Trigemino-reticulo-facial and trigemino-reticulo-hypoglossal pathways in the rat

This study was undertaken to identify premotor neurons in the pontomedullary reticular formation serving as relay neurons between the sensory trigeminal complex and the motor nuclei of the VIIth and XIIth nerves. Trigeminoreticular projections were first investigated after injections of anterogradely transported tracers (biotinylated dextran amine, biocytin) into single subdivisions of the sensory trigeminal complex. The results show that the trigeminoreticular projections were abundant from the pars interpolaris (5i) and caudalis (5c) and moderate from pars oralis (5o) of the spinal trigeminal nucleus. Injections into the 5i and 5c produce dense anterograde labeling (1) in the dorsal medullary reticular field; (2) in the parvocellular reticular field, medially adjacent to the 5i; and (3) more rostral in the region dorsal and lateral to the superior olivary nucleus. Some labeled terminals were also found in the intermediate reticular field, whereas only light anterograde labeling was observed in the gigantocellular and oral pontine reticular formation. The 5o sends fibers and terminals throughout the whole reticular formation, with no clear preferential projections within a particular field. Only light projections originated from the principal nucleus (5P). In a second series of experiments, we examined whether premotor neurons in the reticular formation are afferented by trigeminal fibers. Double labeling was performed by injection of an anterograde tracer in the 5i and 5c and retrograde tracer (gold-horseradish peroxidase complex) into the VII or the XII motor nucleus on the same side. Retrogradely labeled neurons in contact with anterogradely labeled boutons were found throughout the reticular formation with predominance in the parvocellular and intermediate reticular fields. These experiments demonstrate the existence of trigeminal disynaptic influences, via reticular neurons of the pontomedullary reticular formation, in the control of orofacial motor behaviors.     

Axonal trajectories and terminations of on- and off-cells in the cat lower brainstem

Two physiologically defined classes of pontomedullary raphe neurons were intracellularly labeled in order to determine the target nuclei of their axonal projections. In the lightly anesthetized cat, cells either increased (on-cells) or decreased (off-cells) their discharge rate during the paw withdrawal reflex evoked by noxious pinch or heat. On- and off-cells were injected with horseradish peroxidase and the initial course of labeled axons through the lower brainstem was reconstructed. On-cell projections to the pontomedullary raphe and medial reticular nuclei were sparse. On-cells projected densely to regions of the lateral reticular formation and the ventrolateral medulla at both rostral and caudal medullary levels. In general, on-cells had few collaterals and spare axonal swellings. In contrast to on-cells, most off-cells had axons that collateralized densely within the brainstem raphe and adjacent reticular formation. Such collaterals were either local, within the neuron's dendritic field, or distant, involving a projection of 1-8 mm. One off-cell had a dense terminal field within the sensory trigeminal complex, a projection that may subserve the inhibition of trigeminal sensory neurons produced by raphe magnus stimulation. Well-labeled off-cells had numerous collaterals and dense regions of axonal swellings. In summary, off-cells terminated densely in the raphe magnus and adjacent reticular formation whereas on-cells projected predominantly to the ventrolateral medulla, a region implicated in autonomic control. Local off-cell collaterals provide an anatomical substrate that would enable off-cells to coordinate the activity of on- and off-cells through synaptic contacts.     

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.     

GABAergic and glycinergic neurons projecting to the trigeminal motor nucleus: A double labeling study in the rat

The distribution of GABAergic and glycinergic premotor neurons projecting to the trigeminal motor nucleus (Vm) was examined in the lower brainstem of the rat by a double labeling method combining retrograde axonal tracing with immunofluorescence histochemistry. After injection of the fluorescent retrograde tracer, tetramethylrhodamine dextran amine (TRDA), into the Vm unilaterally, neurons labeled with TRDA were seen ipsilaterally in the mesencephalic trigeminal nucleus, and bilaterally in the parabrachial region, the supratrigeminal and intertrigeminal regions, the reticular formation just medial to the Vm, the principal sensory and spinal trigeminal nuclei, the pontine and medullary reticular formation, especially the parvicellular part of the medullary reticular formation, the alpha part of the gigantocellular reticular nucleus, and the medullary raphe nuclei. Some of these neurons labeled with TRDA were found to display glutamic acid decarboxylase (the enzyme involved in GABA synthesis)-like or glycine-like immunoreactivity. Such double-labeled neurons were seen mainly in the supratrigeminal region, the reticular region adjacent to the medial border of the Vm, and the dorsal part of the lateral reticular formation of the medulla oblongata; a number of them were further scattered in the intertrigeminal region, the alpha part of the gigantocellular reticular nucleus, the nucleus raphe magnus, the principal sensory trigeminal nucleus, and the interpolar subnucleus of the spinal trigeminal nucleus. These neurons were considered to be inhibitory (GABAergic or glycinergic) neurons sending their axons to motoneurons in the Vm, or to local interneurons within and around the Vm. © 1996 Wiley-Liss, Inc.     

Brainstem afferents to the omnipause region in the cat: a horseradish peroxidase study

"Omnipause" neurons (OPNs), located in the nucleus raphe pontis and the reticular formation, actively suppress saccadic eye movements during intersaccadic intervals. To determine the brainstem afferents that may inhibit the OPNs and thereby allow a saccade to occur, we injected horseradish peroxidase into the raphe pontis of four cats at the site of physiologically identified OPNs. Labeled neurons were found in a number of brainstem nuclei. The greatest concentrations, composed of small to medium-sized neurons, were located in a group of nuclei around the habenulopeduncular tract, in the rostral mesencephalic reticular formation, in the deep layers of the superior colliculus, and in parts of the subjacent cuneiform and subcuneiform reticular nuclei. Smaller numbers were found in the nucleus reticularis pontis oralis. Caudal to the injection site, labeled neurons were scattered in parts of the nuclei reticularis gigantocellularis, paragigantocellularis dorsalis, and paragigantocellularis lateralis. A few neurons were labeled in a restricted region of the causal part of the nucleus prepositus hypoglossi and in the nucleus reticularis medullaris ventralis. Larger numbers of neurons were labeled in the dorsal column nuclei and in parts of the cochlear nuclei. Smaller numbers were found in the spinal trigeminal nucleus, the lateral nucleus of the superior olive, and the fastigial nucleus of the cerebellum. The nonreticular brainstem projections may contribute sensory information in a number of modalities since OPNs respond to visual, somesthetic, and auditory stimuli. Our findings indicate a number of regions that may contain neural elements impinging on the OPNs. The best prospects for a saccade initiation signal from one of the labeled populations appear to be the meso-diencephalic reticular formation and/or the superior colliculus.     

Brainstem origin of serotonin- and enkephalin-immunoreactive afferents to the opossum's cerebellum

Previous studies have described the distribution of serotonin- and enkephalin-immunoreactive elements in the posterior lobe vermis of the opossum's cerebellum. In the present study we have used a double labeling paradigm which combines the retrograde transport of horseradish peroxidase (HRP) with serotonin and enkephalin immunohistochemistry to determine the brainstem origin of serotoninergic and enkephalinergic neurons that project to the opossum's cerebellar cortex. Subsequent to HRP injections into the posterior lobe vermis, widespread areas of the medulla and pons were found to contain retrogradely labeled neurons. Serotonin-immunoreactive somata are present primarily in the raphe nuclei and the adjacent reticular formation. Enkephalinergic neurons were numerous in the raphe nuclei, medial accessory olive, gigantocellular reticular formation, locus coeruleus, and the nucleus of the trapezoid body. However, serotoninergic neurons that project to the cerebellum were located only in the medullary pyramids and the reticular formation adjacent to the raphe. Double-labeled enkephalinergic neurons were located 1) within the medullary pyramids, 2) throughout the extent of the caudal medial accessory olive, 3) in the rostral subnucleus a of the medial accessory olive, 4) in the nucleus reticularis gigantocellularis pars ventralis, 5) in the nucleus reticularis lateralis, and 6) in the nucleus reticularis ventralis lateral to the inferior olivary complex. These results indicate that although neurons containing serotonin and enkephalin immunoreactivity may be present in some of the same pontine and medullary nuclei, those serotoninergic and enkephalinergic neurons that project to the cerebellum are present primarily in restricted and spatially separate regions of the caudal medulla.     

Brainstem afferents to the oculomotor omnipause neurons in monkey

To determine how saccade-related areas in the brainstem address the saccade generator, we examined the afferents to the nucleus raphe interpositus. This region contains the omnipause neurons, which are pivotal in the generation of saccades. Horseradish peroxidase injected iontophoretically into the nucleus raphe interpositus retrogradely labeled a variety of brainstem nuclei. The greatest numbers of labeled neurons were in the paramedian pontomedullary reticular formation, in the nuclei reticularis gigantocellularis, and paragigantocellularis lateralis. Labeling was more modest but consistent in the interstitial nucleus of Cajal and the adjacent mesencephalic reticular formation, the middle gray of the superior colliculi, the region dorsolateral to the nucleus reticularis tegmenti pontis, and the medial vestibular nucleus. A few neurons were labeled around the habenulopeduncular tract and in the medial portion of the nucleus of the fields of Forel, in the nucleus reticularis medullaris ventralis, and in the spinal nucleus of the trigeminal nerve, the cochlear nucleus, and the superior olivary complex. The distribution and density of labeling suggest that omnipause neurons in the monkey are more intimately connected with other oculomotor structures than those in the cat. In addition, the rhombencephalic reticular afferents to the monkey omnipause neurons are more concentrated in their immediate vicinity than in the cat. The label consistently found dorsolateral to the nucleus reticularis tegmenti pontis may be a newly discovered link in saccade generation.     

Distribution of serotonin 2A, 2C and 3 receptor mRNA in spinal cord and medulla oblongata

It is known that 5-HT receptors have significant roles in nociceptive and motor functions. We have compared the cellular localization of the mRNAs encoding serotonin 5-HT(2A,) 5-HT(2C,) 5-HT(3) receptor subtypes within different levels of the rat spinal cord and medulla. In the spinal cord, 5-HT(2C) receptor mRNA is expressed at high levels in most of the gray matter, except for lamina II. In contrast, 5-HT(2A) receptor mRNA is expressed exclusively in lamina IX. 5-HT(3) receptor mRNA has a low level and diffuse pattern of expression increasing towards the ventral horn. In both gray and white matter, there is a characteristic presence of a few highly stained cells. For each subtype, the expression pattern is similar in all four levels of the spinal cord. In the medulla, 5-HT(2C) receptor mRNA is at high levels in many nuclei including the hypoglossal nucleus, the gigantocellular reticular nucleus alpha and the parvocellular reticular nucleus alpha, the spinal nucleus of the trigeminal tract, the facial, and the dorsal medullary reticular field. Moderate to low levels of expression are seen in the spinal vestibular nucleus, the vagus, the solitary nuclei and the raphe. 5-HT(2A) receptor is expressed at high levels in some nuclei such as the hypoglossal nucleus, the intercalate nucleus, the inferior olive and the lateral reticular nucleus. Moderate to low levels of expression are seen in the facial, the medial vestibular nuclei, the nucleus ambiguous, the vagus, and the gigantocellular reticular nucleus. 5-HT(3) receptor mRNA is present at low levels in most of the nuclei examined, with a few scattered strongly labeled cells. The results show a distinct distribution of the three subtypes of receptors supporting their physiological roles and will help to understand the mechanisms of nociception and motor function.     

Afferent projections to the oral motor nuclei in the rat

Projections to the trigeminal, facial, ambiguus, and hypoglossal motor nuclei were determined by using horseradish peroxidase histochemistry. Most of the afferent projections to these motor nuclei were from the brainstem reticular formation, frequently in areas adjacent to other synergetic motor nuclei. The reticular formation lateral to the hypoglossal nucleus and reticular structures surrounding the trigeminal motor nucleus projected to each of these other brainstem motor nuclei involved in oral-facial function. Afferent projections to these motor nuclei also were organized along the rostrocaudal axis. Within the reticular formation most of the afferent projections to the trigeminal motor nucleus originated rostral to the majority of neurons projecting to the hypoglossal and ambiguus nuclei, which in turn were rostral to the primary source of reticular afferents to the facial nucleus. In comparison, projections from the sensory trigeminal nuclei and nucleus of the solitary tract were sparse. The interneuron pools that project to the orofacial motoneurons provide one further link in understanding the brainstem substrates for integrating oral and ingestive behaviors.     

Brainstem projections to midline and intralaminar thalamic nuclei of the rat

The projections from the brainstem to the midline and intralaminar thalamic nuclei were examined in the rat. Stereotaxic injections of the retrograde tracer cholera toxin beta -subunit (CTb) were made in each of the intralaminar nuclei of the dorsal thalamus: the lateral parafascicular, medial parafascicular, central lateral, paracentral, oval paracentral, and central medial nuclei; in the midline thalamic nuclei-the paraventricular, intermediodorsal, mediodorsal, paratenial, rhomboid, reuniens, and submedius nuclei; and, in the anteroventral, parvicellular part of the ventral posterior, and caudal ventral medial nuclei. The retrograde cell body labeling pattern within the brainstem nuclei was then analyzed. Nearly every thalamic site received a projection from the deep mesencephalic reticular, pedunculopontine tegmental, dorsal raphe, median raphe, laterodorsal tegmental, and locus coeruleus nuclei. Most intralaminar thalamic sites were also innervated by unique combinations of medullary and pontine reticular formation nuclei such as the subnucleus reticularis dorsalis, gigantocellular, dorsal paragigantocellular, lateral, parvicellular, caudal pontine, ventral pontine, and oral pontine reticular nuclei; the dorsomedial tegmental, subpeduncular tegmental, and ventral tegmental areas; and, the central tegmental field. In addition, most intralaminar injections resulted in retrograde cell body labeling in the substantia nigra, nucleus Darkschewitsch, interstitial nucleus of Cajal, and cuneiform nucleus. Details concerning the pathways from the spinal trigeminal, nucleus tractus solitarius, raphe magnus, raphe pallidus, and the rostral and caudal linear raphe nuclei to subsets of midline and intralaminar thalamic sites are discussed in the text. The discussion focuses on brainstem-thalamic pathways that are likely involved in arousal, somatosensory, and visceral functions.     

Nuclei of origin of monoaminergic, peptidergic, and cholinergic afferents to the cat trigeminal motor nucleus: a double-labeling study with cholera-toxin as a retrograde tracer.

The aim of the present study was to determine the brainstem afferents and the location of neurons giving rise to monoaminergic, cholinergic, and peptidergic inputs to the cat trigeminal motor nucleus (TMN). This was done in colchicine treated animals by using a very sensitive double immunostaining technique with unconjugated cholera-toxin B subunit (CT) as a retrograde tracer. After CT injections in the TMN, retrogradely labeled neurons were most frequently seen bilaterally in the nuclei reticularis parvicellularis and dorsalis of the medulla oblongata, the alaminar spinal trigeminal nucleus (magnocellular division), and the adjacent pontine juxtatrigeminal region and in the ipsilateral mesencephalic trigeminal nucleus. We further observed that inputs to the TMN arise from the medial medullary reticular formation (the nuclei retricularis magnocellularis and gigantocellularis), the principal bilateral sensory trigeminal nucleus, and the dorsolateral pontine tegmentum. In addition, the present study demonstrated that the TMN received 1) serotonergic afferents, mainly from the nuclei raphe obscurus, pallidus, and dorsalis; 2) catecholaminergic afferent projections originating exclusively in the dorsolateral pontine tegmentum, including the K?lliker-Fuse, parabrachialis lateralis, and locus subcoeruleus nuclei; further, that 3) methionin-enkephalin-like inputs were located principally in the medial medullary reticular formation (nuclei reticularis magnocellularis and gigantocellularis and nucleus paragigantocellularis lateralis), in the caudal raphe nuclei (Rpa and Rob) and the dorsolateral pontine tegmentum; 4) substance P-like immunoreactive neurons projecting to the TMN were present in the caudal raphe and Edinger-Westphal nuclei; and 5) cholinergic afferents originated in the whole extent of the nuclei reticularis parvicellularis and dorsalis including an area located ventral to the nucleus of the solitary tract at the level of the obex. In the light of these anatomical data, the present report discusses the possible physiological involvement of TMN inputs in the generation of the trigeminal jaw-closer muscular atonia occurring during the periods of paradoxical sleep in the cat.     

Reevaluation of projections from the mesencephalic trigeminal nucleus to the medulla and spinal cord: New projections. A combined retrograde and anterograde horseradish peroxidase study

Microinjection of horseradish peroxidase (HRP) into the medullary parvocellular reticular formation (NPvc) resulted in retrograde labeling of neurons throughout the mesencephalic trigeminal nucleus (Mes V). Labeled cells were large and ovoid and were distributed primarily in the expanded pontine part of the nucleus. However, none of the small neurons in Mes V were labeled. Injections of HRP made into adjacent brainstem structures including the nucleus gigantocellularis, ventrolateral reticular formation, vestibular complex, and the spinal trigeminal nucleus failed to label neurons in Mes V. Injections made into the medullary raphe and into regions reported to receive inputs from Mes V–spinal cord, nucleus tractus solitarius, hypoglossal nucleus, and facial nucleus–were also not followed by transport to Mes V. Anterograde axonal transport of HRP from the region of reticular formation innervated by Mes V also labeled axons projecting to Mes V and to visceral and somatic sensorimotor nuclei in the lower brainstem. Recent reports of afferents from the amygdala to Mes V suggest that reflexes involving the mesencephlic trigeminal nucleus might be modulated by signals from limbic and autonomic as well as somatic centers in the brain.     

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)     

Brainstem projections to the normal and noradrenergically hyperinnervated trigeminal motor nucleus

The noradrenergic innervation of the trigeminal motor nucleus of the rat can be increased severalfold by neonatal treatment with the neurotoxin, 6-hydroxydopamine. The brainstem projections to the nucleus were studied by injecting HRP into the nucleus of normal and noradrenergically hyperinnervated rats. In order to identify the source of the noradrenergic innervation, the fluorescent dye, True Blue, was used as a retrograde tracer in combination with the glyoxylic acid histofluorescence method for catecholamines. In both control and neonatally treated rats, the noradrenergic innervation of the motor nucleus was shown to arise from an ipsilateral group of cells located among the fibers of the lateral lemniscus just rostral to the motor nucleus. Our results confirmed the high degree of specificity of noradrenergic innervation, which arises exclusively from this lateral tegmental noradrenergic cell group. During the process of sprouting, this specificity is maintained since only those noradrenergic cells normally innervating the nucleus were retrogradely labeled in neonatally treated animals. Other noradrenergic projections which are also increased in these animals, such as the nearby locus ceruleus innervation of the main sensory trigeminal nucleus, do not spread to the motor trigeminal nucleus. HRP-labeled nonadrenergic cells were concentrated dorsally, with scattered cells surrounding the nucleus. A similar distribution was observed contralateral to the injection site. The mesencephalic trigeminal nucleus was labeled only ipsilateral to the injection. The motor nucleus also receives an extensive bilateral input from the pontine and medullary reticular formation. The medial reticular formation nuclei, including nucleus pontis caudalis, nucleus gigantocellularis, and nucleus reticularis ventralis contained large labeled cells, which were especially numerous in the retrotrigeminal area. Smaller, lateral reticular formation neurons were concentrated rostrally and ipsilaterally in the nucleus pontis lateralis. HRP retrograde labeling revealed no obvious change in the overall pattern of cells innervating the trigeminal motor nucleus following noradrenergic hyperinnervation.     

Neural pathway for aggressive display in Betta splendens: midbrain and hindbrain control of gill-cover erection behavior

Horseradish peroxidase (HRP) was used to identify parts of the presumptive neural pathway for gill cover erection, a behavioral display pattern performed by Siamese fighting fish (Betta splendens) during aggressive interactions. Motor, motor integration and sensory areas were identified in the medulla and mesencephalon. Motor neurons of the dilator operculi muscle, the effector muscle for gill cover erection, are located in the lateral and medial parts of the caudal trigeminal motor nucleus. Iontophoretic injections of HRP into the lateral trigeminal motor nucleus resulted in labeled cell bodies in two motor areas (medial part of the trigeminal motor nucleus, anterior part of the motor nucleus of cranial nerve IX-X), two parts of the reticular formation (medial and inferior reticular areas), and two nuclei of the octavolateralis system (nucleus medialis, magnocellular octaval nucleus). The HRP injections in the medial part of the caudal trigeminal motor nucleus resulted in labeled cells in the lateral part of the nucleus and in the medial reticular nucleus. Discrete injections of HRP into nucleus medialis revealed a strong axonal projection that terminated in the torus semicircularis. The medial reticular area and both of the octavolateralis nuclei received projections from their contralateral counterparts. Connections between motor areas, and between parts of the reticular formation, may coordinate the performance of gill cover erection with other behavioral patterns used during aggressive display. Connections with the octavolateralis system may provide information on the strength of an opponent's tail beats via the lateral-line system, as well as vestibular information about the fish's own orientation during aggressive display. The organization of inputs to the trigeminal motor nucleus in Betta, a perciform fish, was found to differ from that reported in the common carp, a cypriniform fish. These differences may underlie the different behavioral capabilities of the two groups of fish.     

Anatomical connections of the nucleus prepositus of the cat

The afferent and efferent connections of the nucleus prepositus hypoglossi with brainstem nuclei were studied using anterograde and retrograde axonal transport techniques, and by intracellular recordings and injections of horseradish peroxidase into prepositus hypoglossi neurons. The results of experiments in which horseradish peroxidase was injected into the prepositus hypoglossi suggest that the major inputs to the prepositus hypoglossi arise from the ipsi- and contralateral perihypoglossal nuclei (particularly the prepositus hypoglossi and intercalatus), vestibular nuclei (particularly the medial, inferior, and ventrolateral nuclei), the paramedian medullary and pontine reticular formation, and from the cerebellar cortex (flocculus, paraflocculus, and crus I; the nodulus was not available for study). Regions containing fewer labeled cells included the interstitial n. of Cajal, the rostral interstitial n. of the medial longitudinal fasciculus, the n. of the posterior commissure, the superior colliculus, the n. of the optic tract, the extraocular motor nuclei, the spinal trigeminal n., and the central cervical n. The efferent connections of the prepositus hypoglossi were studied by injecting 3H-leucine into the prepositus hypoglossi, and by following the axons of intracellularly injected prepositus hypoglossi neurons. The results suggest that in addition to the cerebellar cortex, the most important extrinsic targets of prepositus hypoglossi efferents are the vestibular nuclei (particularly the medial, inferior, and ventrolateral nuclei, and the area X), the inferior olive (contralateral dorsal cap of Kooy and ipsilateral subnucleus b of the medial accessory olive), the paramedian medullary and pontine reticular formation, the reticular formation surrounding the parabigeminal n., the contralateral superior colliculus and pretectum, the extraocular motor nuclei (particularly the contralateral abducens nucleus and the ipsilateral medial rectus subdivision of the oculomotor nucleus), the ventral lateral geniculate n., and the central lateral thalamic nucleus. Other areas which were lightly labeled in the autoradiographic experiments were the contralateral spinal trigeminal n., the n. raphe pontis, the Edinger Westphal n., the zona incerta, and the paracentral thalamic n. Many of the efferent connections of the prepositus hypoglossi appear to arise from principal prepositus hypoglossi neurons whose axons collateralize extensively in the brainstem. On the other hand, small prepositus hypoglossi neurons project to the inferior olive, and multidendritic neurons project to the cerebellar flocculus, apparently without collateralizing in the brainstem.(ABSTRACT TRUNCATED AT 400 WORDS)     

Localization of trigeminal, spinal, and reticular neurons involved in the rat blink reflex

Electrical stimulation of the supraorbital nerve (SO) induces eyelid closure by activation of orbicularis oculi muscle motoneurons located in the facial motor nucleus (VII). Neurons involved in brainstem central pathways implicated in rat blink reflex were localized by analyzing c-Fos protein expression after SO stimulation in conjunction with tracing experiments. A retrograde tracer (gold-horseradish peroxidase [HRP]) was injected into the VII. The distribution patterns of activated c-Fos-immunoreactive neurons and of neurons exhibiting both c-Fos immunoreactivity and gold-HRP labeling were determined in the sensory trigeminal complex (STC), the cervical spinal cord (C1), and the pontomedullary reticular formation. Within the STC, c-Fos immunoreactivity labeled neurons in the ipsilateral ventral part of the principal nucleus, the pars oralis and interpolaris, and bilaterally in the pars caudalis. Colocalization of gold-HRP and c-Fos immunoreactivity was observed in neurons of ventral pars caudalis layers I-IV and ventral pars interpolaris. In C1, SO stimulation revealed c-Fos neurons in laminae I-V. After additional injections in VII, the double-labeled c-Fos/gold-HRP neurons were concentrated in laminae IV and V. Although c-Fos neurons were found throughout the pontomedullary reticular formation, most appeared rostrally around the motor trigeminal nucleus and in the ventral parvocellular reticular nucleus medial to the fiber bundles of the seventh nerve. Caudally, c-Fos neurons were in the lateral portion of the dorsal medullary reticular field. In addition, these reticular areas contained double-labeled neurons in electrically stimulated rats that had received gold-HRP injections in the VII. The presence of double-labeled neurons in the STC, C1, and the reticular formation implies that these neurons receive sensory information from eyelids and project to the VII. These double-labeled neurons could then be involved in di- or trisynaptic pathways contributing to the blink reflex.     

Projections from the orbital gyrus in the cat. I. To brain stem structures

Direct projections from the anterior part of the orbital gyrus and its immediate vicinity to the brain stem in the cat were investigated by means of the Nauta and Marchi methods. These experiments were carried out in an attempt to identify the anatomical substrates subserving orbital-gyral inhibitory influences on somatic reflexes. The cortical layers of the anterior orbital gyrus and the immediately surrounding cortex were ablated unilaterally. Preterminal degeneration was found bilaterally in the spinal trigeminal nucleus. The most consoicuous degeneration occurred in the interpoler nuclei, caudal parts of the oral nuclei, and rostral portions of the caudal nuclei of the spinal trigeminal complex. Thus, degeneration was found in the trigeminal sensory nucleus from the level of the inferior olive rostrally to levels just caudal to the obex. Preterminal degeneration was also observed bilaterally in the mesencephalic reticular formation. Heavy preterminal degeneration was found in the reticular formation of the rostral midbrain and moderate degeneration was observed in the caudal mesencephalic reticular formation, the pontile reticular substance and medullary reticular nuclei. Based on these findings, the possible pathways involved in the mediation of orbital-cortically induced inhibition on somatic reflexes were discussed.     

Location,morphology, and central projections of mesencephalic trigeminal neurons innervating rat masticatory muscles studied by axonal transport of choleragenoid-horseradish peroxidase

Retrograde and transganglionic transport of horseradish peroxidase conjugated to the B-fragment of cholera toxin (B-HRP) was used to study the location, morphology, and central projections of mesencephalic trigeminal (Me5) neurons innervating rat masticatory muscles. Labeled Me5 cell bodies were found throughout the Me5 nucleus from a level slightly caudal to the trigeminal motor nucleus to the level of the superior colliculus 5 mm further rostrally. Occasionally, labeled Me5 cells were observed in the anterior medullary velum, in the cerebellum, and in the brainstem contralateral to the B-HRP injection. The vast majority of the labeled Me5 cells were pseudounipolar, but multipolar cells were also found. Extensive central projections from labeled Me5 cells could be seen extending from the nucleus of Darkschewitsch rostrally to the C2 segment caudally. Small but consistent projections from Me5 neurons were observed in nuclear islands among the incoming Me5 root fibers. Trigeminal and hypoglossal motor nuclei received direct projections from Me5 cells, but not the facial motor nucleus. The most prominent Me5 projections appeared in the brainstem reticular formation, including the supratrigeminal nucleus. Smaller projections also extended into the main sensory trigeminal nucleus, trigeminal subnucleus oralis, and the nucleus of the solitary tract. © 1993 Wiley-Liss, Inc.     

Raphespinal and reticulospinal axon collaterals to the hypoglossal nucleus in the rat.

Neurons in the medial tegmental field project directly to spinal somatic motoneurons and to cranial motoneuron pools such as the hypoglossal nucleus. The axons of these neurons may be highly collateralized, projecting to multiple levels of the spinal cord and to many diverse regions at different levels of the neuraxis. We employed a double fluorescent retrograde tracer technique to examine whether medial tegmental neurons that project to the spinal cord also project to the hypoglossal nucleus. Injections of Diamidino Yellow into the hypoglossal nucleus and Fast Blue into the spinal cord produced large numbers of double labeled neurons in the medial tegmental field, particularly in the caudal raphe nuclei and adjacent ventromedial reticular formation. In these structures the number of neurons projecting to both the hypoglossal nucleus and the spinal cord was equivalent to the number of neurons projecting to multiple levels of the spinal cord observed in control animals. Fewer neurons projecting to both the hypoglossal nucleus and the spinal cord were observed in several other nuclei and subregions of the medial tegmental field, while almost no such neurons were observed in the lateral tegmental field or other pontomedullary structures. These results demonstrate that neurons of the caudal raphe nuclei and adjacent ventromedial reticular formation project to both the spinal cord and the hypoglossal nucleus, and support the concept that the diffuse projections to motoneuron pools from the medial tegmental field globally modulate both spinal and cranial somatic motoneuron excitability.