Spinal and trigeminal dorsal horn projections to the parabrachial nucleus in the rat

We studied afferents to the parabrachial nucleus (PB) from the spinal cord and the spinal trigeminal nucleus pars caudalis (SNVc) in the rat by using the anterograde and retrograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Injections of WGA-HRP into medial PB retrogradely labeled neurons in the promontorium and in lamina I of the dorsal rostral SNVc, while injections into lateral PB and the K?lliker-Fuse nucleus retrogradely labeled neurons in these areas as well as in lamina I throughout the caudal SNVc and spinal dorsal horn. Injections of WGA-HRP into the caudal SNVc and dorsal horn of the spinal cord resulted in terminal labeling in the dorsal, central, and external lateral subnuclei of PB and the K?lliker-Fuse nucleus, all of which are known to receive cardiovascular and respiratory afferent information. Injections of WGA-HRP into the promontorium and dorsal rostral SNVc resulted in terminal labeling in the same PB subnuclei, as well as in the medial and the ventral lateral PB subnuclei, which are sites of relay for gustatory information ascending from the medulla to the forebrain. The spinal and trigeminal projection to PB may mediate the convergence of pain, chemosensory, and temperature sensibilities with gustatory and cardiorespiratory systems in PB.     

Spinal and trigeminal projections to the nucleus of the solitary tract: a possible substrate for somatovisceral and viscerovisceral reflex activation

This study used the retrograde transport of a protein-gold complex to examine the distribution of spinal cord and trigeminal nucleus caudalis neurons that project to the nucleus of the solitary tract (NST) in the rat. In the spinal grey matter, retrogradely labeled cells were common in the marginal zone (lamina I), in the lateral spinal nucleus of the dorsolateral funiculus, in the reticular part of the neck of the dorsal horn (lamina V), around the central canal (lamina X), and in the region of the thoracic and sacral autonomic cell columns. The pattern of labeling closely resembled that seen for the cells at the origin of the spinomesencephalic tract and shared some features with that of the spinoreticular and spinothalamic tracts. Labeled cells in lamina IV of the dorsal horn were only observed when injections spread dorsally, into the dorsal column nuclei, and are thus not considered to be at the origin of the spinosolitary tract. They are probably neurons of the postsynaptic fibers of the dorsal column. Retrogradely labeled cells were also numerous in the superficial laminae of the trigeminal nucleus caudalis, through its rostrocaudal extent. The pattern of marginal cell labeling appeared to be continuous with that of labeled neurons in the paratrigeminal nucleus, located in the descending tract of trigeminal nerve. Since the NST is an important relay for visceral afferents from both the glossopharyngeal and vagus nerves, we suggest that the spinal and trigeminal neurons that project to the NST may be part of a larger system that integrates somatic and visceral afferent inputs from wide areas of the body. The projections may underlie somatovisceral and/or viscerovisceral reflexes, perhaps with a significant afferent nociceptive component.     

The ascending input to the midbrain periaqueductal gray of the primate

To obtain a comprehensive map of the brainstem and spinal cord areas that project to the mesencephalic central gray small injections of hors-radish peroxidase were made into various regions of the periaqueductal gray in a series of monkeys. Despite the fact that different regions of the central gray were injected in separate animals, the majority of the brainstem areas containing retrogradely filled neurons remained the same. Labeled neurons were observed in the superior colliculus, periaqueductal gray, lateral parabrachial, locus coeruleus, nucleus raphe magnus and pallidus, and a variety of brainstem reticular nuclei. In contrast to labeled brainstem areas, where labeled neurons were present predominantly ipsilateral to the injection site, the spinal trigeminal nucleus pars caudalis and the spinal cord displayed labeled cells chiefly on the side contralateral to the injection. Also in contrast to the labeled brainstem sites, where medial and lateral injection sites produced a similar pattern of labeling, medial injections in the PAG labeled almost exclusively neurons in the deep laminae (V-X) in the spinal trigeminal nucleus pars caudalis and spinal cord while more lateral injections labeled neurons in both the deep (V-X) and superficial (I) laminae. No consistent differences were noted in the location of labeled neurons in either brainstem or spinal sites after dorsal vs. ventral injections or caudal vs. rostral injection sites. The present study has demonstrated that the central gray receives afferent projections from a number of brainstem and spinal areas which are known to be involved in the modulation andor conduction of nociception, while other inputs are probably involved in the regulation of visceral functions. These data support the hypothesis that the mesencephalic periaqueductal gray functions as a visceral, nociceptive, and cognitive integrator.     

Distribution of trigeminohypothalamic and spinohypothalamic tract neurons displaying substance P receptor-like immunoreactivity in the rat

Primary afferent neurons containing substance P (SP) are apparently implicated in the transmission of noxious information from the periphery to the central nervous system, and SP released from primary afferent neurons acts on second-order neurons with the SP receptor (SPR). In the rat, nociceptive information reached the hypothalamus not only through indirect pathways but also directly through trigeminohypothalamic and spinohypothalamic pathways. Thus, in the present study, the distribution pattern of trigeminohypothalamic and spinohypothalamic tract neurons showing SPR-like immunoreactivity (SPR-LI) was examined in the rat by a retrograde tract-tracing method combined with immunofluorescence histochemistry for SPR. A substantial number of trigeminal and spinal neurons with SPR-LI were retrogradely labeled with Fluoro-Gold (FG) injected into the hypothalamic regions. These neurons were distributed mainly in lamina I of the medullary and spinal dorsal horns, lateral spinal nucleus, regions around the central canal of the spinal cord, and the lateral aspect of the deep part of the spinal dorsal horn. A number of SPR-LI neurons in the spinal parasympathetic nucleus were labeled with FG injected into the area around the paraventricular hypothalamic nucleus. Some SPR-LI neurons in the lateral spinal nucleus and the lateral aspect of the deep part of the spinal dorsal horn were also labeled with FG injected into the septal region. On the basis of the distribution areas of SPR-LI trigeminal and spinal neurons projecting to the hypothalamic and septal regions, it is likely that these neurons are involved in the transmission of somatic and/or visceral noxious information. J. Comp. Neurol. 378:508–521, 1997. © 1997 Wiley-Liss, Inc.     

Cells of origin of the trigeminohypothalamic tract in the rat

Recent studies have demonstrated that a large number of spinal cord neurons convey somatosensory and visceral nociceptive information directly from cervical, lumbar, and sacral spinal cord segments to the hypothalamus. Because sensory information from head and orofacial structures is processed by all subnuclei of the trigeminal brainstem nuclear complex (TBNC) we hypothesized that all of them contain neurons that project directly to the hypothalamus. In the present study, we used the retrograde tracer Fluoro-Gold to examine this hypothesis. Fluoro-Gold injections that filled most of the hypothalamus on one side labeled approximately 1,000 neurons (best case = 1,048, mean = 718 ± 240) bilaterally (70% contralateral) within all trigeminal subnuclei and C1–2. Of these neurons, 86% were distributed caudal to the obex (22% in C2, 22% in C1, 23% in subnucleus caudalis, and 18% in the transition zone between subnuclei caudalis and interpolaris), and 14% rostral to the obex (6% in subnucleus interpolaris, 4% in subnucleus oralis, and 4% in subnucleus principalis). Caudal to the obex, most labeled neurons were found in laminae I–II and V and the paratrigeminal nucleus, and fewer neurons in laminae III–IV and X. The distribution of retrogradely labeled neurons in TBNC gray matter areas that receive monosynaptic input from trigeminal primary afferent fibers innervating extracranial orofacial structures (such as the cornea, nose, tongue, teeth, lips, vibrissae, and skin) and intracranial structures (such as the meninges and cerebral blood vessels) suggests that sensory and nociceptive information originating in these tissues could be transferred to the hypothalamus directly by this pathway. J. Comp. Neurol. 400:125–144, 1998. © 1998 Wiley-Liss, Inc.     

Afferent input to nucleus submedius in rats: retrograde labeling of neurons in the spinal cord and caudal medulla

In cats, spinal and medullary input to the thalamic nucleus submedius (Sm) arises almost exclusively from neurons in the marginal zone. As a result, it has been proposed that Sm may be specifically involved in nociception. In the present study, we determined the locations of neurons in the spinal cord and caudal medulla that project to Sm in rats. Iontophoretic injections of Fluoro-Gold or pressure injections of Fast blue were made into Sm. In each of the 6 rats that received small injections of Fluoro-Gold into Sm, only a small number (mean = 90) of retrogradely labeled neurons were found throughout the 18 segments of the spinal cord examined. Surprisingly, almost no labeled neurons (less than 1%) were counted in the marginal zone of the spinal cord. The majority were located in the deep dorsal horn and intermediate zone/ventral horn. In contrast, many neurons were labeled in the marginal zone of nucleus caudalis. Injections of Fluoro-Gold into any of a number of nuclei near Sm also labeled only a small number of neurons in the spinal cord and almost no neurons in the marginal zone. Using identical injection parameters, we injected Fluoro-Gold into the ventrobasal complex or posterior thalamic group. Hundreds of neurons in the spinal cord, including many in the marginal zone, were labeled following these injections. These results indicate that the techniques used to inject Fluoro-Gold into Sm were capable of labeling many projection neurons, including those in the marginal zone. Larger pressure injections of Fast blue were also made into Sm of 3 rats. The distribution of labeled neurons in nucleus caudalis and the spinal cord was similar to that following iontophoretic injections of Fluoro-Gold. Again, few marginal zone neurons were labeled in the spinal cord in any of these rats. Therefore, our results indicate that few spinothalamic tract neurons appear to project to Sm or any of several adjacent nuclei, and virtually no marginal zone neurons in the spinal cord project to these areas.     

Hypothalamic paraventricular axons projecting to the female rat lumbosacral spinal cord contain oxytocin immunoreactivity

Oxytocin-containing axons project from the hypothalamic paraventricular nucleus to the neurohypophysis and thoracic spinal cord to ultimately influence uterine contractions and autonomic activity, respectively. Whether or not oxytocin-immunoreactive axons project to the female rat lumbosacral spinal cord to influence autonomic outflow to pelvic organs has not been investigated. Thus, the present study was designed to investigate the presence, distribution, and origin of oxytocin-immunoreactive axons in the female rat lumbosacral spinal cord. Immunohistochemistry, spinal cord transections, and axonal tracing with Fluorogold, True Blue, and pseudorabies virus were used. Oxytocin-immunoreactive nerve fibers were present in the L6/S1 segments of the spinal cord. Prominent varicose axons were evident throughout the dorsal horn, along the lateral and medial collateral pathways, in the dorsal intermediate gray area, around the central canal in lamina X, and throughout the sacral parasympathetic nucleus. Injection of retrograde tracer into the L6/S1 spinal cord labeled neurons in the hypothalamic paraventricular nucleus. Transection of the thoracic spinal cord eliminated oxytocin-immunoreactive nerve axons in the L6/S1 spinal cord. In addition, transection of the thoracic spinal cord eliminated transport of retrograde axonal tracer from the L6/S1 spinal cord to the paraventricular nucleus. Pseudorabies virus, a transneuronal retrograde tracer, injected into the uterus and cervix marked uterine-related preganglionic neuronal cell bodies in the sacral parasympathetic nucleus and uterine-related neurons in the hypothalamic paraventricular nucleus. Double immuno-labeling of viral-infected spinal cord sections showed oxytocin-immunoreactive axons closely associated with viral labeled uterine-related preganglionic cell bodies of the sacral parasympathetic nucleus. The results of this study revealed that oxytocin-immunoreactive neurons of the hypothalamic paraventricular nucleus project axons to the lumbosacral spinal cord to areas involved in sensory processing and parasympathetic outflow to the uterus.     

The afferent and efferent connections of the nucleus submedius in the rat.

The afferent and efferent connections of the nucleus submedius (Sm) in the medial thalamus of the rat were examined. Injections of wheat-germ agglutinin conjugated horseradish peroxidase (WGA-HRP) into the Sm resulted in dense terminal labeling in the middle layers of the ipsilateral ventrolateral orbital cortex (VLO). Less dense labeling was also observed in the superficial and deep layers of VLO and in the medial part of the lateral orbital cortex (LO) and in the contralateral VLO. Retrogradely labeled neurons were observed primarily in the deep layers of VLO and the dorsal peduncular cortex (DP). Labeled neurons were also observed bilaterally, in the nucleus of the horizontal limb of the diagonal band, the lateral hypothalamus, the thalamic reticular nucleus (Rt), medial parabrachial nucleus (MPB), and the laterodorsal tegmental nucleus (LDT). Many labeled neurons were also observed in the trigeminal brain-stem complex. Injections of Fluoro-Gold (FG) into Sm resulted in a very similar distribution of retrogradely labeled neurons. Injections of WGA-HRP and FG in the orbital cortex confirmed the ipsilateral Sm projection to VLO and suggested that the middle and deep layers of VLO receive a specific ipsilateral projection from the dorsal Sm and that the superficial layers receive a projection primarily from the ventral Sm. Injections of WGA-HRP into the lateral hypothalamus, LDT, and MPB confirmed the retrograde labeling findings; the lateral hypothalamus was found to send a projection to the medial Sm, the LDT region to the ventromedial Sm and the MPB to the medial and dorsal Sm. These findings confirm and extend the results of previous studies in cat and rat indicating that Sm has a major and specific reciprocal connection with VLO. This finding, in conjunction with previous studies showing direct spinal and trigeminal inputs and the existence of nociceptive neurons in Sm and VLO, provides further support for a role of Sm in nociception.     

Evidence for corticotropin-releasing hormone projections from Barrington's nucleus to the periaqueductal gray and dorsal motor nucleus of the vagus in the rat

The present study used anterograde and retrograde tract tracing and immunohistochemistry to determine the efferent projections of corticotropin-releasing hormone (CRH) neurons of Barrington's nucleus in the rat. Injections of Phaseolus vulgaris-leucoagglutinin into Barrington's nucleus resulted in anterograde labeling in the dorsal motor nucleus of the vagus, periaqueductal gray, medial thalamic nuclei, lateral hypothalamus, paraventricular nucleus of the hypothalamus, lateral preoptic area, and lateral septum. The retrograde tract tracer, fluorogold, injected into the lumbosacral spinal cord labeled many, but not all, CRH-immunoreactive neurons in Barrington's nucleus. Moreover, some Barrington's neurons that were retrogradely labeled from the spinal cord were not CRH-immunoreactive. Several CRH-immunoreactive Barrington's neurons were retrogradely labeled by fluorogold injections into the periaqueductal gray, and these were located predominantly in the dorsal part of the nucleus. Additionally, some CRH-immunoreactive Barrington's neurons were retrogradely labeled from fluorogold injections into the dorsal motor nucleus of the vagus. In contrast, fluorogold injections into the lateral hypothalamus, lateral preoptic area, or lateral septum did not result in double labeling of CRH-immunoreactive neurons in Barrington's nucleus. These results suggest that many, but not all, CRH-containing neurons of Barrington's nucleus project to the lumbosacral spinal cord. In addition to their previously documented projections to the spinal cord, these neurons may be a source of CRH in the periaqueductal gray and dorsal motor nucleus of the vagus. CRH projections of Barrington's nucleus may play a role in behavioral or autonomic aspects of stress responses, in addition to their proposed role in micturition. © 1995 Wiley-Liss, Inc.     

The origins of supraspinal projections to the cervical and lumbar spinal cord at different stages of development in the gray short-tailed Brazilian opossum, Monodelphis domestica.

We have used the retrograde transport of Fast blue (FB) to study the origins of supraspinal projections to the lumbar and cervical spinal cord at different stages of development in the Brazilian, short-tailed opossum, Monodelphis domestica. Monodelphis was chosen for study because its young are born in a very immature state, 14-15 days after copulation, making it possible to manipulate its nervous system in an embryonic state without intra-uterine surgery. When injections of FB were made into the lumbar cord at postnatal day (PD) 1, neurons were labeled within several areas of the reticular formation (the retroambiguus nucleus, the ventral and dorsal reticular nuclei of the medulla, the gigantocellular reticular nucleus, the lateral paragigantocellular reticular nucleus, and the pontine reticular nucleus), the presumptive coeruleus complex, and the lateral vestibular nucleus. In many cases, labeled neurons were also found within the caudal raphe and the presumptive interstitial nucleus of the medial longitudinal fasciculus. The results of immunocytochemical studies provided evidence for catecholaminergic and serotoninergic neurons in the brainstem at PD1 and for axons of both phenotypes in the spinal cord. By PD3, labeled neurons were found within the ventral gigantocellular and ventral pontine nuclei of the reticular formation, the spinal trigeminal nucleus, and the presumptive paraventricular nucleus of the hypothalamus. When injections were made at PD4, neurons were also labeled within the medial and inferior vestibular nuclei, the red nucleus, the mesencephalic nucleus of the trigeminal nerve, the presumptive nucleus of Edinger-Westphal and the lateral hypothalamus. By at least PD7, the pattern of supraspinal labeling was similar to that obtained at older ages and in the adult animal. When FB was injected into the cervical cord at PD1, neurons were labeled in all of the areas labeled by lumbar injections at the same age and in larger numbers. In addition, labeled neurons were found within the ventral gigantocellular and spinal trigeminal nuclei. When cervical injections were made at PD15, labeled neurons were found within the deep cerebellar nuclei and amygdala and by PD17 they were also present within the superior colliculus and cerebral cortex. In some cases, cortical labeling was present outside the areas labeled by comparable injections in adult animals.(ABSTRACT TRUNCATED AT 400 WORDS)     

Efferent projections from temperature sensitive recording loci within the marginal zone of the nucleus caudalis of the spinal trigeminal complex in the cat

The efferent projections from nucleus caudalis of the spinal trigeminal complex in cats were studied with retrograde and anterograde axonal transport techniques combined with localization of recording sites in the thalamus and marginal zone of nucleus caudalis to innocuous skin cooling. Results showed brainstem projections from nucleus caudalis to rostral levels of the spinal trigeminal complex, to the ventral division of the principal trigeminal nucleus, the parabrachial nucleus, cranial motor nuclei 7 and 12, solitary complex, contralateral dorsal inferior olivary nucleus, portions of the lateral reticular formation, upper cervical spinal dorsal horn and, lateral cervical nucleus. Projections to the thalamus included: a dorsomedial region of VPM (bilaterally) and to the main part of VPM and PO contralaterally. Neuronal activity was recorded in the dorsomedial region of VPM to cooling the ipsilateral tongue. HRP injections in this thalamic region retrogradely labeled marginal neurons in nucleus caudalis. These results show that marginal neurons of nucleus caudalis provide a trigeminal equivalent of spinothalamic projections to the ventroposterior nucleus in cats.     

Brain stem projections to the facial nucleus of the rat

Horseradish peroxidase was injected into the medial and lateral columns of the facial nucleus of the rat. Following medial injections, cells were labelled by retrograde transport in the ipsilateral spinal trigeminal nucleus (caudalis) both medial vestibular nuclei, contralateral midbrain reticular formation and nucleus of the lateral lemniscus. The periaqueductal grey, interstitial nucleus and nucleus of Darkschewitch were also labelled ipsilaterally. Injections into the lateral column of the facial nucleus labelled the spinal trigeminal nucleus (oralis) and parabrachial nuclei ipsilaterally and the Darkschewitch and red nuclei contralaterally.     

Distinct lateral and medial projections of the spinohypothalamic tract of the rat

We recently described a direct nociceptive projection from the spinal cord to the hypothalamus in the rat. Several electrophysiological studies of this projection indicated that the axons of some spinohypothalamic tract neurons (SHT) reach the hypothalamus either by a lateral or by a medial route. The purpose of this study was to determine the origin of all SHT neurons that reach the hypothalamus through the lateral and the medial projections, and to investigate the possibility of ablating the SHT without damaging other important sensory and motor tracts by combining retrograde tracing techniques with axonal ablation. As compared with control cases, significant (P < .05) reductions in the number of labeled SHT neurons were encountered, 26% in the ipsilateral spinal cord following lesions of the medial projection, 67% in the contralateral spinal cord following lesions of the lateral projection, and 94% in both contra- and ipsilateral sides following lesions of both the medial and lateral projections. Bilateral lesions of the lateral projections had no effect on the distribution of labeled neurons in the spinal cord and dorsal column nuclei following injections of Fluoro-Gold (FG) into the thalamus, and a small unilateral lesion of the lateral projection reduced the ipsilateral labeling in the motor cortex following injections of FG into the pyramidal decussation. These findings suggest that most SHT neurons ascend through the contralateral lateral projection and that less than half continue in the medial projection to the ipsilateral side. They also suggest a site that can be lesioned without affecting other ascending sensory spinal pathways. © 1996 Wiley-Liss, Inc.     

Evidence for divergent projections to the brain noradrenergic system and the spinal parasympathetic system from Barrington's nucleus

The present study was designed to determine whether Barrington's nucleus, which lies ventromedial to the locus coeruleus (LC) and projects to the sacral parasympathetic nucleus, is a source of afferent projections to the LC. Restricted injections of the anterograde tracer, biocytin, into Barrington's nucleus labeled varicose fibers that extended from the injection site into the LC. Consistent with this, injections of the retrograde tracers, wheatgerm agglutinin conjugated to horseradish peroxidase coupled to gold particles (WGA-Au-HRP) or fluorescein-conjugated latex beads, into the LC labeled numerous (approximately 10%) Barrington's neurons that were also retrogradely labeled by Fluoro-Gold (FG) injections in the spinal cord. Retrograde tracing from the LC combined with corticotropin-releasing hormone (CRH) immunohistochemistry revealed that at least one third of the retrogradely labeled neurons in Barrington's nucleus were CRH-immunoreactive (CRH-IR). Finally, in triple labeling studies, CRH-Barrington's neurons were consistently observed that were retrogradely labeled from both the LC and spinal cord. These findings implicate Barrington's nucleus as an LC afferent and a source of CRH-IR fibers in the LC. Additionally, the results suggest that some Barrington's neurons diverge to innervate both the spinal cord and the LC. This divergent innervation may serve to coregulate the sacral parasympathetic nervous system and brain noradrenergic system, thus providing a mechanism for coordinating pelvic visceral functions with forebrain activity.     

Distribution and origin of corticotropin-releasing factor-immunoreactive axons in the female rat lumbosacral spinal cord.

Corticotropin-releasing factor (CRF) is a neuropeptide traditionally known for its hormonal role in the hypothalamic/pituitary/adrenal stress axis. However, CRF has been reported in axons in sites that may be considered outside of the direct stress axis, e.g., in axons in the lumbosacral spinal cord associated with the micturition response. Whether any of these CRF-immunoreactive axons interacts with uterine-related preganglionic autonomic neurons or projection neurons in the lumbosacral spinal cord is unknown. Thus, immunohistochemistry and retrograde tracing were employed to determine the presence, distribution, and origin of CRF-immunoreactive axons in the L6/S1 spinal cord of the female rat and to ascertain whether these axons are associated with uterine-related neurons. CRF-immunoreactive axons were present in the dorsal horn, medial and lateral collateral pathways, dorsal intermediate gray, laminae VlI and X, and sacral parasympathetic nucleus of the spinal cord. Nitric oxide-synthesizing, i.e., NADPH-d-positive neurons and pseudorabies virus labeled uterine-related neurons were in the sacral parasympathetic nucleus and were closely apposed by CRF-immunoreactive axons. Injection of retrograde tracers (fluorogold or fast blue) into the L6/S1 spinal cord labeled neurons in the hypothalamic paraventricular nucleus and pontine Barrington's nucleus, and some of these neurons were immunoreactive for CRF. This study demonstrates that CRF-immunoreactive axons are present in the L6/S1 spinal cord of the female rat in areas associated with sensory and autonomic processing. Some of these axons originate from the paraventricular nucleus and Barrington's nucleus and are adjacent to uterine-related neurons. These results indicate that CRF may influence neural activity related to the female reproductive system.     

Postsynaptic dorsal column pathway of the rat. III. Distribution of ascending afferent fibers

The distribution in the dorsal column nuclei (DCn) of post-synaptic dorsal column (PSDC) fibers was examined in rats following injections of Phaseolus vulgaris leucoagglutinin (PHA-L) in the spinal cord. Lemniscal neurons in the DCn were retrogradely labeled in the same animals by injecting the thalamus with Fluoro-Gold. In some experiments, primary afferent fibers were also labeled by injecting dorsal root ganglia with choleragenoid-conjugated HRP. Injections of PHA-L into the cervical enlargement labeled many fibers and varicosities throughout most of the ipsilateral cuneate nucleus. Labeled fibers were also present in the external cuneate and internal basilar nuclei. Injections of PHA-L into thoracic cord labeled fibers and varicosities in the medial cuneate and lateral gracile nuclei, as well as the external cuneate nucleus. Injections into the lumbar enlargement labeled fibers and varicosities throughout most of the gracile nucleus. Injections in sacral cord labeled fibers in the most medial part of the gracile nucleus. Dense labeling of PSDC fibers was found in areas with high densities of retrogradely labeled lemniscal neurons and areas with high densities of primary afferent fibers. In all regions of the DCn and in the external cuneate nucleus, fibers and varicosities labeled for PHA-L were seen in apposition to retrogradely labeled lemniscal cells. The distribution of postsynaptic afferent fibers in the DCn of the rat and its relationship to lemniscal neurons and primary afferent fibers contrast sharply with these features in cats.     

Central distribution of trigeminal and upper cervical primary afferents in the rat studied by anterograde transport of horseradish peroxidase conjugated to wheat germ agglutinin

Injections of WGA-HRP were made in the rat trigeminal ganglion and C1-3 dorsal root ganglia (DRGs) to study the central projection patterns and their relations to each other. Trigeminal ganglion injections resulted in heavy terminal labeling in all trigeminal sensory nuclei. Prominent labeling was also observed in the solitary tract nucleus and in the medial parts of the dorsal horn at C1-3 levels, but labeling could be followed caudally to the C7 segment. Contralateral trigeminal projections were found in the nucleus caudalis and in the dorsal horn at C1-3 levels. The C1 DRG was found to be inconstant in the rat. When it was present, small amounts of terminal labeling were found in the external cuneate nucleus (ECN) and the central cervical nucleus (CCN). No dorsal horn projections were seen from the C1 DRG. Injections in the C2 DRG resulted in heavy labeling in the ECN, nucleus X, CCN, and dorsal horn, where it was mainly located in lateral areas. Labeling could be followed caudally to the Th 7 segment. C2 DRG projections also appeared in the cuneate nucleus (Cun), in all the trigeminal sensory nuclei, and in the spinal, medial, and lateral vestibular nuclei. A small C2 DRG projection was observed in the ventral cochlear nucleus. C3 DRG injections resulted in heavy labeling in both medial middle and lateral parts of the dorsal horn, in the ECN, and in nucleus X, whereas the labeling in the CCN was somewhat weaker. Smaller projections were seen to trigeminal nuclei, Cun, and the column of Clarke. Comparisons of the central projection fields of trigeminal and upper cervical primary afferents indicated a somatotopic organization but with a certain degree of overlap.     

Central distribution of afferent pathways from the uterus of the cat.

Afferent pathways from the uterus of the cat were labeled by injections of horseradish peroxidase (HRP), wheat germ agglutinin-HRP, or fluorescent dyes into the uterine cervix and uterine horns. Afferent input to the uterus arises from small to medium size neurons (average size 31 x 28 microns) in dorsal root ganglia at many levels of the spinal cord (T12-S3). The segmental origin correlates with the location of the afferent terminal field in the uterus. Eighty-seven percent of the dorsal root ganglion cells (average, 822 on one side) innervating the cervix are located in sacral ganglia, whereas 97% of the cells innervating the uterine horn (average 479 on one side) are located in lumbar ganglia. Double dye labeling experiments indicate that a small percentage (average 15%) of lumbar neurons innervating the uterine cervix also innervate the uterine horn. The majority (70-80%) of afferent input to the uterine cervix passes through the pelvic nerve and the remainder through the pudendal nerve, whereas afferent input to the uterine horn must travel in sympathetic nerves. Ovariectomy (10-14 days) did not change significantly the number, sizes, or segmental distribution of uterine afferent neurons. In some cats (25%) injections of WGA-HRP into the uterine cervix labeled neurons (90-125 per animal) in lamina VII in the S2 spinal segment in the region of the sacral parasympathetic nucleus. Central projections of uterine horn afferent neurons were not labeled; however, afferent projections from the cervix were detected in the sacral spinal cord. The most prominent labeling was present in Lissauer's tract and in lamina I and outer lamina II on the lateral edge of the dorsal horn. From this region some labeled axons extended through lamina V into the dorsal gray commissure. Very few afferents were labeled on the medial side of the dorsal horn. These results are discussed in regard to the physiological function of uterine afferents and the possible transmitter role of vasoactive intestinal polypeptide, which is present in a large percentage (70%) of cervical afferent neurons.