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.     

Projections of the nucleus of the tractus solitarius in the pigeon (Columba livia)

With the aid of autoradiographic and histochemical (WGA-HRP) tracing techniques, the projections of the nucleus of the tractus solitarius (nTS) in the pigeon have been delineated and related to the viscerotopic organization of the nucleus. As in mammals, nTS projects to both brainstem and forebrain structures. At medullary levels, projections were seen to nTS itself, to the dorsal motor nucleus of the vagus and to the subjacent and more ventral reticular formation. There is a substantial projection to the parabrachial nuclear complex with terminations in all its subnuclei and minor projections to locus coeruleus and several mesencephalic areas, including the ventral area of Tsai, the nucleus of the ascending brachium conjunctivum, and the compact portion of the tegmental pedunculopontine nucleus. At diencephalic levels, projections to the hypothalamus (magnocellular periventricular nucleus, stratum cellulare internum and externum) and dorsal thalamus were seen. Terminal fields within the basal telencephalon included the nucleus of the pallial commissure, the bed nucleus of the stria terminalis, and the nucleus accumbens. The organization of nTS projections in pigeons is correlated with the pattern of inputs to specific nTS subnuclei. Lateral tier subnuclei receiving cardiovascular and pulmonary inputs project upon the ventrolateral reticular formation and the ventrolateral parabrachial complex. Medial tier subnuclei receiving gustatory and gastrointestinal inputs project upon dorsal and medial parabrachial nuclei. Transparabrachial projections arise from nTS subnuclei receiving little or no primary input from the viscera.     

Avian somatosensory system: II. Ascending projections of the dorsal column and external cuneate nuclei in the pigeon

The ascending projections of the dorsal column and external cuneate nuclei (DCN/CuE) in the pigeon were investigated in anterograde tracing experiments by using autoradiography or wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). The results show that the majority of ascending projections decussate via internal arcuate fibers to form a contralateral medial lemniscus which ascends in a ventral position. In the brainstem, terminal fields were observed in the ventral lamella of the inferior olive (OI), the parabrachial nuclei (PB) of the dorsolateral pons, the intercollicular nucleus (ICo) of the midbrain, and the nucleus pretectalis diffusus (PD). In the diencephalon there were terminal fields in the strata cellulare externum and internum (SCE and SCI) of the caudal hypothalamus; in the intercalated (ICT), ventrolateral (VLT), and reticular nuclei of the ventral thalamus; in the nuclei principalis precommissuralis (PPC), spiriform medialis (SpM), and dorsolateralis posterior, pars caudalis (cDLP) of the caudal thalamus; and in the nuclei dorsalis intermedius ventralis anterior (DIVA), dorsolateralis posterior, pars rostralis (rDLP), dorsolateralis anterior (DLA), and dorsolateralis anterior, pars medialis (DLM) of the rostrodorsal thalamus. The origins of these projections within the DCN/CuE complex were verified in retrograde tracing experiments with WGA-HRP and were found to be partly differentiable with respect to their targets. The projections to DIVA, rDLP, DLA, DLM, cDLP, and SpM arise from all rostrocaudal levels of the DCN/CuE complex; those to ICo arise from caudomedial nuclear regions, while those to the hypothalamus and ventral thalamus arise from rostrolateral nuclear regions. Projections to PB arise from lamina I neurons of the dorsal horn of upper cervical spinal cord segments and from CuE. No evidence was found of a projection to the cerebellum. The distribution of the cells of origin of the medial lemniscus (ML) within the DCN/CuE complex was found to be largely coextensive with the areas of termination of primary spinal (Wild: J. Comp. Neurol. 240:377-395, '85) and some trigeminal (Dubbledam and Karten: J. Comp. Neurol. 180:661-678, '78) afferents. Furthermore, the areas of termination of the ML within the rostrodorsal and caudal thalamus are also either coextensive or closely associated with nuclei which provide a somatosensory projection to separate regions of the telencephalon (Wild: Brain Res. 412:205-223, '87). There are thus clear similarities in the overall pattern of somatosensory projections in the pigeon and in many mammalian species.     

Efferent connections of the lateral hypothalamic area of the rat: An autoradiographic investigation

The efferent projections of the lateral hypothalamic area (LHA) at mid-tuberal levels were examined with the autoradiographic tracing method. Connections were observed to widespread regions of the brain, from the telencephalon to the medulla. Ascending fibers course through LHA and the lateral preoptic area and lie lateral to the diagonal band of Broca. Fibers sweep dorsally into the lateral septal nucleus, cingulum bundle and medial cortex. Although sparse projections are found to the ventromedial hypothalamic nucleus, a prominent pathway courses to the dorsal and medial parvocellular subnuclei of the paraventricular nucleus. Labeled fibers in the stria medullaris project to the lateral habenular nucleus. The central nucleus of the amygdala is encapsulated by fibers from the stria terminalis and the ventral amygdalofugal pathway. The substantia innominate, nucleus paraventricularis of the thalamus, and bed nucleus of the stria terminalis also receive LHA fibers. Three descending pathways course to the brainstem: (1) periventricular system, (2) central tegmental tract (CTT), and (3) medial forebrain bundle (MFB). Periventricular fibers travel to the ventral and lateral parts of the midbrain central gray, dorsal raphe nucleus, and laterodorsal tegmental nucleus of the pens. Dorsally coursing fibers of CTT enter the central tegmental field and the lateral and medial parabrachial nuclei. The intermediate and deep layers of the superior colliculus receive some fibers. Fibers from CTT leave the parabranchial region by descending in the ventrolateral pontine and medullary reticular formation; some of these fibers sweep dorsomedially into the nucleus tractus solitarius, dorsal motor nucleus of the vagus, and nucleus commissuralis. From MFB, fibers descend into the ventral tegmental area and to the border of the median raphe and raphe magnus nuclei.     

Differential projections to the intralaminar and gustatory thalamus from the parabrachial area: a PHA-L study in the rat

The organization of projections from the parabrachial (PB) area to the ventral posterior parvicellular (VPpc) "gustatory" and intralaminar nuclei of the thalamus was studied in the rat by using microinjections of Phaseolus vulgaris leucoagglutinin (PHA-L), into subregions of the PB area. The present study is a follow-up of three former studies (Bernard et al. [1993] J. Comp. Neurol. 329:201-229; Aldén et al. [1994] J. Comp. Neurol. 341:289-314; Bester et al. [1997a] J. Comp. Neurol. 383:245-281) that examined PB projections onto the amygdala, the bed nucleus of the stria terminalis, and the hypothalamus. Our data showed that (1) the region centered in the internal lateral PB subnucleus projects densely with a bilateral and symmetric pattern to the caudal portion of the paracentral and, to a lesser extent, to the adjacent portion of the central and parafascicular medial thalamic nuclei; (2) the mesencephalic PB region centered in the ventral lateral subnucleus and scattered neurons in the subjacent brachium conjunctivum project primarily, although diffusely, to the central medial thalamic nucleus. The third region includes two subgroups: (3a) the medial subgroup, including the medial, the waist area, and the ventral lateral subnuclei of the pontine PB area, projects bilaterally but with a weak ipsilateral predominance to the VPpc, terminals bearing large varicosities. Additionally, a diffuse projection with small varicosities spreads in the area between the two VPpc nuclei and the central medial nucleus. (3b) The lateral subgroup, centered in the external medial subnucleus, projects with a contralateral predominance in the periphery of the VPpc nuclei, most terminals being located around the dorsomedial tip. It is suggested that the PB projections to the intralaminar nucleus could be involved in cortical limbic arousal processing in relation with nociceptive, (somatic, visceral, and intraoral) and gustatory aversive stimuli. The projection with large varicosities inside the VPpc could process gustatory discrimination.     

Efferent and collateral organization of paratrigeminal nucleus projections: An anterograde and retrograde fluorescent tracer study in the rat

The paratrigeminal nucleus (PTN) receives primary visceral afferent projections through cranial nerves IX and X and somatic afferent projections through cranial nerve V and dorsal roots as far caudally as C7. Pressure injections of the anterograde tracer tetramethylrhodamine dextran into the PTN in the rat resulted in bilateral labeling in the nucleus of the tractus solitarius, dorsal motor nucleus of the vagus nerve, and parabrachial nucleus. Anterograde labeling in the parabrachial nucleus was strongest in the external medial, external lateral, and ventral lateral subnuclei. Anterograde labeling was also found in the contralateral paratrigeminal nucleus, lamina I of the spinal trigeminal nucleus subnucleus caudalis, and ventroposteromedial nucleus of the thalamus. The collateral organization of PTN neurons was demonstrated by injecting different fluorescent retrograde tracers into the terminal fields of PTN projections as determined by the anterograde tracing experiments. Double-labeled neurons were found in the paratrigeminal nucleus following all combinations of injection sites. The most prominent PTN efferent projections and the most highly collateralized were to the nucleus of the tractus solitarius and parabrachial nucleus. The efferent and collateral connections of the paratrigeminal nucleus may provide a neuroanatomical substrate for integrating convergent visceral and somatic afferent information used to modulate autonomic function and behavior related to thermoregulation, nociception, and gustation. J. Comp. Neurol. 402:93–110, 1998. © 1998 Wiley-Liss, Inc.     

Organization of cortical, basal forebrain, and hypothalamic afferents to the parabrachial nucleus in the rat.

In a previous study (Herbert et al., J. Comp. Neurol. [1990];293:540-580), we demonstrated that the ascending afferent projections from the medulla to the parabrachial nucleus (PB) mark out functionally specific terminal domains within the PB. In this study, we examine the organization of the forebrain afferents to the PB. The PB was found to receive afferents from the infralimbic, the lateral prefrontal, and the insular cortical areas; the dorsomedial, the ventromedial, the median preoptic, and the paraventricular hypothalamic nuclei; the dorsal, the retrochiasmatic, and the lateral hypothalamic areas; the central nucleus of the amygdala; the substantia innominata; and the bed nucleus of the stria terminalis. In general, forebrain areas tend to innervate the same PB subnuclei from which they receive their input. Three major patterns of afferent termination were noted in the PB; these corresponded to the three primary sources of forebrain input to the PB: the cerebral cortex, the hypothalamus, and the basal forebrain. Hypothalamic afferents innervate predominantly rostral portions of the PB, particularly the central lateral and dorsal lateral subnuclei. The basal forebrain projection to the PB ends densely in the external lateral and waist subnuclei. Cortical afferents terminate most heavily in the caudal half of the PB, particularly in the ventral lateral and medial subnuclei. In addition, considerable topography organization was found within the individual projections. For example, tuberal lateral hypothalamic neurons project heavily to the central lateral subnucleus and lightly to the waist area; in contrast, caudal lateral hypothalamic neurons send a moderately heavy projection to both the central lateral and waist subnuclei. Our results show that the forebrain afferents of the PB are topographically organized. These topographical differences may provide a substrate for the diversity of visceral functions associated with the PB.     

Amygdalo-hypothalamic projections in the lizard Podarcis hispanica: A combined anterograde and retrograde tracing study

The cells of origin and terminal fields of the amygdalo-hypothalamic projections in the lizard Podarcis hispanica were determined by using the anterograde and retrograde transport of the tracers, biotinylated dextran amine and horseradish peroxidase. The resulting labeling indicated that there was a small projection to the preoptic hypothalamus, that arose from the vomeronasal amygdaloid nuclei (nucleus sphericus and nucleus of the accessory olfactory tract), and an important projection to the rest of the hypothalamus, that was formed by three components: medial, lateral, and ventral. The medial projection originated mainly in the dorsal amygdaloid division (posterior dorsal ventricular ridge and lateral amygdala) and also in the centromedial amygdaloid division (medial amygdala and bed nucleus of the stria terminalis). It coursed through the stria terminalis and reached mainly the retrochiasmatic area and the ventromedial hypothalamic nucleus. The lateral projection originated in the cortical amygdaloid division (ventral anterior and ventral posterior amygdala). It coursed via the lateral amygdalofugal tract and terminated in the lateral hypothalamic area and the lateral tuberomammillary area. The ventral projection originated in the centromedial amygdaloid division (in the striato-amygdaloid transition area), coursed through the ventral peduncle of the lateral forebrain bundle, and reached the lateral posterior hypothalamic nucleus, continuing caudally to the hindbrain. Such a pattern of the amygdalo-hypothalamic projections has not been described before, and its functional implications in the transfer of multisensory information to the hypothalamus are discussed. The possible homologies with the amygdalo-hypothalamic projections in mammals and other vertebrates are also considered. J. Comp. Neurol. 384:537–555, 1997. © 1997 Wiley-Liss, Inc.     

Neuropeptides and catecholamines in efferent projections of the nuclei of the solitary tract in the rat

This study focuses on the involvement of catecholamines and nine different peptides in efferents of the nucleus of the solitary tract to the central nucleus of the amygdala, the bed nucleus of the stria terminalis, and different parabrachial and hypothalamic nuclei in the rat. A double-labeling technique was used that combines a protein-gold complex as the retrograde tracer with immunohistochemistry. Catecholaminergic projection neurons were the most numerous type observed and projected mainly ipsilaterally to all targets studied. Most projections arose from areas overlying the dorsal motor nucleus, mainly the medial nucleus. Neurons synthesizing somatostatin, met-enkephalin-Arg-Gly-Leu, dynorphin B, neuropeptide Y, and neurotensin projected to all structures examined. Somatostatin and enkephalin immunoreactive projection cells were the most numerous. They were located in close proximity to each other, including all subnuclei immediately surrounding the solitary tract, bilaterally. Most dynorphin and neuropeptide Y immunoreactive projection cells were found rostral to that of enkephalinergic and somatostatinergic projections, and mainly in the ipsilateral medial nucleus. Neurotensinergic projections were sparse and from dorsal and dorsolateral nuclei. Substance P and cholecystokinin contribute to parabrachial afferents. The location of substance P immunoreactive projection cells closely resembled that of enkephalinergic and somatostatinergic projections. Projecting cholecystokinin immunoreactive cells were observed in dorsolateral nucleus. Bombesin immunoreactive cells in dorsal nucleus projected to either the parabrachial or hypothalamic nuclei. No vasoactive intestinal polypeptide-containing cells were detected. Thus, most catecholaminergic and neuropeptidergic efferents originated from different populations of cells. It is proposed that catecholaminergic neurons constitute the bulk of solitary efferents and that they may contribute to autonomic neurotransmission. Peptidergic neurons mainly form other subgroups of projections and may play a role in modulating the physiological state of the target nuclei.     

Efferent connections of the substantia innominata in the rat

The efferent connections of the substantia innominata (SI) were investigated employing the anterograde axonal transport of Phaseolus vulgaris leucoagglutinin (PHA-L) and the retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). The projections of the SI largely reciprocate the afferent connections described by Grove (J. Comp. Neurol. 277:315-346, '88) and thus further distinguish a dorsal and a ventral division in the SI. Efferents from both the dorsal and ventral divisions of the SI descend as far caudal as the ventral tegmental area, substantia nigra, and peripeduncular area, but projections to pontine and medullary structures appear to originate mainly from the dorsal SI. Within the amygdala and hypothalamus, which receive widespread innervation from the SI, the dorsal SI projects preferentially to the lateral part of the bed nucleus of the stria terminalis; the lateral, basolateral, and central nuclei of the amygdala; the lateral preoptic area; paraventricular nucleus of the hypothalamus; and certain parts of the lateral hypothalamus, prominently including the perifornical and caudolateral zones described previously. The ventral SI projects more heavily to the medial part of the bed nucleus of the stria terminalis; the anterior amygdaloid area; a ventromedial amygdaloid region that includes but is not limited to the medial nucleus; the lateral and medial preoptic areas; and the anterior hypothalamus. Modest projections reach the lateral hypothalamus, with at least a slight preference for the medial part of the region, and the ventromedial and arcuate hypothalamic nuclei. Both SI divisions appear to innervate the dorsomedial and posterior hypothalamus and the supramammillary region. In the thalamus, the subparafascicular, gustatory, and midline nuclei receive a light innervation from the SI, which projects more densely to the medial part of the mediodorsal nucleus and the reticular nucleus. Cortical efferents from at least the midrostrocaudal part of the SI are distributed primarily in piriform, infralimbic, prelimbic, anterior cingulate, granular and agranular insular, perirhinal, and entorhinal cortices as well as in the main and accessory olfactory bulbs. The cells of origin for many projections arising from the SI were identified as cholinergic or noncholinergic by combining the retrograde transport of WGA-HRP with histochemical and immunohistochemical procedures to demonstrate acetylcholinesterase activity or choline acetyltransferase immunoreactivity. Most of the descending efferents of the SI appear to arise primarily or exclusively from noncholinergic cells.(ABSTRACT TRUNCATED AT 400 WORDS)     

Projections of the dorsal and lateral terminal accessory optic nuclei and of the interstitial nucleus of the superior fasciculus (posterior fibers) in the rabbit and rat

The projections of the dorsal and lateral terminal accessory optic nuclei (DTN and LTN) and of the dorsal and ventral components of the interstitial nucleus of the superior fasciculus (posterior fibers; inSFp have been studied in the rabbit and rat by the method of retrograde axonal transport following injections of horseradish peroxidase into oculomotor-related brainstem nuclei. The projections of the ventral division of the inSFp have been further investigated in rabbits with the anterograde axonal transport of 3H-leucine. The data show that the projections of the DTN, LTN, and inSFp are remarkably similar in rabbit and rat. The DTN projects heavily to the ipsilateral medial terminal accessory optic nucleus (MTN), nucleus of the optic tract, and dorsal cap of the inferior olive. The DTN projects sparsely to the ipsilateral visual tegmental relay zone and to the contralateral superior and lateral vestibular nuclei. The LTN and dorsal component of the inSFp are found to share the same basic connections; both project heavily to the ipsilateral nucleus of the optic tract and visual tegmental relay zone and send a moderately sized projection to the ipsilateral MTN. However, while the dorsal component of the inSFp sends significant ipsilateral projections to both rostral and caudal portions of the dorsal cap, only a few LTN neurons appear to follow this example and only by projecting to the rostral part of the dorsal cap. In addition, both the LTN and dorsal component of the inSFp send sparse contralateral projections to the MTN, nucleus of the optic tract, and visual tegmental relay zone; and the dorsal component of the inSFp also provides a sparse contralateral projection to both rostral and caudal portions of the dorsal cap. The ventral component of the inSFp projects heavily to the ipsilateral visual tegmental relay zone and moderately to the ipsilateral MTN and nucleus of the optic tract. The ventral inSFp projects sparsely to the contralateral MTN, the nucleus of the optic tract, and the visual tegmental relay zone. A few of its neurons target the ipsilateral dorsal cap of the inferior olive. Unlike the DTN (present study) and the MTN (Giolli et al.: J. Comp. Neurol. 227:228-251, '84; J. Comp. Neurol. 232:99-116, '85a), the LTN and the inSFp of the rabbit and rat lack projections to the superior and lateral vestibular nuclei.(ABSTRACT TRUNCATED AT 400 WORDS)     

Nucleus of the solitary tract in the C57BL/6J mouse: Subnuclear parcellation,chorda tympani nerve projections,and brainstem connections

The nucleus of the solitary tract (NST) processes gustatory and related somatosensory information rostrally and general viscerosensory information caudally. To compare its connections with those of other rodents, this study in the C57BL/6J mouse provides a subnuclear cytoarchitectonic parcellation (Nissl stain) of the NST into rostral, intermediate, and caudal divisions. Subnuclei are further characterized by NADPH staining and P2X₂ immunoreactivity (IR). Cholera toxin subunit B (CTb) labeling revealed those NST subnuclei receiving chorda tympani nerve (CT) afferents, those connecting with the parabrachial nucleus (PBN) and reticular formation (RF), and those interconnecting NST subnuclei. CT terminals are densest in the rostral central (RC) and medial (M) subnuclei; less dense in the rostral lateral (RL) subnucleus; and sparse in the ventral (V), ventral lateral (VL), and central lateral (CL) subnuclei. CTb injection into the PBN retrogradely labels cells in the aforementioned subnuclei; RC and M providing the largest source of PBN projection neurons. Pontine efferent axons terminate mainly in V and rostral medial (RM) subnuclei. CTb injection into the medullary RF labels cells and axonal endings predominantly in V at rostral and intermediate NST levels. Small CTb injections within the NST label extensive projections from the rostral division to caudal subnuclei. Projections from the caudal division primarily interconnect subnuclei confined to the caudal division of the NST; they also connect with the area postrema. P2X₂‐IR identifies probable vagal nerve terminals in the central (Ce) subnucleus in the intermediate/caudal NST. Ce also shows intense NADPH staining and does not project to the PBN. J. Comp. Neurol. 522:1565–1596, 2014. © 2013 Wiley Periodicals, Inc.     

Topographic organization of the projections from the interstitial system of the spinal trigeminal tract to the parabrachial nucleus in the rat

Neurons in the paratrigeminal nucleus are known to project to the parabrachial region, but both these areas are heterogeneous, and the subnuclei that account for these connections are not known. To characterize better these projections, we injected small amounts of fluorogold or latex beads labeled with rhodamine or fluorescein into the parabrachial area in the rat and evaluated the retrograde transport of tracer to the paratrigeminal nucleus and neighboring regions. The results show that the rostral part of the paratrigeminal nucleus projects to the medial subnucleus of the parabrachial nucleus. The intermediary part of the paratrigeminal nucleus projects to both the external lateral and to the external medial subnuclei of the parabrachial nucleus. The caudal part of the paratrigeminal nucleus projects to the ventral lateral subnucleus of the parabrachial nucleus. The dorsal paramarginal nucleus projects to the external lateral and the extreme lateral subnuclei of the parabrachial nucleus. Lamina I and II of the spinal trigeminal nucleus also project to the external lateral and the extreme lateral subnuclei of the parabrachial nucleus. In conclusion, the rostral, intermediate, and caudal parts of the paratrigeminal nucleus and the dorsal paramarginal nucleus each have clearly different projection patterns and presumably have different functions.     

Projections from the rhomboid nucleus of the bed nuclei of the stria terminalis: implications for cerebral hemisphere regulation of ingestive behaviors

The basic organization of an exceptionally complex pattern of axonal projections from one distinct cell group of the bed nuclei of the stria terminalis, the rhomboid nucleus (BSTrh), was analyzed with the PHAL anterograde tract-tracing method in rats. Brain areas that receive a strong to moderate input from the BSTrh fall into nine general categories: central autonomic control network (central amygdalar nucleus, descending hypothalamic paraventricular nucleus, parasubthalamic nucleus and dorsal lateral hypothalamic area, ventrolateral periaqueductal gray, lateral parabrachial nucleus and caudal nucleus of the solitary tract, dorsal motor nucleus of the vagus nerve, and salivatory nuclei), gustatory system (rostral nucleus of the solitary tract and medial parabrachial nucleus), neuroendocrine system (periventricular and paraventricular hypothalamic nuclei, hypothalamic visceromotor pattern generator network), orofaciopharyngeal motor control (rostral tip of the dorsal nucleus ambiguus, parvicellular reticular nucleus, retrorubral area, and lateral mesencephalic reticular nucleus), respiratory control (lateral nucleus of the solitary tract), locomotor or exploratory behavior control and reward prediction (nucleus accumbens, substantia innominata, and ventral tegmental area), ingestive behavior control (descending paraventricular nucleus and dorsal lateral hypothalamic area), thalamocortical feedback loops (medial-midline-intralaminar thalamus), and behavioral state control (dorsal raphé and locus coeruleus). Its pattern of axonal projections and its position in the basal telencephalon suggest that the BSTrh is part of a striatopallidal differentiation involved in modulating the expression of ingestive behaviors, although it may have other functions as well.     

Fiber connections of the compact division of the posterior pallial amygdala and lateral part of the bed nucleus of the stria terminalis in the pigeon (Columba livia)

The compact division of the posterior pallial amygdala (PoAc) and lateral part of the bed nucleus of the stria terminalis (BSTL) are components of the limbic system in the pigeon brain. In this study, we examined the position and fiber connections of these two nuclei by using Nissl staining and tract-tracing methods. PoAc occupies a central division in the posterior pallial amygdala. BSTL faces the ventral horn of the lateral ventricle and extends between A 7.25 and A 10.50. PoAc and BSTL connect bidirectionally by the stria terminalis. PoAc connects reciprocally with two nuclear groups in the cerebrum: 1) a continuum consisting of the caudoventral nidopallium, lateral part of the caudoventral nidopallium (NCVl), subnidopallium beneath NCVl, and piriform cortex and 2) rostral areas of the hemisphere, including the frontolateral and frontomedial nidopallium and the densocellular part of the hyperpallium. Extratelencephalic projections of PoAc terminate in the dorsomedial thalamic nuclei and reach the lateral hypothalamic area via the hypothalamic part of the occipito-mesencephalic tract. BSTL also connects reciprocally with two main regions: 1) the same continuum as for PoAc projections, except the piriform cortex and 2) rostral areas of the hemisphere, including the olfactory tubercle and nucleus accumbens. Extratelencephalic reciprocal connections are with the substantia nigra, nucleus subceruleus dorsalis, parabrachial nucleus, locus coeruleus, and nucleus of the solitary tract. The dorsomedial subdivision of the hippocampal formation projects massively to PoAc and BSTL. These findings indicate that PoAc and BSTL are important components of an interconnected neural circuit involving widespread regions of the neuraxis and mediating limbic-visceral functions.     

Diagonal ventral forebrain continuum has overlapping telencephalic inputs and brainstem outputs which may represent loci for limbic/autonomic integration

Growing evidence indicates that three areas within the mammalian basal forebrain share many common features. Based on the similarity of connections and their adjacent spacial proximity, three forebrain nuclei are referred to as a continuum. The components of this diagonal ventral forebrain continuum (DVFC) are the central nucleus of the amygdala, the sublenticular portion of the substantia innominata, and the lateral bed nucleus of the stria terminalis. A primary concern and terminal goal of this study is to determine whether the region of this continuum which projects to the brainstem autonomic nuclei such as the vagal nuclei or the parabrachial nuclei also receives inputs from the basolateral amygdala. The first phase of this study involved determining what autonomic regions receive projections from the basal forebrain. The vagal complex and the parabrachial nuclei were found to receive the densest inputs from the DVFC. The topographic distribution of the respective retrogradely labeled cells and their collateral status is described. The second phase involved looking at afferent inputs from brainstem nuclei. The parabrachial nucleus sends reciprocal projections back to the continuum, which generally overlap the neurons which project back to the brainstem visceral nuclei. The third phase of the study indicated that the cells of the basolateral amygdala contribute a major terminal field which overlaps those cells of the basal forebrain continuum which in turn project to either the nucleus of the solitary tract of the parabrachial nucleus. The possibility that the circuits implied in this study represent the neural circuitry whereby emotional stimuli result in changes in visceral activity is addressed.     

Bidirectional connections of the medial amygdaloid nucleus in the Syrian hamster brain: Simultaneous anterograde and retrograde tract tracing

In the male Syrian hamster, mating is dependent on chemosensory and hormonal stimuli, and interruption of either input prevents copulation. The medial amygdaloid nucleus (Me) is a key nodal point in the neural circuitry controlling male sexual behavior because it relays both odor and steroid cues. Me is comprised of two major subdivisions, anterior (MeA) and posterior (MeP), which have distinct, although overlapping efferent projections. The present study investigated the afferents and efferents of MeA and MeP by using combined anterograde and retrograde tract tracing. Phaseolus vulgaris–leucoagglutinin and cholera toxin B were injected by iontophoresis through a single glass micropipette and detected by immunohistochemistry. MeA has widespread connections with olfactory structures, whereas MeP is heavily interconnected with steroid-responsive brain regions. The efferent projections of MeA and MeP were similar to those reported previously for the rat and hamster. In particular, MeP projects to the posteromedial subdivision of the bed nucleus of the stria terminalis (BNST) and to the medial preoptic nucleus, whereas MeA projects to adjacent subnuclei in BNST and the preoptic area. MeA and MeP also have distinct patterns of afferent input. Furthermore, the combination of anterograde and retrograde tract tracers shows that MeA and MeP are each bidirectionally connected with each other and with limbic nuclei. These results demonstrate that subnuclei of Me are interconnected with limbic structures in hamster brain. These connections may contribute to chemosensory and hormonal integration to control male sexual behavior. J. Comp. Neurol. 399:189–209, 1998. © 1998 Wiley-Liss, Inc.     

Organization of cerebellar and area "y" projections to the nucleus reticularis tegmenti pontis in macaque monkeys

Axonal projections to the nucleus reticularis tegmenti pontis (RTP) were studied in 11 macaque monkeys by mapping axonal degeneration from lesions centered in the dentate and interpositus anterior (IA) nuclei and by mapping anterograde transport of tritiated amino acid precursors injected into the dentate nucleus. Projections from the dentate and IA nuclei overlap in central parts of the body of RTP, but the terminal field of dentate axons extends dorsomedial and rostral to the terminal field of IA axons, and IA terminal fields extend more ventrolaterally. A caudal to rostral topography of projections from each nucleus onto dorsal to ventral parts of RTP was seen. Projections from rostral parts of both nuclei terminate in a sublemniscal part of the nucleus. The topography of dentate and IA projections onto central to ventrolateral RTP appears to match somatotopic maps of these cerebellar nuclei with the somatotopic map of projections to RTP from primary motor cortex. Projections from caudal and ventral parts of the dentate nucleus appear to overlap oculomotor inputs to rostral, dorsal, and medial RTP from the frontal and supplementary eye fields, the superior colliculus, and the oculomotor region of the caudal fastigial nucleus. Projections to the paramedian part of RTP from vestibular area "y" were also found in two cases that correlated with projections to vertical oculomotor motoneurons. The maps of dentate and IA projections onto RTP correlate predictably with maps of dentate and IA projections to the ventrolateral thalamus and subnuclei of the red nucleus that were made from these same cases (Stanton [1980b] J. Comp. Neurol. 192:377-385).     

Organization of axonal projections from the anterolateral area of the bed nuclei of the stria terminalis

The anterolateral group of the bed nuclei of the stria terminalis (BSTalg) contains four distinct cell groups embedded within an undifferentiated anterolateral area (BSTal) that architectonically resembles a subjacent subcommissural zone (BSTsc). The overall distributions of axonal projections from various regions of the BSTal and from the BSTsc were determined with the Phaseolus vulgaris-leucoagglutinin (PHAL) anterograde tracer method and found to be identical. The BSTal and BSTsc share dense bidirectional connections, and also project heavily within the BST to the rhomboid and fusiform nuclei and the anteroventral and anterodorsal areas. They project less densely to the juxtacapsular, oval, magnocellular, ventral, and interfascicular BST nuclei. Outside the BST, brain areas receiving strong to moderate inputs from the BSTal and BSTsc fall into several functional groups: somatomotor system (nucleus accumbens, substantia innominata, ventral tegmental area, and retrorubral area and adjacent midbrain reticular nucleus), central autonomic control system (central amygdalar nucleus, dorsal lateral hypothalamic area, ventrolateral periaqueductal gray, parabrachial nucleus, and nucleus of the solitary tract), neuroendocrine system (paraventricular and supraoptic nuclei, hypothalamic visceromotor pattern generator network), and thalamocortical feedback loops (midline, medial, and intralaminar nuclei). The results indicate that the BSTal and BSTsc are parts of the same cell group (dorsal and ventral to the anterior commissure), which plays a role in coordinating visceral and somatic motor responses (during ingestive behaviors, for example), especially in response to noxious stimuli (learned anorexia associated with noxious stimuli). BSTal projections are distinct from those of the adjacent juxtacapsular, oval, fusiform, and rhomboid nuclei.