Organization of projections from the ventromedial nucleus of the hypothalamus: A Phaseolus vulgaris-Leucoagglutinin study in the rat

The organization of projections from the four parts of the ventromedial nucleus (VMH) and a ventrolaterally adjacent region tentatively identified as the tuberal nucleus (TU) have been analyzed with small injections of the anterograde axonal tracer Phaseolus vulgaris-leucoagglutinin (PHA-L). Extrinsic and intranuclear projections of each part of the VMH display clear quantitative differences, whereas the overall patterns of outputs are qualitatively similar. Overall, the VMH establishes massive intrahypothalamic terminal fields in other parts of the medial zone, tending to avoid the periventricular and lateral zones. The ventrolateral VMH is more closely related to other parts of the hypothalamus that also express gonadal steroid hormone receptors, including the medial preoptic, tuberal, and ventral premamillary nuclei, whereas other parts of the VMH are more closely related to the anterior hypothalamic and dorsal premammillary nuclei. All parts of the VMH project to the zona incerta (including the A13 region) and parts of the midline thalamus, including the paraventricular and parataenial nuclei and nucleus reuniens. The densest inputs to the septum are to the bed nuclei of the stria terminalis, where the ventrolateral and central VMH innervate the anteroventral and anterodorsal areas and transverse and interfascicular nuclei, whereas the anterior and dorsomedial VMH innervate the latter two. The central, lateral, and medial amygdalar nuclei receive substantial inputs from various parts of the VMH. Other regions of the telencephalon, including the nucleus accurmbens and the piriform-amygdaloid, infralimbic, prelimbic, anterior cingulate, agranular insular, piriform, perirhinal, entorhinal, and postpiriform transition areas, also receive sparse inputs. All parts of the VMH send a massive, topographically organized projection to the periaqueductal gray. Other brainstem terminal fields include the superior colliculus, peripeduncular area, locus coeruleus, Barrington's nucleus, parabrachial nucleus, nucleus of the solitary tract, and the mesencephalic, pontine, gigantocellular, paragigantocellular, and parvicellular reticular nuclei. The projections of the TU are similar to, and a subset of, those from the VMH and are thus not nearly as widespread as those from adjacent parts of the lateral hypothalamic area. Because of these similarities, the TU may eventually come to be viewed most appropriately as the lateral component of the VMH itself. The functional implications of the present findings are discussed in view of evidence that the VMH plays a role in the expression of ingestive, affective, and copulatory behaviors. © 1994 Wiley-Liss, Inc.     

Subcortical projections to the centromedian and parafascicular thalamic nuclei in the cat

The primary objective of this study is to identify the totality of input to the centromedian and parafascicular (CM-Pf) thalamic nuclear complex. The subcortical projections upon the CM-Pf complex were studied in the cat with three different retrograde tracers. The tracers used were unconjugated horseradish peroxidase (HRP), horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP), and rhodamine-labeled fluorescent latex microspheres (RFM). Numerous subcortical structures or substructures contained labeled neurons with all three tracing techniques. These labeled structures included the central nucleus of the amygdala; the entopeduncular nucleus; the globus pallidus; the reticular and ventral lateral geniculate nuclei of the thalamus; parts of the hypothalamus including the dorsal, lateral, and posterior hypothalamic areas and the ventromedial and parvicellular nuclei; the zona incerta and fields of Forel; parts of the substantia nigra including the pars reticularis and pars lateralis, and the retrorubral area; the pretectum; the intermediate and deep layers of the superior colliculus; the periaqueductal gray; the dorsal nucleus of the raphe; portions of the reticular formation, including the mesencephalic, pontis oralis, pontis caudalis, gigantocellularis, ventralis, and lateralis reticular nuclei; the nucleus cuneiformis; the marginal nucleus of the brachium conjunctivum; the locus coeruleus; portions of the trigeminal complex, including the principal sensory and spinal nuclei; portions of the vestibular complex, including the lateral division of the superior nucleus and the medial nucleus; deep cerebellar nuclei, including the medial and lateral cerebellar nuclei; and lamina VII of the cervical spinal cord. Moreover, the WGA-HRP and rhodamine methods (known to be more sensitive than the HRP method) revealed several afferent sources not shown by HRP: the anterior hypothalamic area, ventral tegmental area, lateral division of the superior vestibular nucleus, nucleus interpositus, and the nucleus praepositus hypoglossi. Also, the rhodamine method revealed labeled neurons in laminae V and VI of the cervical spinal cord.     

Ascending projections of the posterior nucleus of the hypothalamus: PHA-L analysis in the rat

With the exception of a report by R. B. Veazey, D. G. Amaral, and W. M. Cowan (1982, J. Comp. Neurol. 207:135–156) that examined the projections of the posterior hypothalamic area in the monkey by using the autoradiographic technique, the ascending projections of the posterior nucleus (PH) of the hypothalamus have not been systematically examined in any species. The present report describes the ascending projections of PH in the rat by using the anterograde anatomical tracer, Phaseolus vulgaris -leucoagglutinin (PHA-L). The major ascending route for PH fibers is the medial forebrain bundle. PH fibers project densely to several subcortical and cortical sites. The subcortical sites are the subthalamus/hypothalamus (zona incerta, the supramammillary nucleus, lateral, perifornical, dorsal, and anterior nuclei/areas), the thalamus (lateroposterior, laterodorsal, parafascicular, reuniens, paraventricular, central medial, paracentral, central lateral and intermediodorsal nuclei), the amygdala (central, lateral, and medial nuclei), the septal area (bed nucleus of atria terminalis, medial and lateral septum), and the basal forebrain (horizontal/vertical limbs of diagonal band nuclei and lateral preoptic area). The cortical sites are the perirhinal, insular, frontal (lateral agranular), prelimbic, and infralimbic cortices. The diversity of PH projections to subcortical and cortical “limbic-related” sites and to several structures with direct input to the hippocampus (supramammillary nucleus, reuniens, paraventricular and laterodorsal nuclei of the thalamus, medial and lateral septum, and perirhinal cortex) suggest that the PH may serve a critical role in various components of emotional behavior, including mnemonic processes associated with significant emotional events. © 1995 Wiley-Liss, Inc.     

Projections of the medial preoptic nucleus: a Phaseolus vulgaris leucoagglutinin anterograde tract-tracing study in the rat

The projections of the medial preoptic nucleus (MPN) were examined by making injections of the anterogradely transported lectin Phaseolus vulgaris leucoagglutinin (PHA-L) into the MPN and charting the distribution of labeled fibers. The evidence indicates that the MPN projects extensively to widely distributed regions in both the forebrain and brainstem, most of which also supply inputs to the nucleus. An important neuroendocrine role for the MPN is underscored by its extensive projections to almost all parts of the periventricular zone of the hypothalamus, including the anteroventral periventricular, anterior part of the periventricular, paraventricular (PVH), and arcuate nuclei, and a role in autonomic mechanisms is indicated by projections to such regions as the dorsal and lateral parvicellular parts of the PVH, the lateral parabrachial nucleus, and the nucleus of the solitary tract. Other projections of the MPN suggest participation in the initiation of specific motivated behaviors. For example, inputs to two nuclei of the medial zone of the hypothalamus, the ventromedial and dorsomedial nuclei, may be related to the control of reproductive and ingestive behaviors, respectively, although the possible functional significance of a strong projection to the ventral premammillary nucleus is presently unclear. The execution of these behaviors may involve activation of somatomotor regions via projections to the substantia innominata, zona incerta, ventral tegmental area, and pedunculopontine nucleus. Similarly, inputs to other regions that project directly to the spinal cord, such as the periaqueductal gray, the laterodorsal tegmental nucleus, certain medullary raphe nuclei, and the magnocellular reticular nucleus may also be involved in modulating somatic and/or autonomic reflexes. Finally, the MPN may influence a wide variety of physiological mechanisms and behaviors through its massive projections to areas like the ventral part of the lateral septal nucleus, the bed nucleus of the stria terminalis, the lateral hypothalamic area, the supramammillary nucleus, and the ventral tegmental area, all of which have extensive connections with regions along the medial forebrain bundle. Although the PHA-L method does not allow a clear demonstration of possible differential projections from each subdivision of the MPN, our results suggest that each of them does give rise to a unique pattern of outputs.(ABSTRACT TRUNCATED AT 400 WORDS)     

Direct projections from the dorsolateral pontine tegmentum to pudendal motoneurons innervating the external urethral sphincter muscle in the rat

Direct projections from the dorsolateral pontine tegmentum to pudendal motoneurons innervating the external urethral sphincter and the external anal sphincter muscles were examined in the rat by the tract-tracing methods utilizing retrograde transport of cholera toxin B subunit and anterograde transport of biotinylated dextran amine. The dorsolateral pontine tegmental region, corresponding to the micturition reflex center of Barrington, was confirmed to send bilaterally, with an ipsilateral dominance, projection fibers to the spinal parasympathetic nucleus (inferior intermediolateral nucleus). The micturition reflex center of Barrington, however, did not seem to send many projection fibers to the ventral horn of the lumbosacral cord segments, whereas the region immediately ventral to the micturition reflex center of Barrington was found to send bilaterally, with a contralateral dominance, projection fibers to the dorsolateral group of pudendal motoneurons in both the male and female rats. In the female rat, the dorsolateral group of pudendal motoneurons are comprised primarily of motoneurons that innervate the external urethral sphincter muscle. The dorsomedial group of pudendal motoneurons, which contain motoneurons that innervate the external anal sphincter and the bulbocavernosus muscles, did not seem to receive major projections from the dorsolateral pontine tegmental regions. It was also observed that the locus coeruleus sent some projection fibers bilaterally to the spinal parasympathetic nucleus but only a few to the ventral horn of the lumbosacral cord segments. Thus, the present results indicate that the dorsolateral group of pudendal motoneurons containing those innervating the external urethral sphincter muscle receive pontospinal projection fibers mainly from the dorsolateral pontine tegmental region immediately ventral to the micturition reflex center of Barrington. © 1995 Wiley-Liss, Inc.     

Dorsal mesencephalic projections to pons, medulla, and spinal cord in the cat: limbic and non-limbic components.

The vertebrate dorsal mesencephalon consists of the superior colliculus, the dorsal portion of the periaqueductal gray, and the mesencephalic trigeminal neurons in between. These structures, via their descending pathways, take part in various behavioral responses to environmental stimuli. This study was undertaken to compare the origins and trajectories of these pathways in the cat. Injections of horseradish peroxidase into the cervical spinal cord and upper medullary medial tegmentum retrogradely labeled cells mainly in the contralateral intermediate and deep superior colliculus, and in the ipsilateral dorsal and lateral periaqueductal gray and adjacent tegmentum. Only injections in the medullary lateral tegmental field labeled mesencephalic trigeminal neurons ipsilaterally. Autoradiographic tracing results, based on injections across the dorsal mesencephalon, revealed three efferent fiberstreams. A massive first fiberstream (limbic pathway), consisting of thin fibers, descended ipsilaterally from the dorsal and lateral periaqueductal gray and adjacent superior colliculus through the mesencephalic and pontine lateral tegmentum, terminating in these areas as well as in the ventral third of the caudal pontine and medullary medial tegmentum. A few fibers from the dorsal periaqueductal gray matter (PAG) were distributed bilaterally to the dorsal vagal, solitary, and retroambiguus nuclei. The second fiberstream (the predorsal bundle) descended contralaterally from the superior colliculus (SC) and consisted of both thick and thin labeled fibers. The thin fibers terminated bilaterally in the dorsomedial nucleus reticularis tegmenti pontis and the medial half of the caudal medial accessory inferior olive. The thick fibers targeted the contralateral dorsal two thirds of the caudal pontine and medullary medial tegmental fields, and the facial, abducens, lateral reticular, subtrigeminal, and prepositus hypoglossi nuclei. A few fibers recrossed the midline to terminate in the ipsilateral medial tegmentum. Caudal to the obex, fibers terminated laterally in the tegmentum and upper cervical intermediate zone. From the lateral SC, fibers terminated bilaterally in the lateral tegmental fields of the pons and medulla and lateral facial subnuclei. The third fiberstream (mesencephalic trigeminal or Probst tract) terminated in the supratrigeminal and motor trigeminal nuclei, and laterally in the tegmentum and upper cervical intermediate zone. In summary, neurons in the PAG and in the deep layers of the SC give rise to a massive ipsilateral descending pathway, in which a medial-to-lateral organization exists. A similar topographical pattern occurs in the crossed SC projections. The possibility that these completely different descending systems cooperate in producing specific defensive behaviors is discussed.     

Projections from the periaqueductal gray to the rostromedial pericoerulear region and nucleus locus coeruleus: anatomic and physiologic studies.

Previous studies showed that the nucleus locus coeruleus (LC) receives two major afferent inputs from 1) nucleus paragigantocellularis and 2) nucleus prepositus hypoglossi, both in the rostral medulla. Recent reports suggested that the midbrain periaqueductal gray (PAG) projects to the rostromedial pericoerulear area and LC. Since the PAG is a major site for control of central antinociception, and since descending noradrenergic fibers have been implicated in pain modulation, we have investigated in detail the functional anatomy of projections from PAG to the dorsolateral pontine tegmentum. A combined anatomical and electrophysiological approach was used to assess the organization and synaptic influence of PAG on neurons in the rostromedial pericoerulear region and in LC proper. Injections of the tracer wheatgerm agglutinin conjugated to horseradish peroxidase encompassing LC proper and the rostromedial pericoerulear area retrogradely labeled neurons in PAG located lateral and ventrolateral to the cerebral aqueduct; injections restricted to LC proper did not consistently label PAG neurons. Deposits of the anterograde axonal tracer Phaseolus vulgaris leucoagglutinin into this same region of PAG labeled axons that robustly innervated the zone rostral and medial to LC. Only sparse fibers were observed in LC proper. Consistent with these results, focal electrical stimulation of LC antidromically activated only a few PAG neurons (6 of 100); all of these driven cells were located lateral and ventrolateral to the cerebral aqueduct. The majority of neurons in the rostromedial pericoerulear area were robustly activated by single pulse stimulation of PAG. In contrast, single pulse electrical stimulation of lateral PAG produced weak to moderate synaptic activation of some LC neurons; stimulation of ventrolateral PAG produced predominant inhibition of LC discharge, perhaps through recurrent collaterals subsequent to antidromic activation of neighboring LC cells. Taken together, these results indicate that PAG strongly innervates the region rostral and medial to LC, including Barrington's nucleus, but only weakly innervates LC proper. Although recent studies indicate that the dendrites of LC neurons ramify heavily and selectively in the rostromedial pericoerulear region, the results of the present physiological studies suggest that PAG preferentially targets rostromedial pericoerulear neurons rather than LC dendrites.     

Distribution of locus coeruleus axons within the rat brainstem demonstrated by Phaseolus vulgaris leucoagglutinin anterograde tracing in combination with dopamine-beta-hydroxylase immunofluorescence

Projections of the locus coeruleus (LC) to the midbrain and hindbrain were analyzed by anterograde transport of the lectin Phaseolus vulgaris leucoagglutinin (PHA-L). Following iontophoretic application of PHA-L into the LC, the distribution of labeled axons was analyzed in sections processed for the immunoperoxidase method and in sections processed for double-immunofluorescence staining using antibodies to PHA-L and to dopamine-beta-hydroxylase. This combined staining approach proved to be necessary for the unequivocal identification of LC axons in the brainstem since all injections labeled many non-noradrenergic axons whose distribution was different from that of LC fibers. The major new finding of the present study was the observation that large territories of the brainstem that receive a dense noradrenergic input are very sparsely innervated by the LC. Numerous labeled LC axons were observed in somatic afferent nuclei, tectum, pontine nuclei, interpenduncular nucleus, and inferior olivary complex. In contrast, very few labeled fibers were observed in autonomic and motor nuclei, and throughout the brainstem reticular formation, including raphe nuclei. Our data show that the distribution of LC axons in the brainstem is far less prominent than the projections of this nucleus to the forebrain and spinal cord. Our findings suggest that the dense NA projections to the core of the brainstem originate principally in non-LC NA neurons. On the basis of the present anatomical findings, a prominent role of the LC in motor and integrative functions of the brainstem appears unlikely.     

Descending projections of the basal forebrain in the rat demonstrated by the anterograde neural tracer Phaseolus vulgaris leucoagglutinin (PHA-L)

Descending pathways from the mediobasal forebrain were studied in the rat by injecting anterograde axonal tracer Phaseolus vulgaris leucoagglutinin into the substantia innominata and diagonal band of Broca. From both areas, positive fibers which varied in density were observed in the mediodorsal and ventral parts of the ventroposterior and ventromedial thalamic nuclei, the lateral habenula, the stria medullaris, the lateral hypothalamus and the ventral tegmental area. This descending complex appeared predominantly course through the medial forebrain bundle from which positive fibers ramified into the fasciculus thalamicus to distribute in the thalamic nuclei. A minor descending pathway through the stria medullaris was also noted which terminated in the lateral habenula and the mediodorsal thalamic nucleus. An obvious difference in terminal distribution in the medial habenula, mediodorsal thalamic nucleus and pons could be observed following substantia innominata or diagonal band injection.     

Electrophysiological mapping of brainstem projections of spinal cord lamina I cells in the rat

The course and terminations of the spinal and supraspinal projections of rat dorsal horn lamina I cells have been determined by antidromically activating the cells with a roving stimulating microelectrode which was used to map systematically the brainstem of the animals. The cells studied are the most common type of projection cell in lamina I of the rat with ascending axons coursing in the contralateral dorsolateral funiculus at C2. In the present study we have found that the axons decussate within 1–5 mm of the cell body. The ascending axons occupy a peripheral site within the dorsolateral funiculus at C2. Within the brainstem it was possible to identify a main or ‘parent’ axon which showed a progressive drop in conduction velocity as it coursed rostrally. All these terminated in the midbrain. In addition to the main branch each axon gave rise to several collaterals and these terminated in: (a) ventrolateral periaqueductal grey and the immediately lateral n. cuneiformis; and (b) the dorsal medullary reticular formation. (i.e. most fibres had two different projection zones). Since these cells did not project to the thalamus, it may be that the information they carry is used other than for purely sensory processes. One possibility is that they impinge on an antinociceptive system believed to originate in the periaqueductal grey.     

Distribution of spinal cord projections from the medullary subnucleus reticularis dorsalis and the adjacent cuneate nucleus: A phaseolus vulgaris- leucoagglutinin study in the rat

The distribution and organization of descending spinal projections from the dorsal part of the caudal medulla were studied in the rat following injections of Phaseolus vulgaris leucoagglutinin into small areas of the subnucleus reticularis dorsalis (SRD) and the adjacent cuneate nucleus (Cu). The caudal aspect of the Cu projected only to the dorsal horn of the ipsilateral cervical cord via the dorsal funiculus. These projections were mainly to laminae I, IV, and V. More ventrally located reticular structures projected to the full length of the cord. Fibers originating from the SRD travelled through the ipsilateral dorsolateral funiculus and terminated within the deep dorsal horn and upper layers of the ventral horn, mainly in laminae V–VII. Fibers originating from subnucleus reticularis ventralis (SRV) travelled ipsilaterally through the lateral and ventrolateral funiculi and bilaterally through the ventromedial funiculus. These fibers terminated within the ventral horn. The density of labeling within the gray matter varied at different levels of the cord was as follows: cervical > sacral > thoracic > lumbar. The reciprocal connections between the caudal medulla and the spinal cord suggest that the former is an important link in feedback loops that regulate spinal outflow. © 1995 Wiley-Liss, Inc.     

PHA-L analysis of projections from the supramammillary nucleus in the rat.

The projections of the supramammillary nucleus (SUM) were examined in the rat by the anterograde anatomical tracer Phaseolus vulgaris leucoagglutinin (PHA-L). The majority of labeled fibers from SUM ascended through the forebrain within the medial forebrain bundle. SUM fibers were found to terminate heavily in the hippocampal formation, specifically within the granule cell layer and immediately adjoining molecular layer of the dentate gyrus. In addition, SUM fibers were shown to distribute densely to several structures with strong connections with the hippocampus, namely, the nucleus reunions of the thalamus, the medial and lateral septum, the entorhinal cortex, and the endopiriform nucleus. SUM fibers were also shown to project significantly to several additional subcortical and cortical sites. The subcortical sites were the dorsal raphe nucleus, the midbrain central gray, the fields of Forel/zona incerta, the dorsomedial hypothalamic area, midline/intralaminar nuclei of the thalamus (posterior paraventricular, rhomboid, central medial, intermediodorsal, and mediodorsal), the medial and lateral preoptic areas, the bed nucleus of the stria terminalis, the substantia innominata, the vertical limb of the diagonal band nucleus, and the claustrum. The cortical sites were the occipital, temporal, parietal, and frontal cortices. Some notable differences were observed in projections from the lateral as compared to the medial SUM. For example, fibers originating from the lateral SUM distributed heavily to the hippocampal formation and parts of the cortex, whereas those from the medial SUM projected sparsely to these two regions. The SUM projections to the hippocampal formation and associated structures may serve as the substrate for a SUM involvement in the generation of the theta rhythm of the hippocampus and the gating of information flow through the hippocampal formation.     

Ascending projections from the pedunculopontine tegmental nucleus and the adjacent mesopontine tegmentum in the rat

Ascending projections from the pedunculopontine tegmental nucleus (PPT) and the surrounding mesopontine tegmentum to the forebrain in the rat are here examined by using both retrograde and anterograde tracing techniques combined with choline acetyltransferase (ChAT) immunohistochemistry. The anterogradely transported lectin Phaseolus vulgaris-leukoagglutinin (PHA-L) was iontophoretically injected into the PPT in 12 rats. Anterogradely labelled fibers and varicosities were observed in the thalamic nuclei, confirming the findings of our previous retrograde studies (Hallanger et al: J. Comp. Neurol. 262:105-124, '87). In addition, PHA-L-labelled fibers and varicosities suggestive of terminal fields were observed in the anterior, tuberal, and posterior lateral hypothalamic regions, the ventral pallidum in the region of the nucleus basalis of Meynert, the dorsal and intermediate lateral septal nuclei, and in the central and medial nuclei of the amygdala. To determine whether these were cholinergic projections, the retrograde tracer WGA-HRP was injected into terminal fields in the hypothalamus, septum, ventral pallidum, and amygdala. Numerous ChAT-immunoreactive neurons in the PPT and laterodorsal tegmental nucleus (LDT) were retrogradely labelled from the lateral hypothalamus. These cholinergic neurons constituted over 20% of those retrogradely labelled in the dorsolateral mesopontine tegmentum; the balance consisted of noncholinergic neurons of the central tegmental field, retrorubral field, and cuneiform nucleus. Following placement of WGA-HRP into dorsal and intermediate lateral septal regions, the vast majority (greater than 90%) of retrogradely labelled neurons were cholinergic neurons of the PPT and LDT, with few noncholinergic retrogradely labelled neurons in the adjacent tegmentum. In contrast, fewer cholinergic neurons were retrogradely labelled following placement of tracer into the nucleus basalis of Meynert or into the central, medial, and basolateral nuclei of the amygdala, while numerous noncholinergic neurons of the central tegmental field rostral to the PPT and of the retrorubral field adjacent to the PPT were retrogradely labelled in these cases. These anterograde and retrograde studies demonstrate that cholinergic PPT and LDT neurons provide a substantial proportion of mesopontine tegmental afferents to the hypothalamus and lateral septum, while projections to the nucleus basalis and the amygdala are minimal.     

Orbitomedial prefrontal cortical projections to hypothalamus in the rat

A previous study in the rat revealed that distinct orbital and medial prefrontal cortical (OMPFC) areas projected to specific columns of the midbrain periaqueductal gray region (PAG). This study used anterograde tracing techniques to define projections to the hypothalamus arising from the same OMPFC regions. In addition, injections of anterograde and retrograde tracers were made into different PAG columns to examine connections between hypothalamic regions and PAG columns projected upon by the same OMPFC regions. The most extensive patterns of hypothalamic termination were seen after injection of anterograde tracer in prelimbic and infralimbic (PL/IL) and the ventral and medial orbital (VO/MO) cortices. Projections from rostral PL/IL and VO/MO targeted the rostrocaudal extent of the lateral hypothalamus, as well as lateral perifornical, and dorsal and posterior hypothalamic areas. Projections arising from caudal PL/IL terminated within the dorsal hypothalamus, including the dorsomedial nucleus and dorsal and posterior hypothalamic areas. There were also projections to medial perifornical and lateral hypothalamic areas. In contrast, it was found that anterior cingulate (AC), dorsolateral orbital (DLO), and agranular insular (AId) cortices projected to distinct and restricted hypothalamic regions. Projections arising from AC terminated within dorsal and posterior hypothalamic areas, whereas DLO and AId projected to the lateral hypothalamus. The same OMPFC regions also projected indirectly, by means of specific PAG columns, to many of the same hypothalamic fields. In the context of our previous findings, these data indicate that, in both rat and macaque, parallel but distinct circuits interconnect OMPFC areas with specific hypothalamic regions, as well as PAG columns.     

Reticulo-olivary circuits: An intracellular HRP study in the rat

Intracellular recordings were obtained from medullary reticular neurons subsequent to electrical stimulation of the ipsilateral or contralateral inferior cerebellar peduncle (ICP) and/or the midbrain. After recording physiological data, the neurons were intracellularly injected with horseradish peroxidase (HRP). Thirty-four HRP filled neurons were subjected to light microscopic analysis. They could be divided into two general groups: those which extend dendritic processes into the neuropil of the inferior olivary complex (n = 19); and those that have no anatomical relationship to the inferior olive (n = 15). These two populations of reticular neurons differ in their distribution, morphological characteristics and physiological responses. Neurons which extend dendritic processes into the inferior olive are located within 200 microns of the dorsal border of this nuclear complex, between the exiting fibers of the XIIth nerve and the raphe. The cell bodies are located in the nucleus reticularis gigantocellularis and are fusiform or multipolar in shape. Their dendrites extend for long distances in the mediolateral direction; are thin and relatively spine-free except at their distal tips where spines and varicose appendages are evident. Physiologically, midbrain stimulation elicits a fast rising hyperpolarization which is identified as an inhibitory postsynaptic potential. However, only rarely is a response observed subsequent to stimulation of either the ipsilateral or contralateral ICP. Dendrites from 4 neurons from the first group of reticular cells were analyzed at the ultrastructural level. Based on random and serial thin sections, the following features were noted: they contain numerous mitochondria when compared to olivary dendrites; they contribute to the postsynaptic elements within olivary synaptic clusters (glomeruli); and they exhibit focal clusters of synaptic vesicles although conventional synaptic complexes have not been observed. Reticular neurons of the second group, those that do not extend dendritic processes into the inferior olive, are located either lateral to the XIIth nerve or at distances greater than 200 microns from the dorsal border of the inferior olivary complex. Their cell bodies and dendrites are comparable morphologically to the reticular neurons whose dendrites do arborize in the inferior olive. However, rarely are the distal tips of their dendrites characterized by spines or varicose appendages. Physiologically, this population of reticular neurons respond to midbrain stimulation with a low amplitude, short latency depolarizing potential which is interrupted by a hyperpolarizing potential.(ABSTRACT TRUNCATED AT 400 WORDS)     

Projections of the dorsal raphe nucleus to the brainstem: PHA-L analysis in the rat

Early studies that used older tracing techniques reported exceedingly few projections from the dorsal raphe nucleus (DR) to the brainstem. The present report examined DR projections to the brainstem by use of the anterograde anatomical tracer Phaseolus vulgaris leucoagglutinin (PHA-L). DR fibers were found to terminate relatively substantially in several structures of the midbrain, pons, and medulla. The following pontine and midbrain nuclei receive moderate to dense projections from the DR: pontomesencephalic central gray, mesencephalic reticular formation, pedunculopontine tegmental nucleus, medial and lateral parabrachial nuclei, nucleus pontis oralis, nucleus pontis caudalis, locus coeruleus, laterodorsal tegmental nucleus, and raphe nuclei, including the central linear nucleus, median raphe nucleus, and raphe pontis. The following nuclei of the medulla receive moderately dense projections from the DR: nucleus gigantocellularis, nucleus raphe magnus, nucleus raphe obscurus, facial nucleus, nucleus gigantocellularis-pars alpha, and the rostral ventrolateral medullary area. DR fibers project lightly to nucleus cuneiformis, nucleus prepositus hypoglossi, nucleus paragigantocellularis, nucleus reticularis ventralis, and hypoglossal nucleus. Some differences were observed in projections from rostral and caudal parts of the DR. The major difference was that fibers from the rostral DR distribute more widely and heavily than do those from the caudal DR to structures of the medulla, including raphe magnus and obscurus, nucleus gigantocellularis-pars alpha, nucleus paragigantocellularis, facial nucleus, and the rostral ventrolateral medullary area. A role for the dorsal raphe nucleus in several brainstem controlled functions is discussed, including REM sleep and its events, nociception, and sensory motor control. © Wiley-Liss, Inc.     

Amygdalonigral pathway: an anterograde study in the rat with Phaseolus vulgaris leucoagglutinin (PHA-L)

The projection from the central nucleus of the amygdala to the substantia nigra was labeled by injections of the anterograde tracer Phaseolus vulgaris leucoagglutinin into different subregions of the nucleus. A sparse projection of labeled bouton-like swellings was observed in the rostral, medial substantia nigra pars compacta and ventral tegmental area from all subregions of the central nucleus of the amygdala that were injected. A dense projection of labeled axons and bouton-like swellings was observed in the lateral part of the substantia nigra pars compacta and pars lateralis when the injection site included the dorsal and rostral central nucleus. Heavy labeling was also seen in the lateral retrorubral field in these cases. In no instances were labeled terminals observed in the substantia nigra pars reticulata. The same pattern of labeling in the lateral substantia nigra and retrorubral field was seen after injections rostral to the central nucleus or dorsal and medial to it in the sublenticular region. The results suggest that the amygdalonigral pathway contributes to the innervation of extensive areas of the substantia nigra pars compacta. The major component of the pathway, however, projects only to a subregion of the substantia nigra. The origin of this pathway is confined to a discrete region of the dorsal central nucleus of the amygdala but extends rostrally into an area that is part of the "extended amygdala."     

Descending projections from the nucleus retroambiguus to the iliopsoas motoneuronal cell groups in the female golden hamster: Possible role in reproductive behavior

In the cat, the nucleus retroambiguus (NRA) projects to expiratory motoneurons in the brainstem and spinal cord. Recently, it has been demonstrated that the NRA sends fibers to a specific set of motoneurons in the lumbosacral cord, which pathway is thought to play a crucial role in mating behavior. The question is whether such projections exist in the hamster, because the female of this species displays a very distinctive receptive behavior. In the hamster, lumbosacral cord injections of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) combined with hemisection 1 or 2 segments rostral to injection sites in three of the five cases demonstrated retrogradely labeled neurons in the NRA at levels 1.0–2.25 mm caudal to the obex, contralateral to the injection sites. Injections of WGA-HRP into the NRA and adjoining reticular formation revealed that NRA fibers crossed the midline in the caudal medulla and descended in the contralateral lateral and ventrolateral funiculi to terminate bilaterally, but mainly contralaterally, in the motoneuronal cell groups of the abdominal wall and iliopsoas muscles. NRA projections to levels caudal to lumbar segment 5 were virtually absent. Electron microscopic examination revealed that, of the 162 labeled NRA terminal profiles found in the ultrathin sections, 144 (89%) made monosynaptic contacts with retrogradely labeled dendrites of iliopsoas motoneurons. These NRA terminals formed asymmetrical synapses and contained spherical vesicles indicative of an excitatory function. The results indicate that, in the hamster, direct contralateral NRA projections exist to iliopsoas motoneurons. A concept is discussed in which this pathway plays a crucial role in mating behavior. J. Comp. Neurol. 403:219–228, 1999. © 1999 Wiley-Liss, Inc.