In subnuclei medialis superficialis pars posterior and lateralis dorsalis pars posterior, 87% of the retrogradely dye-labeled cells were also immunoreactively labeled, whereas in the caudal nucleus of the solitary tract (at levels caudal to the obex), 68% of the retrogradely labeled cells were immunoreactively labeled. Not all tyrosine hydroxylase containing cells had projections to the implantation site in the anteromedial hypothalamus since only 40% of the immunoreactive cells in the caudal nucleus of the solitary tract and 59% of the immunoreactive cells in the subnucleus medialis superficialis pars posterior were retrogradely labeled with Fast Blue.(ABSTRACT TRUNCATED AT 400 WORDS)     

Distribution,parabrachial region projection,and coexistence of neuropeptide and catecholamine cells of the nucleus of the solitary tract in the pigeon

The chemical nature of the cells of the nucleus of the solitary tract (NTS) that project to the parabrachial nucleus (PB) was investigated in the pigeon by the use of fluorescent bead retrograde tracer and immunofluorescence for the detection of substance P (SP), leucineenkephalin (LENK), cholecystokinin (CCK), neurotensin (NT), somatostatin (SS), and tyrosine hydroxylase (TH). Cells immunoreactive for CCK were located in subnuclei lateralis dorsalis pars anterior (LDa) and medialis superficialis pars posterior, and caudal NTS (cNTS); 22–26.5% of these cells were double-labeled bilaterally. Immunoreactive SP cells were found in ventral NTS subnuclei; 24–25% of these cells were double-labeled bilaterally. Cells immunoreactive for LENK and NT were concentrated in the anterior NTS; 5.5–7.5% of the LENK cells were double-labeled bilaterally, while 11% (ipsilateral) and 21% (contralateral) of the NT immunoreactive cells were double-labeled. Many SS immunoreactive cells were found in peripherally located subnuclei; 5.5–6.5% of these cells were double-labeled bilaterally. Catecholamine cells were distributed in LDa, peripheral subnuclei, and cNTS; 23% of these cells were double-labeled ipsilaterally and 8.5% contralaterally. A two-color double-labeling immunofluorescence technique revealed many cells immunoreactive for both NT and LENK, only a rare cell immunoreactive for both SS and SP, and no cells immunoreactive for both TH and SP. Cells immunoreactive for SP, CCK, NT, and TH are major contributors to NTS projections to PB. The confinement of these substances to specific NTS subnuclei, which receive visceral sensory information from specific organs, may contribute to the chemical encoding of ascending visceral information. 1993 Wiley-Liss, Inc.     

Distribution of bombesin-like immunoreactivity in the nucleus of the solitary tract and dorsal motor nucleus of the rat and human: Colocalization with tyrosine hydroxylase

Bombesin is a peptide neurotransmitter/neuromodulator with important autonomic and behavioral effects that are mediated, at least in part, by bombesin-containing neurons and nerve terminals in the nucleus of the solitary tract (NTS) and the dorsal motor nucleus of the vagus (DMV). The distribution of bombesin-like immunoreactive nerve terminals/fibers and cell bodies in relation to a viscerotopically relevant subnuclear map of this region was studied by using an immunoperoxidase technique. In the rat, bombesin fiber/terminal staining was heavy in an area that included the medial subnucleus of the NTS and the DMV over their full rostral-caudal extent. Distinctly void of staining were the gelatinous, central, and rostral commissural subnuclei and the periventricular area of the NTS, regions to which gastric, esophageal, cecal, and colonic primary afferents preferentially project. The caudal commissural and dorsal subnuclei had light bombesin fiber/terminal staining, as did the intermediate, interstitial, ventral, and ventrolateral subnuclei. With colchicine pretreatment, numerous cell bodies were stained in the medial and dorsal subnuclei, with fewer neurons in the caudal commissural, intermediate, interstitial, ventral, and ventrolateral subnuclei. Bombesin-like immunoreactive neurons were found in numerous other areas of the brain, including the ventrolateral medulla, the parabrachial nucleus, and the medial geniculate body. In the human NTS/DMV complex, the distribution of bombesin fiber/terminal staining was very similar to the rat. In addition, occasional bombesin-like immunoreactive neurons were labeled in a number of subnuclei, with clusters of neurons labeled in the dorsal and ventrolateral subnuclei. Double immunofluorescence studies in rat demonstrated that bombesin colocalizes with tyrosine hydroxylase in neurons in the dorsal subnucleus of the NTS. Bombesin does not colocalize with tyrosine hydroxylase in any other location in the brain. In conclusion, the distribution of bombesin in the NTS adheres to a viscerotopically relevant map. This is the anatomical substrate for the effects of bombesin on gastrointestinal function and satiety and its likely role in concluding a meal. The anatomic similarities between human and rat suggest that bombesin has similar functions in the visceral neuraxis of these two species. Bombesin coexists with catecholamines in neurons in the dorsal subnucleus, which likely mediate, in part, the cardiovascular effects of bombesin. © 1996 Wiley-Liss, Inc.     

Cholecystokinin-, galanin-, and corticotropin-releasing factor-like immunoreactive projections from the nucleus of the solitary tract to the parabrachial nucleus in the rat.

The parabrachial nucleus (PB) is the main relay for ascending visceral afferent information from the nucleus of the solitary tract (NTS) to the forebrain. We examined the chemical organization of solitary-parabrachial afferents by using combined retrograde transport of fluorescent tracers and immunohistochemistry for galanin (GAL), cholecystokinin (CCK), and corticotropin-releasing factor (CRF). Each peptide demonstrated a unique pattern of immunoreactive staining. GAL-like immunoreactive (-ir) fibers were most prominent in the "waist" area, the inner portion of external lateral PB, and the central and dorsal lateral PB subnuclei. Additional GAL-ir innervation was seen in the medial and external medial PB subnuclei. GAL-ir perikarya were observed mainly rostrally in the dorsal lateral, superior lateral, and extreme lateral PB. CCK-ir fibers and terminals were most prominent in the outer portion of the external lateral PB; some weaker labeling was also present in the central lateral PB. CCK-ir cell bodies were almost exclusively confined to the superior lateral PB and the "waist" area, although a few cells were seen in the K?lliker-Fuse nucleus. The distribution of CRF-ir terminal fibers in general resembled that of GAL, but showed considerably less terminal labeling in the lateral parts of the dorsal and central lateral PB, and the external medial and K?lliker-Fuse subnuclei. The CRF-ir cells were most numerous in the dorsal lateral PB and the outer portion of the external lateral PB; rostrally, scattered CRF-ir neurons were seen mainly in the central lateral PB. After injecting the fluorescent tracer Fast Blue into the PB, the distribution of double-labeled neurons in the NTS was mapped. GAL-ir cells were mainly located in the medial NTS subnucleus; 34% of GAL-ir cells were double-labeled ipsilaterally and 7% contralaterally. Conversely, 17% of the retrogradely labeled cells ipsilaterally and 16% contralaterally were GAL-ir. CCK-ir neurons were most numerous in the dorsomedial subnucleus of the NTS and the outer rim of the area postrema. Of the CCK-ir cells, 68% in the ipsilateral and 10% in the contralateral NTS were double-labeled, whereas 15% and 10%, respectively, of retrogradely labeled cells were CCK-ir. In the area postrema, 36% of the CCK-ir cells and 9% of the Fast Blue cells were double-labeled. CRF-ir neurons were more widely distributed in the medial, dorsomedial, and ventrolateral NTS subnuclei, but double-labeled cells were mainly seen in the medial NTS.(ABSTRACT TRUNCATED AT 400 WORDS)     

Projections of the lateral hypothalamus and bed nucleus of the stria terminalis to the dorsal vagal complex in the pigeon

The dorsal vagal complex is composed of the nucleus tractus solitarii (Nts) and the dorsal motor nucleus of the vagus (DMN X). In the pigeon, these nuclei are composed of cytoarchitectonically well-defined subnuclear groups, which have connections that are partially segregated to specific organs (Katz and Karten: J. Comp. Neurol. 218:42-73, '83b, J. Comp. Neurol. 242:397-414, '85). The present study sought to determine whether forebrain afferents to Nts-DMN X are differentially distributed to specific subnuclei and thereby modulate the functions of specific organs. Forebrain afferents to the dorsal vagal complex were determined by retrograde tracing techniques. Labeled perikarya were found in the bed nucleus of the stria terminalis (BNST), ventral paleostriatum, and stratum cellulare externum (SCE) of the lateral hypothalamus, and in the medial hypothalamus, nucleus periventricularis magnocellularis (PVM), which is the avian homologue to a portion of the mammalian paraventricular nucleus. The pattern of axonal distribution to Nts-DMN X subnuclei from the BNST-ventral paleostriatum and SCE were investigated by anterograde tracing techniques. These experiments revealed axonal projections distributed to specific Nts-DMN X subnuclei. However, there is a high degree of overlap of the axonal projections to Nts-DMN X subnuclei from BNST-ventral paleostriatum and SCE, as well as from PVM (Berk and Finkelstein: J. Comp. Neurol. 220:127-136, '83). Labeled fibers from BNST-ventral paleostriatum and SCE project heavily to Nts subnuclei medialis superficialis, lateralis dorsalis, and medialis ventralis and to DMN X subnucleus ventralis parvicellularis. Fewer labeled fibers were found in Nts subnucleus medialis intermedius and extremely sparse labeling was found in Nts subnucleus medialis dorsalis. The Nts and DMN X subnuclei that receive forebrain projections also have peripheral connections with the aortic nerve, crop, esophagus, glandular stomach, and caudal abdominal organs. Thus, the forebrain could modulate the functions of these segments of the cardiovascular and digestive systems.     

The carotid body connections: a WGA-HRP study in the cat

Previous neuroanatomical studies described the central representation of the carotid sinus nerve, but did not differentiate the projections of the baroreceptors from the chemoreceptors present in the carotid bifurcation. In this research we investigated the individual territories occupied by the primary afferents from the carotid body in the brainstem of the cat. We also studied the distribution of afferent and efferent neurons to the carotid body. We injected into the carotid body lectin coupled to horseradish peroxidase. We found labeled axons only in the nucleus of the tractus solitarius; in particular, we found strong projections to the following ipsilateral subnuclei: dorsal, interstitial, and medial part of the commissural subnucleus. Moderate labeling was found in the ipsilateral medial and intermediate subnuclei and in the contralateral dorsal subnucleus and the medial region of the commissural subnucleus. We found a mean of 256 +/- 79 (S.E.M.) labeled afferent ganglion cells in the petrosal ganglia, and no evidence of efferent neurons in the brainstem that innervate the carotid body; conversely, about 4000 efferent neurons of the superior cervical ganglion send terminals to the ipsilateral carotid body.     

Subnuclear organization of the human caudal nucleus of the solitary tract

The caudal human nucleus of the solitary tract (NTS) is composed of 10 subnuclei. The commissural subnucleus spans the midline below the obex, merging rostrally into the medial subnucleus. The other subnuclei of the NTS are best seen just above the obex. The ventrolateral subnucleus contains large, darkly staining neurons. The interstitial subnucleus consists of neurons lying in groups intermingled with the fibers of the tract. The lateral subnucleus is small at caudal levels, merging with the interstitial subnucleus more rostrally. The dorsal subnucleus contains large melanotic neurons and encircles the substantia gelatinosus, a round, cell-poor subnucleus. The ventromedial subnucleus curls around the medial and ventral edge of the tract. The intermediate subnucleus, laying ventrolateral to the dorsal motor nucleus of the vagus, also contains melanotic neurons. The subpostremal subnucleus separates the area postrema from the NTS proper. The medial subnucleus is the largest subnucleus in the caudal NTS, containing medium-sized fusiform neurons. Adoption of a uniform cytoarchitectural map of the caudal NTS will permit more accurate comparisons between human and nonhuman studies.     

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.     

Autoradiographic localization of thyrotropin-releasing hormone and substance P receptors in the rat dorsal vagal complex

We utilized quantitative autoradiography to localize receptors for thyrotropin-releasing hormone (TRH) and substance P in individual subnuclei of the rat nucleus tractus solitarii (NTS) and the dorsal vagal complex. Within the NTS, TRH receptor concentrations were highest within the gelatinosus and centralis subnuclei and the medial subnucleus rostral to the area postrema, moderate within the intermediate subnucleus and the medial subnucleus adjacent to the area postrema, and low within the ventrolateral and commissural subnuclei and the medial subnucleus caudal to the area postrema. In contrast, substance P receptor concentrations were high throughout the medial subnucleus, moderate in all other subnuclei medial to the tractus solitarius, and relatively low in subnuclei lateral to the tractus solitarius. The dorsal motor nucleus of the vagus contained high concentrations of both TRH and substance P receptors, whereas we observed low TRH and moderate substance P receptors in the area postrema. High TRH and moderate substance P receptors were observed in the adjacent hypoglossal nucleus. In addition, we compared the concentrations of TRH receptors between chloroform-defatted and nondefatted tissue sections, and noted little effect of white matter tritium quench upon the observed TRH receptor concentrations. These results suggest that neurotransmitter receptors within the rat dorsal vagal complex are organized in a manner consistent with previous cytoarchitectural and hodological partitioning of the NTS and that the distribution of an individual neurotransmitter receptor in the NTS may correspond to the role of that transmitter in modulating autonomic function.     

Observations on the afferent and efferent organization of the vagus nerve and the innervation of the stomach in the squirrel monkey

Horseradish peroxidase was injected into the cervical vagus nerve or stomach wall of adult squirrel monkeys. Following cervical vagus nerve injections, labelled afferent fibres were present in the tractus solitarius and labelled fibres and terminals were present in medial and lateral parts of the nucleus of the tractus solitarius (NTS) ipsilaterally. Afferent labelling was also seen in the ipsilateral commissural nucleus and in the area postrema. Labelling was present contralaterally in caudal levels of the medial parts of the NTS, in the commissural nucleus, and in the area postrema. Afferent projections to the ipsilateral pars interpolaris of the spinal trigeminal nucleus and to the substantia gelatinosa of the C1 segment of the spinal cord were also labelled. Following injections of HRP into the anterior and posterior stomach walls, the tractus solitarius was labelled bilaterally. Afferent labelling was concentrated bilaterally in the dorsal parts of the medial division of the NTS, i.e., in the subnucleus gelatinosus, and in the commissural nucleus. The regions of NTS immediately adjacent to the tractus solitarius were largely unlabelled. Injections of HRP into the cervical vagus nerve resulted in heavy retrograde labelling of neurons in the ipsilateral dorsal nucleus of the vagus (DMX) and in the nucleus ambiguus (NA). In addition a few neurones were labelled in the intermediate zone between these two nuclei. Retrogradely labelled neurons were also present in the nucleus dorsomedialis in the rostral cervical spinal cord and in the spinal nucleus of the accessory nerve. Injections of HRP into the left cricothyroid muscle in two cases resulted in heavy retrograde labelling of large neurons in the left NA. Following stomach wall injections of HRP retrograde labelling of neurons was seen throughout the rostrocaudal and mediolateral extent of the DMX; there was no apparent topographical organization of the projection. In these cases, a group of labelled smaller neurons was found lying ventrolateral to the main part of the NA through its rostral levels. This study in a primate indicates that a large vagal afferent projection originates in the stomach wall and terminates primarily in the subnucleus gelatinosus of the NTS and in the commissural nucleus with a distribution similar to that described previously in studies in several subprimate mammalian species. The present results and those of other studies suggest some degree of segregation of visceral input within different subnuclei of the NTS.(ABSTRACT TRUNCATED AT 400 WORDS)     

Distribution and neurochemical phenotypes of caudal medullary neurons activated to express cFos following peripheral administration of cholecystokinin

Immunocytochemical localization of the protein product of the proto-oncogene C-fos allows anatomical identification of physiologically activated neurons. The present study examined the subnuclear distribution of cFos protein in the rat caudal medulla following peripheral administration of cholecystokinin octapeptide, which reduces feeding and gastric motility by a vagally mediated mechanism. To begin phenotypic characterization of neurons activated to express cFos following cholecystokinin treatment, double-labeling techniques were used to identify vagal motor neurons and neurons immunoreactive for tyrosine hydroxylase, neuropeptide Y, and neurotensin. Activated cells were most prevalent in the subnucleus medialis of the nucleus of the solitary tract, less prevalent in the subnucleus commissuralis, and virtually absent in the subnuclei centralis and gelatinosus. Many activated cells occupied the caudal area postrema; some of these were catecholaminergic. In contrast, activated cells were sparse within the medial rostral area postrema. Other activated cells occupied the dorso- and ventrolateral medulla and the midline raphe nuclei. Retrograde labeling of vagal motor neurons confirmed that very few were activated. Those that were activated occupied the caudal dorsal motor nucleus. In the dorsomedial medulla, 51% of catecholaminergic neurons and 39% of neurons positive for neuropeptide Y were activated, but no neurotensin-positive neurons were activated. In the ventrolateral medulla, 25% of catecholaminergic neurons and 27% of neuropeptide Y-positive neurons were activated. By characterizing the subnuclear distribution and chemical phenotypes of neurons activated by exogenous cholecystokinin, these data contribute to elucidation of the neural circuits mediating the behavioral, physiological, and neuroendocrine effects produced by this peptide. © 1993 Wiley-Liss, Inc.     

Substance P innervation of neurons projecting to the paraventricular hypothalamic nucleus in the rat nucleus tractus solitarius

After injection of WGA-HRP-colloidal gold in the rat paraventricular nucleus (PVN), retrogradely labeled neurons were found mainly in the medial and commissural subnuclei of the nucleus tractus solitarius (NTS) around 0.5 mm caudal to the obex which is closely related to cardiovascular function. Electron microscopic immunohistochemistry in these areas demonstrated synaptic contacts between retrogradely labeled neurons and substance P-immunoreactive terminals. Innervation of NTS-PVN projection systems by substance P is suggested.     

Immunohistochemical localisation of a gastrin-releasing peptide-like material in area postrema, nucleus of the solitary tract and vagal motor nucleus in the brainstem of rat

The presence of an endogenous gastrin-releasing peptide (GRP)-like peptide in the hindbrain of rat was demonstrated immunohistochemically using antisera directed against the N-terminus and C-terminus of GRP. N-terminal and C-terminal-like immunoreactive material were distributed throughout the nucleus of the solitary tract (NTS), dorsal motor nucleus of the vagus (DMV) and tractus solitarius (TS), as well as in areas postrema (AP) and substantia gelatinosa separating AP from NTS. Positive immunostaining was localised to a dense network of nerve fibres which project longitudinally along the neuraxis. Immunolabelled cell bodies were observed rostral to the obex, principally in the mediolateral subnucleus of NTS. These immunopositive neurones project their axons caudally and longitudinally towards the commissural subnucleus of the NTS. Immunolabelled cell bodies also were found in AP; they projected their axons caudally and ventrally towards NTS. Positive immunostaining was blocked by pre-adsorbing antisera with either GRP (1 nmol/ml) or bombesin (3 nmol/ml), but was unaffected by substance P (30 nmol/ml) and spared by capsaicin pretreatments which deplete sensory nerves of their peptide content. The results indicate that NTS neurons containing a GRP-like peptide connect the rostral and caudal regions of the dorsal vagal complex by way of longitudinal nerve tracts descending NTS and TS. Some neurons in AP also contain a GRP-like peptide and appear to connect with the dorsal vagal complex.     

Distribution of neuropeptide immunoreactive nerve terminals within the subnuclei of the nucleus of the tractus solitarius of the rat

The location of substance P, enkephalin and somatostatin (SRIF), and neurophysin II immunoreactive nerve terminals and preterminal processes in the caudal part of the nucleus of the tractus solitarius (nTS) was examined by the indirect immunofluorescence method for immunocytochemistry combined with cytoarchitectural identification of nuclear subgroups in the same tissue. In 22 Sprague-Dawley rats we examined 14-micrometers-thick serial sections of the dorsal medulla at levels from 1 mm caudal to 2 mm rostral to the obex. These sections were incubated with substance P, enkephalin, somatostatin, and neurophysin II antisera. All four peptides were examined in each case and five typical levels (two caudal and three rostral to the obex) were selected for comparison of terminal distribution between peptides. All sections were photographed under the fluorescence microscope and then counter-stained with cresyl violet. This method of analysis revealed distinct patterns of neuropeptide immunoreactivity in the subnuclei of the nTS that varied according to the level of the section. The nTS is responsible for integrating respiratory, cardiovascular (baroreceptor and cardiac), and gastrointestinal functions. The ventrolateral subnucleus (Vl)nTS, ventral subnucleus (v)nTS, interstitial subnucleus (ni)nTS, and intermediate subnucleus (nI)nTS are the major respiratory subnuclei with vlnTS and vnTS prominently associated with pulmonary afferents, ni associated with laryngeal afferents, and nI with tracheal afferents. The vlnTS, vnTS, and ni showed a moderate density of somatostatin-positive nerve terminals, scattered substance P and enkephalin immunoreactivity, and no neurophysin II-positive terminals. The nI showed moderate density of substance P immunoreactive nerve terminals. The subnuclei of the nTS receiving baroreceptor and chemoreceptor afferents--dorsolateral and dorsal (dl and d) subnuclei of nTS--showed scattered substance P immunoreactive nerve terminals. The commissural nucleus of nTS (ncom), which receives most of the cardiac afferents, showed a moderate density of enkephalin-positive immunoreactive nerve terminals. The medial subnucleus (m)nTS at levels rostral to the obex, the primary site for the termination of gastrointestinal afferents, showed substance P immunoreactivity in moderate amounts and weak immunoreactivity for all the other neuropeptides. An important result of these experiments was the observation that regions of the medulla adjacent to the nTS, i.e., the ventral parasolitarius region (vPSR), dorsal (d)PSR, and the periventricular region (PVR) showed the densest amounts of immunoreactive nerve terminals.(ABSTRACT TRUNCATED AT 400 WORDS)     

The carotid sinus connections: a WGA-HRP study in the cat

The neural connections of the carotid sinus were studied by administration of horseradish peroxidase or a lectin conjugate to the adventitia of the carotid sinus of cats. The carotid sinus afferents project exclusively to the nucleus of the tractus solitarius (NTS). Rostral to the obex the projection is mainly ipsilateral with a strong contralateral component caudal to the obex. The carotid sinus projects to several NTS territories that do not receive afferents from the carotid body chemoreceptors; they are: the dorsolateral, the lateral extension of the commissural, the caudal intermediate, the ventrolateral and the gelatinosus subnuclei. In addition the carotid sinus central representation includes territories occupied also by carotid body terminals: dorsal, interstitial, rostral intermediate, medial and the medial part of commissural subnuclei (see previous paper). Labeled cell bodies were found in the petrosal (216 +/- 37, mean +/- S.E.M.) nodose (825 +/- 434) and superior cervical ganglia (3583 +/- 1227) demonstrating the sympathetic efferent innervation of the carotid sinus and a dual sensory innervation via both the glossopharyngeal and vagus nerves.     

Phenylethanolamine N-methyltransferase-immunoreactive neurons in the medulla oblongata of the pigeon (Columba livia) projecting to the hypothalamic paraventricular nucleus

The distribution and ascending projections to the hypothalamic paraventricular nucleus of phenylethanolamine N-methyltransferase (PNMT)-immunoreactive perikaria were studied in adult pigeons using a combination of retrograde transport of Fluorogold injected into the paraventricular nucleus, and double immunohistochemical procedures for PNMT, tyrosine hydroxylase and neuropeptide Y. PNMT-immunoreactive cell bodies were found in the subtrigeminal reticular nucleus of the ventrolateral medulla and in the nucleus of the solitary tract, mainly in the subnuclei: medialis superficialis, pars posterior, and medialis ventralis, pars posterior. PNMT-immunoreactive perikaria were also tyrosine hydroxylase immunoreactive, and are located within the rostral tyrosine hydroxylase immunoreactive cell groups of these areas. No perikaria double-labeled for neuropeptide Y and PNMT were found. Retrograde labeled cell bodies were observed in the subtrigeminal reticular nucleus and in the nucleus of the solitary tract. PNMT-immunoreactive retrogradely labeled cells were mainly observed in the subtrigeminal reticular nucleus. These data suggest the presence in the pigeon of medullary adrenergic cell groups partially comparable to mammalian C1 and C2 groups. Comparison of these results with data previously obtained in amphibians and reptiles suggests that the presence of a hypothalamically-projecting C1-like group might be a plesiomorphic medullary attribute in amniotes, whereas the variable presence of C2 and C3-like groups, as well as the content of NPY in the putative adrenergic perikaria, seem to be species-specific.     

Long descending projections of the hypothalamus in the pigeon, Columba livia

An autoradiographic analysis was performed on the descending projections of nucleus periventricularis magnocellularis (PVM) of the hypothalamus in the pigeon. A PVM-medullospinal pathway was observed coursing posteriorly through the lateral hypothalamus, ventrolateral midbrain tegmentum, and into the spinal lemniscus (ls) in the ventrolateral pons and medulla. In the pons, some fibers course dorsomedially from ls and terminate at the lateral border of the locus coeruleus. At medullary levels, fibers from ls sweep dorsomedially in the plexus of Horsley and project to certain regions of the nucleus of the solitary tract (NTS) and the dorsal motor nucleus of the vagus (NX). Specifically, PVM fibers project heavily into NTS subnuclei medialis superficialis, medialis ventralis, and lateralis (sulcalis) dorsalis as well as into the ventral parvocellular subnucleus of NX. Fibers in ls were traced caudally into the lateral funiculus as far as upper cervical levels of the spinal cord. Although autoradiographs of lower cervical or thoracic spinal cord sections were not available, PVM fibers do descend to thoracic spinal cord levels, as evidenced by the retrograde transport of horseradish peroxidase. In addition to the medullospinal pathway, the autoradiographs demonstrated PVM projections to septum, diencephalon, and midbrain. Labeled PVM fibers are found in the lateral septal nucleus, nucleus of the anterior pallial commisure, dorsomedial thalamic nucleus, dorsolateral anterior thalamic nucleus (pars ventralis), median eminence, medial and lateral hypothalamus, medial mammillary area, and nucleus intercollicularis and central gray of the midbrain. The projection of fibers to medullospinal regions and median eminence suggests that PVM is homologous to the mammalian paraventricular nucleus. These projections to specific subnuclei of NTS and NX denote hypothalamic control over certain autonomic functions.     

The dorsal vagal complex of the ferret: anatomical and immunohistochemical studies

To further the understanding of gastrointestinal function in this species, and in particular to advance our own work concerning central emetic pathways, the cytoarchitecture and the distribution of eight neurochemicals were studied in the ferret dorsal vagal complex (DVC; area postrema, nucleus of the solitary tract [nTS] and dorsal motor nucleus of the vagus). The cytoarchitectural features of this region in the ferret were similar to those seen in other species; however, the ferret possesses a particularly large and distinct subnucleus gelatinosus of the nTS. Dense calcitonin gene-related peptide-immunoreactivity was found in the gelatinous, interstitial and commissural subnuclei of the nTS, with lesser amounts in other regions of the DVC. Enkephalin-immunoreactivity of varying densities was found throughout the DVC. Moderate to dense galanin-immunoreactivity was observed throughout the DVC, with the exception of the subnucleus gelatinosus of the nTS, from which it was virtually absent. Dense neuropeptide Y-immunoreactivity was observed in the subnucleus gelatinosus and interstitial subnucleus, with moderate staining in other regions of the DVC. Neurotensin immunoreactivity was very sparse or absent. Immunoreactivity for serotonin was sparsely distributed throughout the DVC. Moderate somatostatin-immunoreactivity was observed over a large portion of the DVC, but was virtually absent from the gelatinosus and interstitial subnuclei. Substance P immunoreactivity was observed throughout the DVC and was particularly dense in the dorsal/dorsolateral subnucleus and the dorsal aspects of the medial and commissural subnuclei. In terms of its cytoarchitecture the DVC of the ferret is more similar to the cat than the rat, especially with regard to the area postrema and the subnucleus gelatinosus of the nTS. The distribution of neuroactive substances was largely similar to other species; however, differences were present particularly in patterns of immunoreactivity for enkephalin, serotonin, neuropeptide Y and somatostatin.     

Neuropeptide Y-like immunoreactivity in the brain stem respiratory nuclei of the cat

The distribution of fibres and cell bodies containing neuropeptide Y-like immunoreactivity in respiratory nuclei of the brain stem was studied using an indirect immunoperoxidase technique. In order to visualize immunoreactive perikarya, intraventricular or intratissue injections of colchicine were carried out. The richest cluster of immunoreactive perikarya was localized in the B?tzinger complex, whereas in the nuclei ambiguus, retroambiguus, parabrachialis medialis and K?lliker-Fuse area, a moderate density of cell bodies was observed. The ventrolateral subnucleus of the nucleus tractus solitarii had the lowest number of immunoreactive neurons. Neuropeptide Y-like immunoreactive fibres were abundant in the K?lliker-Fuse area, and scarce in the nuclei ambiguus, parabrachialis medialis and K?lliker-Fuse area. A moderate network of immunoreactive fibres was observed in the nuclei retroambiguus and the ventrolateral subnucleus of the nucleus tractus solitarii. The presence of neuropeptide Y-like immunoreactive areas suggests a role for this peptide in the control of respiratory mechanisms. Alternatively such a cluster of cell bodies and fibre networks might be also related with neighbouring cardiovascular control areas.     

Mapping of carotid baroreceptor subtype projections to the nucleus tractus solitarius using c-fos immunohistochemistry

This study has combined physiological pressure stimulation of carotid baroreceptors via a vascularly isolated carotid sinus and anodal block of baroreceptor afferent fibers in the carotid sinus nerve to examine the medullary projections of type I vs. type II (large A- vs. small A- and C-fiber afferent) baroreceptors. The control distribution of cells in the nucleus tractus solitarius expressing c-fos in response to physiological activation of carotid baroreceptors in the isolated sinus was compared to that during anodal block of large A-fibers in the carotid sinus nerve. Carotid baroreceptor stimulation primarily activated neurons in the ipsilateral commissural and medial subnuclei of the caudal nucleus tractus solitarius and the dorsolateral, dorsomedial and medial subnuclei in the intermediate and rostral levels of the nucleus tractus solitarius. Elimination of large A-fiber carotid baroreceptor afferents, during similar carotid baroreceptor stimulation resulted in a decrease in the nmber of cells expressing c-fos in the dorsomedial subnucleus of the rostral nucleus tractus solitarius. These data indicate that projections of larger A-fiber (type I) carotid baroreceptors are localized primarily to the rostral dorsomedial subnucleus, while those of smallr A- and C-fiber baroreceptors are more widely distributed to the commissural, medial and dorsal subnuclei of the nucleus tractus solitarius.