The mormyrid brainstem. I. Distribution of brainstem neurones projecting to the spinal cord in Gnathonemus petersii. An HRP study

The sources of the descending spinal tracts were identified in the teleost fish Gnathonemus petersii by retrograde HRP transport. HRP injections were made at two spinal levels, either at level of the caudal end of the dorsal fin, anterior to the electric organ, or at the pectoral fin. In both cases all labeled cells were found in the rhombencephalon and the mesencephalic tegmentum. No labeled cells were observed either in the cerebellum and lateral line lobes or in the dorsal mesencephalon i.e. torus semicircularis and mesencephalic tectum or in the telencephalon. Following caudal spinal injections, the majority of the labeled cells were grouped in a median and a ventrolateral column of the rhombencephalic reticular formation. The latter is composed of three parts corresponding to the nucleus reticularis inferior, medius and superior. Both Mauthner cells, all the cells in the medullary relay nucleus controlling the electric organ discharge and a few cells in the posterior part of the magnocellular octaval nucleus were labeled. In the mesencephalon, four nuclei were identified by HRP labeling: the nucleus of the medial longitudinal fasciculus, the nucleus reticularis mesencephali and the anterior and posterior tegmental mesodiencephalic nuclei. The rostral injections revealed several additional spinal projections from the descending vestibular and tangential nuclei, from the medial part of the magnocellular nucleus and, finally, from the rostral periventricular gray of the mesencephalon. Also, after such injections, a greater number of cells were labeled in the reticular formation, especially in the median column and in the inferior reticular nucleus. The results suggest that the rostral spinal cord has a larger connection with the acoustico-vestibular area and the medullary reticular formation than the caudal spinal cord. In contrast, the mesencephalic nuclei, probably linked to the mesencephalic tectum and the pretectal area, appears to be a coordinating apparatus between the visual system and the trunk/tail musculature. Thus, it appears that teleost fish possess the same basic equipment of descending spinal pathways as higher vertebrates.     

Spinovestibular projections in the rat,with particular reference to projections from the central cervical nucleus to the lateral vestibular nucleus

Projections from the spinal cord to the vestibular nuclei were examined following injections of Phaseolus vulgaris–leucoagglutinin, cholera toxin subunit B, or biotinylated dextran at various levels of the spinal cord in the rat. Labeled terminals were abundant after injections of the tracers into the C2 and C3 segments containing the central cervical nucleus. Labeled terminals were seen in the descending vestibular nucleus and the parvocellular, magnocellular, and caudal parts of the medial vestibular nucleus throughout its rostrocaudal extent. Labeled terminals were most numerous in the lateral vestibular nucleus throughout its rostrocaudal extent. The projections from the central cervical nucleus to the vestibular nuclei were exclusively contralateral to the cells of origin because the axons of the central cervical nucleus neurons cross in the spinal cord. Following tracer injections in the cervical enlargement, many labeled terminals were seen in the magnocellular part of the medial vestibular nucleus, but a few were seen in the lateral and the descending vestibular nucleus. Injections into more caudal segments resulted in sporadic terminal labeling in the magnocellular part of the medial vestibular nucleus, the descending vestibular nucleus, and the caudal part of the lateral vestibular nucleus. The results indicate that primary neck afferent input relayed at the central cervical nucleus is mediated directly to the contralateral vestibular nuclei. It is suggested that this projection serves as an important linkage from the upper cervical segments to the lateral vestibulospinal tract in the tonic neck reflex. © 1995 Wiley-Liss, Inc.     

Morphology of the cells of origin of descending pathways to the spinal cord in Rana esculenta. A tracing study using cobaltic-lysine complex

The localization and morphological characteristics of neurons projecting to the spinal cord were studied with cobalt-filling technique. Cobaltic-lysine complex was iontophorized into descending pathways in the lumbar and cervical intumescence of the spinal cord and in some parts of the rhombencephalon. Well-filled cells located mainly in the rhombencephalon and telencephalon could be characterised as fusiform, triangular, multipolar and irregular neurons. Piriform and pyramidal cells predominated among projection neurons in the mesencephalon. Bilateral descending spinal pathways originate from the reticular nuclei of the rhombencephalon, nucleus vestibularis lateralis, nucl. anterodorsalis- and anteroventralis mesencephali, nucl. posterior thalami, nucl. ventralis hypothalami, area preoptica anterior and the striatum ventrale. Crossed pathways descend from the nucl. vestibularis descendens, nucl. tractus solitarii, nucl. cerebelli and the nucleus ruber. Uncrossed fibres originate from the nucl. tractus spinalis nervi trigemini, nucl. posteroventralis tegmenti, nucl. profundus mesencephali, nucl. fasciculi longitudinalis medialis and the nucleus ventrolateralis thalami. The organization of the frog's descending pathways is very similar to those in reptiles and in many ways to those in mammals. The possible synaptic connections of projection neurons have been discussed.     

Developmental sequence in the origin of descending spinal pathways. Studies using retrograde transport techniques in the North American opossum (Didelphis virginiana)

The origin of descending pathways to thoracic and cervical levels of the spinal cord has been investigated with retrograde tracing techniques in a series of pouch young and adult opossums. The opossum was chosen because it is born in a very immature state, 12-13 days after conception, and has a protracted development in an external pouch. A few neurons in the pontine reticular formation and nucleus coeruleus were labeled by horseradish peroxidase (HRP) injections of the thoracic cord as early as postnatal day (PND) 3. By PND 5, similar injections labeled neurons in the same areas as well as in the medullary reticular formation, the raphe nuclei of the caudal pons and medulla, the spinal trigeminal nuclei, the vestibular complex, the accessory oculomotor nuclei and the interstitial nucleus of Cajal. When Nuclear Yellow (NY) was employed, neurons were also labeled in the red nucleus, the hypothalamus and possibly in the nucleus of the solitary tract. Regardless of the technique employed, neurons in the dorsal column nuclei were not labeled by thoracic injections until at least PND 14. Axons from the nucleus ambiguus, the fastigial and interposed nuclei of the cerebellum as well as the intermediate and deep layers of the superior colliculus reach cervical levels of the cord, where they are specifically targeted, by at least PND 17. They do not significantly overgrow those levels during development. Corticospinal axons are the last of the major descending pathways to innervate the spinal cord. Cortical neurons cannot be labeled by cervical injections of either HRP or NY until at least PND 30. Evidence for transient brainstem-spinal and corticospinal projections was obtained.     

Topographic organization of descending projection neurons to the spinal cord of the goldfish, Carassius auratus

Several neurons from different nuclei give rise to descending spinal tracts and project to various levels in the spinal cord of goldfish, Carassius auratus. These were visualized by retrograde transport of horseradish peroxidase (HRP) administered to the bilaterally transected spinal cord at 6 levels, corresponding to 1st, 5th, 10th, 15th, 20th and 25gh segments of the vertebral column. As many as 16 brain nuclei or neuronal aggregations and the Mauthner cells project posteriorly up to the 20th spinal segment. Restricted neurons of the dorsolateral area in the nucleus preopticus magnocellularis and those of the ventromedial nucleus of the thalamus projected up to the 20th and 25th segments respectively. In the mesencephalon, the nucleus ruber and the nucleus of the medial longitudinal fasciculus revealed retrogradely labeled somata; the former extended up to the 20th segment, while the latter projected up to the 25th segment. The remaining descending projected neurons of the brain belonged to the rhombencephalon. The nucleus of the lateral lemniscus; anterior, magnocellular, descending and posterior divisions of the octaval nucleus: raphe nucleus; Mauthner cell and the neurons located adjacent to the trigeminal tract and those in the vicinity of the secondary gustatory tract sent their processes up to 20th segmental level. However, somata of the superior, medial and inferior divisions of the reticular nucleus and restricted neurons of the facial lobe extended up to 25th segmental level. The pattern of neuronal projections into the spinal cord suggests a topographic organization.     

The location of spinal projection neurons in the cerebellar nuclei (cerebellospinal tract neurons) of the cat. A study with the horseradish peroxidase technique

The distribution of spinal projections neurons was studied in the cerebellar nuclei of the cat following injections of horseradish peroxidase (HRP) into the cervical, thoracic and lumbar cord. HRP-positive (labeled) neurons were found in the medial (fastigial) and the posterior interpositus nuclei on the side contralateral to the cervical injections, being most numerous in cases with injections between the C2 and the C3 segments.In the medial nucleus (M) labeled neurons were distributed in the central to the caudal portions, and there was a conspicuous group of labeled small neurons extending from the ventolateral part to the intermediate zone between the M and the anterior interpositus nucleus. With an increasing number of medium-sized neurons, this neuronal group persisted caudally in a similar position, ventromedial to the posterior interpositus nucleus (IP). Labeled large neurons were seen in the medial third of the IP. In the two cases labeled neurons of medium and small sizes were equal iin number, and the neurons of the IP constituted about 10% of the total number of the spinal projection neurons. The present study suggests that the neurons of the M and the IP, including those of the intermediate group located between the two, project the bulk of the crossed descending fibers as far caudally as the C2 and the C3 segments.     

Localization of met-enkephalin-like immunoreactivity within pain-related nuclei of cervical spinal cord, brainstem and midbrain in the cat

Met-enkephalin immunoreactivity was investigated with an indirect immunoperoxidase technique in the cervical spinal cord, brainstem and midbrain of the cat, paying special attention to pain-related nuclei. Different technical conditions were used to reveal preferentially met-enkephalin-containing fibres and terminals or perikarya. Immunoreactive fibres and terminals were revealed optimally in sections from control animals incubated with detergent (Triton X-100). Immunoreactive perikarya were revealed in colchicine treated animals. Comparison between different routes of administration showed that local injections of colchicine are needed to reveal optimally immunoreactive perikarya in nuclei located far from the ventricles. Met-enkephalin-containing fibres and terminals are widely distributed in the posterior brain and spinal cord. The densest network of immunoreactive fibres are observed in the superficial layers of the cervical spinal cord and the caudal trigeminal nucleus, in the nucleus of the solitary tract, the nucleus of the facial nerve, the nucleus of the prepositus hypoglossi, the nucleus raphe pallidus, the medial vestibular nucleus, the interpedoncular nucleus and the substantia nigra. A moderate staining of fibres is observed in various nuclei including the ventral horn of the spinal cord and caudal trigeminal nucleus, the brainstem and midbrain reticular formation, the inferior olivary complex, the nucleus of the descending trigeminal tract and the periaqueductal grey. Met-enkephalin-containing perikarya are present in all the nuclei cited before, except in the inferior olivary complex. The densest aggregation of enkephalin-like perikarya is observed in the nucleus raphe magnus, nucleus raphe obscurus, nucleus raphe pallidus, nucleus reticularis gigantocellularis pars α and nucleus reticularis lateralis. The general distribution of enkephalin-containing structures in the cervical spinal cord, brainstem and midbrain of the cat appears very similar to that of the rat except in the substantia nigra where met-enkephalin cell bodies are found in the cat but not in the rat. In particular the pain-related nuclei present a similar distribution of the peptide in the two species; however, met-enkephalin-containing cell bodies are much more numerous in the cat than in the rat (notably in the reticular formation). Similar types of metenkephalin innervation occur in the dorsal and intermediate grey of the spinal cord and of the caudal trigeminal nucleus supporting further that the functional organizations of these regions are closely related.     

Calbindin-D28k immunoreactivity in the spinal cord of Xenopus laevis and its participation in ascending and descending projections

Immunohistochemistry for calbindin-D28k (CB) revealed that the spinal cord of Xenopus laevis possess a large number of CB-containing neurons widely distributed in both the dorsal and ventral horns, including areas which possess long ascending projections to supraspinal structures. In addition, the presence of CB-immunoreactive axons in the spinal funiculi suggested that descending projections containing this calcium binding protein may originate in different brainstem nuclei. Apart from mapping CB-containing elements in the spinal cord, a double labeling approach was used that combined the retrograde transport of dextran amines with CB immunohistochemistry. Thus, dextran amine injections into the lateral reticular region of the rhombencephalon, the parabrachial region, the mesencephalon and the dorsal thalamus revealed many retrogradely labeled cells in the spinal cord, a few number of which were double labeled for CB and found in the superficial dorsal horn and in the ventral medial region of the ventral horn. Their axons passed mainly via the lateral funiculus. Tracer application into the cervical spinal cord, combined with CB immunohistochemistry, resulted in retrogradely labeled cells throughout the brain, five groups of which showed CB immunoreactivity: (1) the mesencephalic trigeminal nucleus, (2) the laterodorsal tegmental nucleus, (3) the raphe nucleus, (4) the middle reticular nucleus and (5) the inferior reticular nucleus. The presence of CB in spinal pathways suggests that CB may play a role in controlling spinal cells, mainly subserving visceroceptive and nociceptive information to supraspinal levels, and might also modulate reticulospinal pathways.     

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.     

Light and electron microscopic study of neurones in the feline lateral cervical nucleus with a descending projection

Nerve cells in the feline lateral cervical nucleus (LCN) with a descending projection were labelled by injections of horseradish peroxidase, in most cases conjugated to wheat germ agglutinin. After small injections into different spinal cord segments in 16 cats the labelled cells were found mainly in the rostral and ventral portions of the ipsilateral LCN, without a detectable topographic organization. Massive bilateral injections were made into the cervical or lumbar enlargement in 6 cats. Unilateral lesions of the dorsolateral funiculus rostral to the injections generally prevented labelling of LCN cells on the same side. The other side was used to calculate the total number of descending LCN neurones and it was estimated that approximately 500 cells projected down the spinal cord. Comparison with the number of labelled cells after the small injections revealed that the descending LCN neurones projected to an average of at least two spinal segments. Quantitative ultrastructural analyses were made of 17 descending neurones from 7 cats subjected to massive unilateral injections into the cervical or lumbar enlargement. The labelled cells constituted a fairly homogeneous population with respect to the investigated somatic and dendritic features. The morphology and relative frequency of the descending neurones indicate that they may constitute parts of the subpopulation of small LCN cells that have previously been assumed to consist of locally ramifying interneurones.     

Cells of origin of pathways descending to the spinal cord in two chondrichthyans,the shark Scyliorhinus canicula and the ray Raja clavata

The cells of origin of pathways descending to the spinal cord in the shark Scyliorhinus canicula and in the ray Raja clavata have been demonstrated by using the horseradish peroxidase (HRP) technique. Following HRP injections in the spinal cord of Scyliorhinus (fourth to sixth segment) and of Raja (15th to 20th segment) labeled neurons could be identified in the rhombencephalon, the mesencephalon, and in the diencephalon. Cells of origin of diencephalic nuclei, which project to the spinal cord, were observed in the nucleus periventricularis hypothalami and in the thalamus ventralis pars medialis which can in this respect be considered hypothalamic. Descending pathways from mesencephalic structures originate from the interstitial nucleus of the fasciculus longitudinalis medialis, the tectum mesencephali, the nucleus intercollicularis, the tectotegmental junction zone, and from diffusely arranged tegmental neurons. A contralateral rubrospinal pathway could be recognized in Raja, but not in Scyliorhinus. Rhombencephalic cells of origin of pathways descending to the spinal cord were found in all parts of the reticular formation, i.e., the nucleus raphes inferior, the nucleus reticularis inferior, medius, superior, and isthmi, in two vestibular nuclei, and in three nuclei, which have been tentatively indicated as nucleus B, F, and G. Furthermore cells of origin of descending pathways have been found in the nucleus tractus descendens nervi trigemini, in the nucleus funiculi lateralis, and in the nucleus tractus solitarii. The descending pathways of the two species studied have been compared with those of other vertebrates. It is concluded that the basic pattern in the organization of descending pathways to the spinal cord, as proposed by ten Donkelaar ('76) for terrestrial vertebrates, also holds for cartilaginous fishes.     

On the organization of cerebellar efferent pathways in the nurse shark (Ginglymostoma cirratum)

The organization of the efferent fiber systems of the nurse shark cerebellum was studied with the Nauta and Fink-Heimer techniques. Lesions of the cerebellar cortex produce a pattern of terminal degeneration restricted to select regions of the cortex, as well as ipsi- and contralateral cerebellar nuclei. We found no evidence of cortical axons projecting beyond the nuclei, to the medulla and spinal cord as has often been reported. Lesions involving the cerebellar nucleus reveal three efferent pathways. The ipsilateral descending cerebello-bulbar (IDC) pathway extends as far caudal as the first spinal segment and issues fibers to the lateral parts of the reticular formation. The brachium conjunctivum splits into a larger ascending limb (BCA) and a smaller descending one (BCD). The latter descends to the caudal medulla terminating along its course in the medial reticular formation. The ascending limb provides an input to (1) a nucleus we have tentatively labelled the “red nucleus,” (2) the trochlear nucleus, (3) the oculomotor nuclei, (4) the midbrain central gray, and (5) a poorly differentiated nucleus in the dorsal thalamus.     

Cervical primary afferent input to vestibulospinal neurons projecting to the cervical dorsal horn: An anterograde and retrograde tracing study in the cat

Vestibulospinal neurons in the caudal half of the medial and descending vestibular nuclei terminate in the cervical spinal cord, not only in the ventral horn and intermediate zone but also in the dorsal horn. The purpose of the present study was to examine whether the areas containing these vestibulospinal neurons are reached by cervical primary afferents. In one group of experiments, wheat germ agglutinin-horseradish peroxidase conjugate and horseradish peroxidase were pressure injected into spinal ganglia C₂-C₈ and revealed anterogradely labeled fibers and boutons in the caudal part (caudal to the dorsal cochlear nucleus) of the ipsilateral medial and descending vestibular nuclei. This projection was verified in experiments in which wheat germ agglutinin-horseradish peroxidase conjugate was microiontophoretically injected into the caudal half of either the medial or the descending vestibular nuclei and revealed retrogradely labeled cells only in ipsilateral spina ganglia C₂-C₇, with a maximum of cells in C₃. In another group of experiments, after microiontophoretic injections of Phaseolus vulgaris leucoagglutinin or Biocytin into either the medial or the descending vestibular nuclei, anterogradely labeled fibers and boutons were present in the cervical spinal cord, mainly bilaterally in the dorsal horn (laminae I–VI) but also, to a lesser extent, in the ventral horn and intermediate zone. The existence of a loop that relays cervical primary afferent information to vestibulospinal neurons projecting to the cervical spinal cord, in particular the dorsal horn, may have implications for vestibular control over local information processing in the cervical dorsal horn. © 1995 Wiley-Liss, Inc.     

The development of serotonergic raphespinal projections in Xenopus laevis

The development of serotonin-immunoreactive neurons in the central nervous system of Xenopus laevis larvae has been studied with special emphasis on the development of the raphe nuclei and raphespinal projections. The first serotonergic neurons were observed in the rostral part of the brain stem at stage 25, only 28 hr after fertilization. By stage 28 some 20 serotonin-immunoreactive neurons were found in the rostral part of the brain stem, bearing small protrusions on the ventromedial side of the soma. These initial axonal outgrowths reach the rostral part of the spinal cord at stage 32. By stage 35/36 the growth cones of the descending serotonergic axons in the spinal cord have reached the level of the anus (10th to 15th myotome). Up to stage 45 the majority of the descending serotonergic axons was found in the dorsolateral part of the marginal zone of the spinal cord. After stage 45 some serotonergic axons were also found scattered over other parts of the spinal marginal zone. Collateral branches were first observed in the caudal part of the brain stem at stage 35/36. Later they occurred also in the rostral (stage 43) and caudal (stage 50) spinal cord, usually on fibers in the ventral half of the spinal cord. The number of serotonergic neurons in the central nervous system (brain stem and hypothalamus) increased steadily throughout development until stage 45. After that the total number of serotonergic neurons in the central nervous system increased about two times faster than the number of serotonergic neurons in the raphe nuclei, due to a massive increase of serotonergic neurons in the hypothalamus. The present study shows that young, just differentiated raphe neurons already contain serotonin. The generation of these neurons appears to take place in the ventricular zone (matrix) of the brain stem between the caudal border of the mesencephalon and the entrance of the nervus octavus. From here these neurons seem to migrate to their final destination. The distribution of serotonin-immunoreactive neurons in the brain stem suggests that a superior (not described so far in Anura) and an inferior raphe nucleus can be distinguished in Xenopus. A rostrocaudal gradient seems to be present in the production of serotonergic neurons which project to the spinal cord. Spinal projections from the raphe nuclei are particularly extensive from the nucleus raphes inferior and gradually decrease rostralwards. In the rostral part of the nucleus raphes superior almost no neurons projecting to the spinal cord are found.     

The spinomesencephalic tract in the cat: its cells of origin and termination pattern as demonstrated by the intraaxonal transport method

The cells of origin and terminal areas of the feline spinomesencephalic tract were investigated by the intraaxonal transport method. Following injection of wheat germ agglutinin-horseradish peroxidase conjugate into the cervical and lumbar enlargements, anterograde labelling was observed in several regions of the dorsal midbrain. The main terminal areas were the periaqueductal gray matter, the intercollicular nucleus, the posterior pretectal nucleus and the nucleus of Darkschewitsch. In addition, there was a sparse projection to the cuneiform nucleus and the anterior pretectal nucleus. The superior colliculus was virtually devoid of labelling except for a weak termination in the caudal part of the deep layers. Although there was a considerable overlap, the projection from the cervical spinal cord to the periaqueductal gray matter terminated more rostrally than that from the lumbar segments, indicating the presence of a somatotopic organization. The retrograde labelling seen after tracer injection into the midbrain terminal areas revealed that the cells of origin were located mainly in the upper cervical segments and in the cervical and lumbar enlargements; in the latter parts of the cord an overwhelming majority were situated in lamina I, with smaller fractions in laminae IV and V, whereas in the upper cervical segments and in the less densely labelled thoracic and sacral segments a much larger proportion of the peroxidase-positive neurons were found in the deep laminae. About 75% of the labelled cells were located contralateral to the injection site. The functional implication of the present results are discussed in relation to somatosensory activity in the mesencephalon. It is suggested that several regions of the dorsal midbrain have an important somesthetic function including that of pain.     

Descending projections to the rat sacrocaudal spinal cord.

Descending supraspinal and propriospinal neurons projecting to the female rat sacrocaudal spinal cord, the portion of the spinal cord that innervates the tail, were identified following injection of Fluoro-Gold into the S1-Ca2 spinal cord segments. This study attempted to determine anatomical substrates for propriospinal and supraspinal control of the tail. Propriospinal neurons were identified throughout laminae V-VIII and X at all levels of the spinal cord. The greatest density of labeling was in the lumbar enlargement, followed by the cervical enlargement, with least in the thoracic spinal cord. Within a given cord level, labeling was greatest within the intermediate zone. In addition, other prominent spinal cord collections included neurons in 1) lamina V of the lumbar enlargement, 2) dorsal lamina X of the cervical enlargement, and 3) the lateral spinal nucleus within the cervical enlargement. Supraspinal cells were identified within raphe nuclei, reticular formation nuclei, dorsal column nuclei, vestibular nuclei, noradrenergic groups, the red nucleus, the periaqueductal gray, the hypothalamus, and the motor cortex. These data indicate that there are significant descending projections to the sacrocaudal spinal cord, with distributions similar to those of other cord levels. Functionally, important supraspinal and propriospinal influences on tail, pelvic viscera and limbs, such as with locomotion, balance, defense, micturition, defecation, and sexual functions, may be mediated by these connections.     

Limb specific connections of the cat magnocellular red nucleus

Afferent and efferent connections of the limb specific divisions of the cat magnocellular red nucleus (RNm) were traced using the bidirectional transport of wheatgerm agglutinin-horseradish peroxidase complex (WGA-HRP). Injection sites within forelimb or hindlimb RNm regions were identified by microelectrode recording and confirmed by the position of labeled rubrospinal terminals. Additional injections into structures that project to, or receive input from, RNm confirmed the somatotopic organization of these pathways. The forelimb region of RNm receives input from the posteriolateral part of the anterior interpositus nucleus (NIA) and the intermediate part of the posterior interpositus nucleus (NIP). The hindlimb region of RNm receives input from anteriomedial NIA and medial NIP. Terminals of NIA cells densely fill all of RNm, but terminals of NIP cells form a half shell on the medial, ventral, and posterior borders of RNm without encroaching on RNm's lateral edge or central core. Forelimb and hindlimb RNm are reciprocally connected with the caudal cuneate and gracile nuclei respectively. There is little or no input to RNm from the medial or lateral cerebellar nuclei. Forelimb RNm, which also contains a face representation, projects to the lateral reticular nucleus, cell group f of the inferior vestibular nucleus, the facial nucleus, the main sensory nucleus of the trigeminal nerve, the caudal cuneate nucleus, the parvicellular reticular formation, and cervical segments of the spinal cord. A few fibers from forelimb RNm project directly to motor neurons in the lower cervical cord. Hindlimb RNm projects to only the lateral reticular nucleus, gracile nucleus, and lower spinal segments. Forelimb and hindlimb RNm project to different regions of the lateral reticular nucleus with some overlap.     

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

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

Comparative topography of projections from the mesodiencephalic junction to the inferior olive, vestibular nuclei, and upper cervical cord in the cat

Distributions of neurons located in the central rostral mesencephalon and caudal diencephalon that project to the upper cervical spinal cord, vestibular nuclei, or inferior olive were studied in the cat by using retrograde axonal transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Afferent sources to all of these targets were observed in the interstitial nucleus of Cajal (INC), the region surrounding the fasciculus retroflexus (PF), and the nucleus of the fields of Forel (NFF). Three-dimensional reconstruction revealed differences in densities of cells projecting from these common areas. Spinal projecting cells were present in slightly greater numbers in the caudal two-thirds of the INC, whereas those projecting to the vestibular complex were more numerous in the rostral two-thirds of this nucleus. A relatively smaller number of olivary projecting cells were dispersed throughout the INC. Olivary afferent sources outnumber those with spinally directed or vestibularly directed axons in the PF region. In the fields of Forel, cells projecting to the vestibular nuclei or inferior olive were concentrated medially, whereas cells projecting to the spinal cord appeared both medially and laterally. Each type of afferent source was also seen in the nucleus of the posterior commissure and the posterior ventral lateral hypothalamic area. Unique sources of afferents to the inferior olive were observed in the parvicellular red nucleus (ipsilateral to the injections) and the anterior and posterior pretectal nuclei. A large number of labeled neurons was seen in the nucleus of Darkschewitsch after injections of tracer into the inferior olive, but this projection did not appear to be unique, as small numbers of labeled cells were also seen after injections into the cervical spinal cord. The Edinger-Westphal nucleus and the adjacent somatic oculomotor nucleus contained cells which projected separately to the spinal cord or the vestibular complex, and the superior colliculus contained cells which projected separately to the contralateral spinal cord or the contralateral inferior olive. In this study, it was also noted that neurons in the medial terminal nucleus of the accessory optic tract projected to the ipsilateral inferior olive or to the contralateral vestibular complex. These differences in locations and densities of cells projecting to the cervical spinal cord, vestibular complex, and inferior olive may underlie functional specializations in these areas in relation to vertical eye and head movement control and to neural systems controlling postural adjustments accompanying limb movements.     

Brainstem projections to the rat cuneate nucleus

Neurons in the pontomedullary tegmentum have been proposed as a final common pathway subserving descending inhibition in the dorsal column nuclei. To investigate the anatomical substrate for these descending effects, brainstem projections to the cuneate nucleus of rats were studied with injections of lectin-conjugated horseradish peroxidase. In rats with iontophoretic tracer injections in this nucleus, many labeled neurons were detected near the injection site, especially ventral and caudal to it. Intrinsic reciprocal projections were observed after injections in caudal, middle, or rostral levels of the cuneate nucleus. Neurons were labeled in the red nucleus, in agreement with previous anatomical studies, and also in the trigeminal, vestibular, and cochlear nuclei. An ipsilateral dorsomedial group of neurons was labeled in the upper cervical segments and scattered neurons were also labeled bilaterally near the central canal. Sparse retrograde labeling in the tegmentum was focused in the lateral paragigantocellular nucleus and caudal raphe. Consistent with the retrograde experiments, anterograde labeling after pressure injections of lectin-conjugated horseradish peroxidase in the pontomedullary tegmentum was very sparse within the dorsal column nuclei; labeling was dense, however, in the region immediately ventral to these nuclei. These results confirm previous work indicating that the activity of cuneate neurons is modulated by brainstem sensory nuclei. However, it appears that direct projections to the cuneate nucleus from pontine and rostral medullary regions are sparser than previously suggested. The last link of a polysynaptic descending inhibitory pathway may include GABAergic neurons immediately adjacent to the dorsal column nuclei and/or intrinsic to these nuclei.