Central distribution of cervical primary afferents in the rat, with emphasis on proprioceptive projections to vestibular, perihypoglossal, and upper thoracic spinal nuclei

The projections of primary afferents from rostral cervical segments to the brainstem and the spinal cord of the rat were investigated by using anterograde and transganglionic transport techniques. Projections from whole spinal ganglia were compared with those from single nerves carrying only exteroceptive or proprioceptive fibers. Injections of horseradish peroxidase (HRP) or wheat germ agglutinin-horseradish peroxidase conjugate (WGA-HRP) were performed into dorsal root ganglia C2, C3, and C4. Free HRP was applied to the cut dorsal rami C2 and C3, greater occipital nerve, sternomastoid nerve, and to the C1/2 anastomosis, which contains afferents from suboccipital muscles and the atlanto-occipital joint. WGA-HRP injections into ganglia C7 and L5 were performed for comparative purposes. Injections of WGA-HRP or free HRP into rostral cervical dorsal root ganglia and HRP application to C2 and C3 dorsal rami produced labeling in dorsal and ventral horns at the level of entrance, the central cervical nucleus, and in external and main cuneate nuclei. From axons ascending to pontine and descending to upper thoracic spinal levels, medial collaterals were distributed to medial and descending vestibular, perihypoglossal and solitary nuclei, and the intermediate zone and Clarke's nucleus dorsalis in the spinal cord. Lateral collaterals projected mainly to the trigeminal subnucleus interpolaris and to lateral spinal laminae IV and V. Results from HRP application to single peripheral nerves indicated that medial collaterals were almost exclusively proprioceptive, whereas lateral collaterals were largely exteroceptive with a contribution from suboccipital proprioceptive fibers. WGA-HRP injections into dorsal root ganglia C7 and L5 failed to produce significant labeling within vestibular and periphypoglossal nuclei, although they demonstrated classical projection sites within the brainstem and spinal cord. The consistent collateralisation pattern of rostral cervical afferents along their whole rostrocaudal course enables them to contact a variety of precerebellar, vestibulospinal, and preoculomotor neurons. These connections reflect the well-known significance of proprioceptive neck afferents for the control of posture, head position, and eye movements.     

Organization of spinal inputs to the perihypoglossal complex in the cat

First- and second-order spinal afferents to the perihypoglossal complex were sought by using axonal transport of WGA-HRP. Injections in C1, 2, and 3 dorsal root ganglia resulted in axonal labeling in the nucleus intercalatus and the external cuneate nucleus, with a number of retrogradely labeled cells seen as well in the latter. A similar pattern of axonal labeling in the nucleus intercalatus as well as several retrogradely labeled cells were found after spinal cord injections at levels C1, 2, and 3. A prominent field of labeled axons was also present in the rostral main cuneate nucleus. No labeling was seen in the perihypoglossal nuclei after injections in the spinal cord or dorsal root ganglia at levels caudal to C3. After injections of HRP into the perihypoglossal nucleus we were able to identify labeled neurons within Rexed's laminae V-VIII and the central cervical nucleus. Anterograde labeling in the main cuneate nucleus was observed after C1 to C5 ganglion and C1 to C6 cord injections. The pattern and extent of labeling in the perihypoglossal nuclei and adjacent structures seen after cerebellar injections into lobules V and VI were comparable to those previously reported and permitted evaluation of the relay from dorsal root ganglia through the intercalatus to the vermis. Topography of the cervical projections to the nucleus intercalatus is considered with respect to that of the perihypoglossal-collicular projection. A discussion is offered of the apparent importance of nucleus intercalatus as a relay of cervical and vestibular afferent information to premotor structures involved in neck motor control. The perihypoglossal complex is viewed as being organized in such a fashion as to allow the nuclei intercalatus and prepositus hypoglossi to function as key structures in the integration of inputs related to neck and ocular motor control, respectively.     

Brainstem terminations of extraocular muscle primary afferent neurons in the monkey

The central terminations of afferent nerve fibers from the extraocular muscles of the monkey were investigated by means of transganglionic transport of wheat germ agglutinin-conjugated horseradish peroxidase (WGA/HRP). Following injections of selected extraocular muscles with WGA/HRP, terminal labeling was apparent in the ipsilateral trigeminal sensory and cuneate nuclei. The density of trigeminal projections varied markedly from one rostrocaudal level to the next, being heaviest within the ventrolateral portion of pars interpolaris of the spinal trigeminal nucleus. A second extraocular muscle afferent representation was noted in ventrolateral portions of the cuneate nucleus. This projection was restricted to rostral portions of pars triangularis of the cuneate nucleus, partially overlapping the afferent termination from dorsal neck muscles. It is likely that some of the problems encountered in formulating conclusions regarding the functional role of extraocular muscle proprioception are due to a lack of detailed information of the central termination pattern of muscle afferents. Taken together, the present findings should provide a basis for further anatomical and physiological studies designed to elucidate the role played by extraocular muscle proprioceptors in vision and oculomotor control.     

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.     

Afferent fibers in the hypoglossal nerve: a horseradish peroxidase study in the cat

The existence of afferent fibers in the cat hypoglossal nerve was studied by transganglionic transport of horseradish peroxidase (HRP). Injections of wheat germ agglutinin-conjugated HRP (WGA-HRP) into the hypoglossal nerve resulted in some retrograde labeling of cell bodies within the superior ganglia of the ipsilateral glossopharyngeal and vagal nerves. A few labeled cell bodies were also present ipsilaterally within the inferior ganglion of the vagal nerve and the spinal ganglion of the C1 segment. Some of the labeled glossopharyngeal and vagal fibers reached the nucleus of the solitary tract by crossing the dorsal portion of the spinal trigeminal tract. Others distributed to the spinal trigeminal nucleus pars interpolaris and to the ventrolateral part of the medial cuneate nucleus by descending through the dorsal portion of the spinal trigeminal tract. In the spinal cord these descending fibers, intermingling with labeled dorsal root fibers, distributed to laminae I, IV-V and VII-VIII of the C1 and C2 segments. Additional HRP experiments revealed that the fibers in laminae VII-VIII originate mainly from dorsal root of the C1 segment.     

The central projections of the great auricular nerve primary afferent fibers--an HRP transganglionic tracing method

A study of the central distribution of the primary afferent fibers of the great auricular nerve (GAN) was made in 18 rabbits by means of transganglionic transport of horseradish peroxidase (HRP). HRP applied to the cut central end of the GAN was detected ipsilaterally in the dorsal root ganglion cells (segments C2-C3) and the superior cervical ganglion cells. The transganglionically labeled fibers were seen in the dorsal column of the upper 4 cervical segments and in the cranial nerve nuclei of the medulla oblongata. The afferent projections were rather strong in the regions of laminae I-V of C2, the caudal subnucleus of the nucleus of the spinal tract of the trigeminal nerve (NVSpc), the solitary nucleus (SN), the medial and lateral cuneate nuclei, etc. The results showed that the primary afferent impulses of the GAN and the peripheral nerves which supply the head, face, trunk and viscera might converge on the upper cervical cord, the NVSpc and the SN, and play a certain role of modulation on the transmission of somaticovisceral sensations, especially pain.     

Immunohistochemical localization of calretinin in the rat hindbrain

The localization of calretinin in the rat hindbrain was examined immunohistochemically with antiserum against calretinin purified from the guinea pig brain. Calretinin immunoreactivity was found within neuronal elements. The distribution of calretinin-immunoreactive cell bodies and fibers is presented in schematic drawings and summarized in a table. Major calretinin-immunoreactive neurons were found in the lateral and medial geniculate nuclei, substantia nigra, ventral tegmental area, interpeduncular nucleus, periaqueductal gray, mesencephalic trigeminal nucleus, superior and inferior colliculi, pontine nuclei, parabrachial nucleus, dorsal and laterodorsal tegmental nuclei, cochlear nuclei, vestibular nuclei, medullary reticular nuclei, nucleus of the solitary tract, area postrema, substantia gelatinosa of the spinal trigeminal nucleus, and cerebellum. These results show that distinct calretinin-immunoreactive neurons are widely distributed in the rat hindbrain.     

Immunocytochemical localization of the choline acetyltransferase containing neuron system in the rat lower brain stem

This distribution of choline acetyltransferase (CHAT) immunoreactivity (CHAT-I) in the rat lower brain stem was analyzed using a highly sensitive avidin-biotin immunocytochemical method and 3-amino-9-ethyl-carbazole visualization. A much wider and more abundant distribution of CHAT-I structures in the lower brain stem was demonstrated than in earlier studies. The following areas were newly identified as areas rich in CHAT-I fibers: the interpeduncular nucleus, medial geniculate body, central gray matter of pons, pontine nucleus, parabigeminal nucleus, dorsal tegmental nucleus of Gudden, lateral trapezoid nucleus, inferior colliculus, dorsal and ventral cochlear nuclei, medial and lateral vestibular nuclei, reticular formation of medulla oblongata, and gelatinosa of caudal trigeminal spinal tract nucleus. In addition to the areas in which they have been known to exist, CHAT-I perikarya were found in the caudal portion of substantia nigra pars reticulata, the area between trigeminal motor nucleus and superior olivary nucleus, the medial and spinal vestibular nucleus, prepositus hypoglossal nucleus, raphe magnus and obscurus, ventromedial portion of solitary tract nucleus and its just ventral reticular formation, and caudal trigeminal spinal tract nucleus.     

Trigeminal primary afferent projections to "non-trigeminal" areas of the rat central nervous system.

The central projections of rat trigeminal primary afferent neurons to various "non-trigeminal" areas of the central nervous system were examined by labeling the fibers with wheat germ agglutinin-horseradish peroxidase (WGA-HRP) transported anterogradely from the trigeminal ganglion. This technique produced a clear and comprehensive picture of trigeminal primary afferent connectivity that was in many ways superior to that which may be obtained by using degeneration, autoradiography, cobalt labeling, or HRP transganglionic transport techniques. Strong terminal labeling was observed in all four rostrocaudal subdivisions of the trigeminal brainstem nuclear complex, as well as in the dorsal horn of the cervical spinal cord bilaterally, numerous brainstem nuclei, and in the cerebellum. Labeling in the ipsilateral dorsal horn of the cervical spinal cord was very dense at C1, moderately dense at C2 and C3, and sparse at C4-C7. Numerous fibers crossed the midline in the medulla and upper cervical spinal cord and terminated in the contralateral pars caudalis and dorsal horn of the spinal cord from C1-C5. The latter axons terminated most heavily in the mandibular and ophthalmic regions of the contralateral side. Extremely dense terminal labeling was observed in the ipsilateral paratrigeminal nucleus and the nucleus of the solitary tract, moderate labeling was seen in the supratrigeminal nucleus and in the dorsal reticular formation, and small numbers of fibers were observed in the cuneate, trigeminal motor, lateral and superior vestibular nuclei, and in the cerebellum. The latter fibers entered the cerebellum in the superior cerebellar peduncle and projected to the posterior and anterior lobes as well as to the interposed and lateral deep cerebellar nuclei. Most projections in this study originated from fibers in the dorsal part of the spinal tract of V, suggesting a predominantly mandibular origin for these fibers. Projections from the ophthalmic and maxillary divisions, in contrast, were directed mainly to the cervical spinal cord bilaterally, to contralateral pars caudalis, and to certain areas of the reticular formation. In conclusion, this study has demonstrated that somatosensory information from the head and face may be transmitted directly to widespread and functionally heterogeneous areas of the rat central nervous system, including the spinal cord dorsal horn, numerous brainstem nuclei, and the cerebellum.(ABSTRACT TRUNCATED AT 400 WORDS)     

Primary afferent projections from the upper respiratory tract in the muskrat.

The central projections of the ethmoidal, glossopharyngeal, and superior laryngeal nerves were determined in the muskrat by use of the transganglionic transport of a mixture of horseradish peroxidase (HRP) and wheat germ agglutinin (WGA)-HRP. The ethmoidal nerve projected to discrete areas in all subdivisions of the ipsilateral trigeminal sensory complex. Reaction product was focused in ventromedial portions of the principal nucleus, subnucleus oralis, and subnucleus interpolaris. The subnucleus oralis also contained sparse reaction product in its dorsomedial part. Projections were dense to ventrolateral parts of laminae I and II of the rostral medullary dorsal horn, with sparser projections to lamina V. Label in laminae I and V extended into the cervical dorsal horn. A few labeled fibers were followed to the contralateral dorsal horn. The interstitial neuropil of the ventral paratrigeminal nucleus was densely labeled. Extratrigeminal primary afferent projections in ethmoidal nerve cases involved the K?lliker-Fuse nucleus and ventrolateral part of the parabrachial nucleus, the reticular formation surrounding the rostral ambiguous complex, and the dorsal reticular formation of the closed medulla. Retrograde labeling in the brain was observed in only the mesencephalic trigeminal nucleus in these cases. The cervical trunk of the glossopharyngeal and superior laryngeal nerves also projected to the trigeminal sensory complex, but almost exclusively to its caudal parts. These nerves terminated in the dorsal and ventral paratrigeminal nuclei as well as lamina I of the medullary and cervical dorsal horns. Lamina V received sparse projections. The glossopharyngeal and superior laryngeal nerves projected to the ipsilateral solitary complex at all levels extending from the caudal facial nucleus to the cervical spinal cord. At the level of the obex, these nerves projected densely to ipsilateral areas ventral and ventromedial to the solitary tract. Additional ipsilateral projections were observed along the dorsolateral border of the solitary complex. Near the obex and caudally, the commissural area was labeled bilaterally. Labeled fibers from the solitary tract projected into the caudal reticular formation bilaterally, especially when the cervical trunk of the glossopharyngeal nerve received tracer. Labeled fibers descending further in the solitary tract gradually shifted toward the base of the cervical dorsal horn. The labeled fibers left the solitary tract and entered the spinal trigeminal tract at these levels. Retrogradely labeled cells were observed in the ambiguous complex, especially rostrally, and in the rostral dorsal vagal nucleus after application of HRP and WGA-HRP to either the glossopharyngeal or superior laryngeal nerves. In glossopharyngeal nerve cases, retrogradely labeled neurons also were seen in the inferior salivatory nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Central projections from cat suboccipital muscles: a study using transganglionic transport of horseradish peroxidase

Central projections of suboccipital muscle nerves were examined following exposure of cut peripheral nerves to the tracer horseradish peroxidase. Labelled fibers entered the C1 and C2 dorsal roots and accumulated in the dorsolateral part of the dorsal funiculus. Many labelled fibers entered the grey matter of C1 to C3 in ventrally directed bundles which passed medially to the base of the dorsal horn. No terminal labelling was apparent in superficial layers of the dorsal horn. However, labelled fibers ramified extensively throughout medial parts of the intermediate laminae, in and around the central cervical nucleus. Labelled fibers also projected toward the ventral horn. In cats subjected to ventral root section at the time of peripheral nerve exposure, a modest distribution of reaction product was observed deep in the ventral horn. In cats which did not undergo ventral root section, anterograde projections in the ventral horn were obscured by the simultaneous retrograde filling of motoneurons both in the ventromedial nucleus and on the medial and lateral borders of the gray matter. Labelled axons also coursed rostrally into the medulla where they formed a circumscribed bundle between the main cuneate nucleus and the spinal nucleus of V. Three consistent regions of HRP deposition could be identified at medullary levels. Dense accumulations of reaction product were present in circumscribed regions of the external cuneate nucleus (ECN) throughout its rostrocaudal extent. A second zone of dense labelling occurred in the intermediate nucleus of Cajal, where it appeared to form a continuing column rostral to the central cervical nucleus in C1-C3. Sparse labelling was restricted to a third zone in the ventrolateral part of the main cuneate nucleus.     

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.     

The pattern of spinal and medullary projections from a cutaneous nerve and a muscle nerve of the forelimb of the cat: a study using the transganglionic transport of HRP

The transport of HRP into the spinal cord and medulla in the cat has been examined from a forelimb cutaneous nerve, the lateral superficial radial nerve (LSR), and from the muscle nerves supplying both heads of the forelimb muscle, extensor carpi radialis (ECR). HRP transported by the LSR was widely distributed in the spinal cord throughout laminae I-IV in the vicinity of the root entry zone and from spinal segments T1 to C5. HRP was also transported from the LSR to the medulla where there was intense patchy, discontinuous labelling in the main cuneate nucleus. The pattern of labelling in the cuneate nucleus did not follow any simple somatotopic plan. Exposure of the muscle nerve to HRP led to labelling in the spinal dorsal horn in lamina I, in the deep dorsal horn on the lamina V/VI border, and in lateral and medial lamina VI at sites that contain cells of origin of spinocerebellar tracts. The medial lamina VI label was contiguous with a deposit that extended medially to the central canal. The label in lateral lamina VI was patchy and formed a discontinuous column from T1 to C5. HRP transported by the muscle nerve also produced label in the more ventral regions of the cuneate nucleus where it had a lacy appearance, in part due to its extensive distribution around dendrites. A relatively dense, patchy, and discontinuous deposit of reaction product was also present in the external cuneate nucleus after muscle nerve exposure. This deposit was most intense on the dorsomedial surface of this nucleus, but another, less intense, deposit was also present ventrally.     

Localization of preproenkephalin mRNA in the rat brain and spinal cord by in situ hybridization

To determine the localization in rat brain and spinal cord of individual neurons that contain the messenger RNA coding for the opioid peptide precursor preproenkephalin, we performed in situ hybridization with a tritiated cDNA probe complementary to a protion of preproenkephalin mRNA. We observed autoradiographic signal over the cytoplasm of neurons of many regions of the central nervous system. Several types of controls indicated specificity of the labeling. Neurons containing preproenkephalin mRNA were found in the piriform cortex, ventral tenia tecta, several regions of the neocortex, nucleus accumbens, olfactory tubercle, caudate-putamen, lateral septum, bed nucleus of the stria terminalis, diagonal band of Broca, preoptic area, amygdala (especially central nucleus, with fewer labeled neurons in all other nuclei), hippocampal formation, anterior hypothalamic nucleus, perifornical region, lateral hypothalamus, paraventricular nucleus, dorsomedial and ventromedial hypothalamic nuclei, arcuate nucleus, dorsal and ventral premamillary nuclei, medial mamillary nucleus, lateral geniculate nucleus, zona incerta, periaqueductal gray, midbrain reticular formation, ventral tegmental area of Tsai, inferior colliculus, dorsal and ventral tegmental nuclei of Gudden, dorsal and ventral parabrachial nuclei, pontine and medullary reticular formation, several portions of the raphe nuclei, nucleus of the solitary tract, nucleus of the spinal trigeminal tract (especially substantia gelatinosa), ventral and dorsal cochlear nuclei, medial and spinal vestibular nuclei, cuneate and external cuneate nuclei, gracile nucleus, superior olive, nucleus of the trapezoid body, some deep cerebellar nuclei, Golgi neurons in the cerebellum, and most laminae of the spinal cord. In most of these brain regions, the present results indicate that many more neurons contain preproenkephalin mRNA than have been appreciated previously on the basis of immunocytochemistry.     

Central projections of C4-C8 dorsal root ganglia in the rat studied by anterograde transport of WGA-HRP.

Injections of WGA-HRP were made in the rat C4-C8 dorsal root ganglia (DRGs) individually to study the central projections and their relations to each other. The main dorsal horn projections from these DRGs to the dorsal horn lamina II extended for about two segments rostrally and caudally to the injected DRG, whereas the projections to laminae I, III, and IV were less restricted rostrocaudally. Comparisons of the dorsal horn projections from the DRGs investigated indicated a tendency for a somatotopic organization, which was most prominent in lamina II. Labeled central branches from the C4-8 DRGs could be traced in the dorsal column as far caudally as 12-17 segments caudal to the level of entrance. Most of these fibers appeared to end in the medial dorsal horn base, including the column of Clarke. Labeling of primary afferents in the ventral horn generally extended for at least 3-4 segments rostral and caudal to the level of the injected DRG. Projections to the central cervical nucleus were most prominent from the C4 DRG and gradually became less prominent from the more caudal DRGs. Heavy projections to the cuneate nucleus (Cun) originated from the C7 and C8 DRG, whereas those from the C4-C6 DRGs were less extensive. The Cun projections from the different DRGs appeared to overlap, and the same was true for the projections to the external cuneate nucleus. Projections to the gracile nucleus, the vestibular nuclear complex, including nucleus X, and to trigeminal sensory nuclei were seen from all DRGs investigated.     

The origin of brainstem-spinal pathways in the north american opossum (didelphis virginiana). Studies using the horseradish peroxidase method

Brainstem neurons which project to lumbar, thoracic and cervical spinal levels have been identified in the North American opossum by the horseradish peroxidase (HRP) technique. Neurons which relay to all of the levels studied are located within the medullary and pontine reticular formation as well as within the nucleus cuneatus, the nucleus of the tractus solitarius, the lateral reticular nucleus, the medullary and caudal pontine raphe nuclei, the lateral, medial and inferior vestibular nuclei, the nucleus “F,” the nucleus coeruleus, and the nucleus coeruleus, para α, the red nucleus, and the interstitial nucleus of Cajal. The lateral vestibulospinal and rubrospinal systems are topographically organized, although neurons projecting to different cord levels show considerable intermingling. Our material also provides evidence that raphe-spinal and reticulospinal connections are organized to some degree. Neurons which backfill after cervical and thoracic placements, but not after lumbar injections, are distributed within the caudal spinal trigeminal nucleus, the nucleus intercalatus, the dorsal vagal nucleus, the cuneiform area of Castaldi, the fields of Forel, and the nucleus of Darkschewitsch. Reactive neurons are present within the lateral, dorsal and posterior hypothalamic areas as well as within the periventricular and paraventricular nuclei after thoracic placements and within the superior colliculus after injections within the cervical cord. Additionally, neurons are reactive in the nucleus ambiguus, the interpolar division of the spinal trigeminal nucleus and the rostral division of the oculomotor nucleus (Oswaldo-Cruz and Rocha-Miranda, '68) after HRP placements into the third and fourth cervical segments.     

An HRP study of the central course of sensory intermediate and vagal fibers in peripheral facial nerve branches in the cat

Previous studies have shown that sensory fibers of intermediate and vagal nerve origin are present in facial nerve branches to the mimetic muscles in the cat. In the present study the central course of these fibers has been examined by transganglionic transport of horseradish peroxidase (HRP). In some of these experiments the facial nerve proper was transected central to the site of HRP application. In this way, the central course of the vagal fibers could be studied separately. For comparison HRP-conjugated wheat germ agglutinin was injected into the geniculate ganglion, revealing the central course of the entire afferent component of the intermediate nerve. The results show that labeled sensory intermediate nerve fibers, at their level of entrance in the brainstem, form a tract at the dorsal margin of the spinal trigeminal tract (5T). While some fibers ascend from this level to terminate in the main sensory trigeminal nucleus, and a few fibers terminate in the rostral part of the solitary tract nucleus, the majority take a descending course. The main site of termination for these descending fibers is in the medial part of the C2 dorsal horn. Terminal labeling is also seen in the ventrolateral part of the cuneate nucleus (CUN) and in a small area of gray substance between CUN and trigeminal nucleus caudalis. After entering the brainstem some sensory vagal fibers project to the trigeminal nucleus interpolaris and to an interstitial nucleus within the 5T, but the larger part joins the descending tract of intermediate nerve fibers. These descending vagal fibers have a terminal distribution pattern similar to the intermediate nerve fibers.