Response properties of periodontal mechanosensitive neurons in the trigeminal spinal tract nucleus of the cat.

Periodontal mechanosensitive (PM) units were recorded from the trigeminal spinal tract nucleus (Vst) of the cat. The Vst is divided into three subnuclei: oralis (Vo), interpolaris (Vi), and caudalis (Vc). The receptive fields of PM units in Vo and Vi were arranged in a dorsoventral sequence in the mandibular to maxillary divisions, and those in Vc were arranged in a mediolateral sequence. The majority of Vo units were single-tooth ones, whereas more than half the Vi units and all the Vc ones were multitooth units. The PM units in each subnucleus were predominantly responsive to canine tooth stimulation. Most of the PM units in Vo and Vi gave sustained responses to pressure applied to the tooth, were directionally selective, and were most actively excited by canine tooth stimulation in the caudomedial or rostrolateral direction. Vc units, however, were transient. The threshold intensity for firings by canine tooth stimulation was less than 0.05 N. These findings indicate that only the response properties of PM units in the rostral part of Vst resemble those of the trigeminal main sensory nucleus neurons and primary afferent nerves.     

SCN2B in the Rat Trigeminal Ganglion and Trigeminal Sensory Nuclei

The beta-2 subunit of the mammalian brain voltage-gated sodium channel (SCN2B) was examined in the rat trigeminal ganglion (TG) and trigeminal sensory nuclei. In the TG, 42.6 % of sensory neurons were immunoreactive (IR) for SCN2B. These neurons had various cell body sizes. In facial skins and oral mucosae, corpuscular nerve endings contained SCN2B-immunoreactivity. SCN2B-IR nerve fibers formed nerve plexuses beneath taste buds in the tongue and incisive papilla. However, SCN2B-IR free nerve endings were rare in cutaneous and mucosal epithelia. Tooth pulps, muscle spindles and major salivary glands were also innervated by SCN2B-IR nerve fibers. A double immunofluorescence method revealed that about 40 % of SCN2B-IR neurons exhibited calcitonin gene-related peptide (CGRP)-immunoreactivity. However, distributions of SCN2B- and CGRP-IR nerve fibers were mostly different in facial, oral and cranial structures. By retrograde tracing method, 60.4 and 85.3 % of TG neurons innervating the facial skin and tooth pulp, respectively, showed SCN2B-immunoreactivity. CGRP-immunoreactivity was co-localized by about 40 % of SCN2B-IR cutaneous and tooth pulp TG neurons. In trigeminal sensory nuclei of the brainstem, SCN2B-IR neuronal cell bodies were common in deep laminae of the subnucleus caudalis, and the subnuclei interpolaris and oralis. In the mesencephalic trigeminal tract nucleus, primary sensory neurons also exhibited SCN2B-immunoreactivity. In other regions of trigeminal sensory nuclei, SCN2B-IR cells were very infrequent. SCN2B-IR neuropil was detected in deep laminae of the subnucleus caudalis as well as in the subnuclei interpolaris, oralis and principalis. These findings suggest that SCN2B is expressed by various types of sensory neurons in the TG. There appears to be SCN2B-containing pathway in the TG and trigeminal sensory nuclei.     

Gamma-aminobutyric acid-immunoreactive neurons in the rat trigeminal nuclei

The distribution of GABAergic neurons in the rat trigeminal nuclei was studied using a highly specific monoclonal antibody (mAb3A12) to gamma-aminobutyric acid (GABA). Immunopositive cells were relatively abundant in the marginal and gelatinosa beds of the caudal part of the trigeminal spinal tract nucleus, and in the dorsomedial areas of the oral subnucleus and the principal nucleus. A high density of GABA-immunoreactive somata was also found in the rostral part of the oral subnucleus and in the adjacent parvicellular reticular formation as well as in the supratrigeminal and intertrigeminal regions. Thus, the distribution of the GABAergic cells showed a relatively high density in areas related to the convergence of sensory stimuli, and in zones that contain interneurons inhibiting masticatory motorneurons. The results suggest, therefore, that GABA might play an important role both in discriminative sensory processing and in reflex modulation of the orofacial region.Abbreviations RF reticular formation - FRp parvicellular reticular formation - Vc trigeminal nucleus of the spinal tract, subnucleus caudalis - Vmes mesencephalic nucleus - Vmo trigeminal motor nucleus - Vo trigeminal nucleus of the spinal tract, subnucleus oralis - Vp principal trigeminal nucleus - Vsp spinal trigeminal nucleus - Vsup supratrigeminal nucleus     

Chemoanatomical separation of vibrissal trigeminal primary afferents in the rat: a special central representation of supraorbital vibrissae

A choleratoxin B subunit transganglionic labelling technique and NPY immunohistochemistry were applied in the rat to achieve the chemoanatomical separation of myelinated vibrissal primary afferents, previously considered to be morphologically indistinguishable. Further, a special central representation pattern of supraorbital vibrissae was observed in the trigeminal brainstem nuclear complex: (1) Choleratoxin-labelled supraorbital vibrissal primary afferents terminated densely in their appropriate barrelettes in the trigeminal principal sensory nucleus, in the spinal oral subnucleus, in the caudal part of the spinal interpolar subnucleus, and in lamina IV of the caudal part of the spinal caudal subnucleus. (2) A second population of choleratoxin-labelled vibrissal afferents was also observed, terminating only in lamina III of the caudal subnucleus. (3) After peripheral nerve transection, NPY-immunoreactive supraorbital vibrissal primary afferent fibres appeared in their appropriate barrelettes in the principal sensory nucleus and the caudal part of the interpolar subnucleus, while in the caudal part of the caudal subnucleus NPY-immunoreactive vibrissal primary afferent terminals were found exclusively in the inner part of lamina II, extending over the outer part of lamina III. NPY-immunoreactive supraorbital vibrissal primary afferents were never found in the oral subnucleus. In contrast with the rules of the central representation of the mystacial (infraorbital) vibrissae, the multiple representation of the supraorbital vibrissae in the caudal subnucleus and the dense, barrelette-like terminal arborization of the choleratoxin-labelled supraorbital vibrissal primary afferents in the oral subnucleus apparently indicate an enhanced role of supraorbital vibrissae in reflexes that protect the eyes and the head from damage.     

Somatotopic organization of second order neurons of the eye muscle proprioception

Unit activity was recorded extracellularly from lamb's nucleus principalis and pars oralis of trigeminal nuclear complex following moderate manual stretching of individual extraocular muscles. The oral portion of the spinal trigeminal nucleus and the main sensory nucleus have been investigated by systematic exploration of the second-order neurons of the eye muscle proprioception. Such responses were somatotopically organized. In particular, each single extraocular muscle was represented along the main dorso-ventral axis in this manner: superior oblique and superior rectus in a dorsal layer; inferior rectus and inferior oblique in an intermediate layer, while the medial rectus and the lateral rectus were represented more ventrally. In a few experiments this representation was not observed, due to intermingling of the units. The topographic organization of eye muscle proprioception in the trigeminal nuclear complex described above closely corresponds to that reported by previous authors in the Gasser ganglion.     

Ulex europaeus agglutinin-I binding to dental primary afferent projections in the spinal trigeminal complex combined with double immunolabeling of substance P and GABA elements using peroxidase and colloidal gold

Ulex europaeus agglutinin I (UEA-I) is a plant lectin with an affinity for L-fucosyl residues in the chains of lactoseries oligosaccharides associated with medium- and smaller-diameter dorsal root ganglion neurons and their axonal processes. These enter Lissauer's tract and terminate within the superficial laminae of the spinal cord overlapping projections known to have a nociceptive function. This implies that the surface coatings of neuronal membranes may have a relationship with functional modalities. The present investigation further examined this concept by studying a neuronal projection with a nociceptive function to determine whether fucosyl-lactoseries residues were incorporated in its primary afferent terminals. Transganglionic transport of horseradish peroxidase (HRP) following injection into tooth pulp chambers was employed to demonstrate dental pulp terminals in the trigeminal spinal complex, while peroxidase and fluorescent tags were used concomitantly to stain for UEA-I. Double immunolabeling for substance P (SP) and gamma-aminobutyric acid (GABA) using peroxidase and colloidal gold allowed a comparison of the distribution of a known excitatory nociceptive transmitter with that of UEA-I binding in specific subnuclei. Synaptic interrelationships between UEA-I positive dental pulp primary afferent inputs and specific inhibitory terminals were also examined. SP immunoreactivity occurred in laminae I and outer lamina II (IIo) of subnucleus caudalis (Vc) and in the ventrolateral and lateral marginal region of the caudal half of subnucleus interpolaris (Vi), including the periobex area in which Vi is slightly overlapped on its lateral aspect by cellular elements of Vc. The adjacent interstitial nucleus (IN) also showed an intense immunoreactivity for this peptide antibody. UEA-I binding displayed a similar distribution pattern in both Vc and Vi, but extended into lamina IIi and the superficial part of Lamina III in Vc. Dental pulp terminals were found to have a comparable distribution; however, many extended into the dorsal portion of the caudal half of Vi and the ventromedial quadrant of rostral Vi. Electron-microscopic analysis showed that transganglionically labeled dental pulp terminals contained ovoid, complex membrane-bound vacuoles laden with transported HRP. The preterminal axon and synaptic membranes of those dental pulp terminals located in zones of Vc and Vi displaying an affinity for UEA-I were usually characterized by a patchy, electron-dense coating of the peroxidase tag. SP was demonstrated ultrastructurally with Protein-A colloidal gold (3-nm particles), whereas GABA immunoreactivity was revealed by the avidin-biotin-peroxidase method.(ABSTRACT TRUNCATED AT 400 WORDS)     

Neurobiology of trigeminal pain

The brainstem trigeminal complex integrates somatosensory inputs from orofacial areas and meninges. Recent studies have shown the existence of a double representation of pain within the brainstem, at the level of both caudalis and oralis subnuclei. Noxious messages are mainly conveyed by C-fibers that activate the subnucleus caudalis neurons. These neurons in turn activate the subnucleus oralis whose neurons share similar features with the deep spinal dorsal horn neurons. In contrast with the nearness of the laminar organization of the dorsal horn, the vertical organization of the trigeminal complex offers an easier access for the study of segmental mechanisms of nociceptive processing. This model allowed us to show the existence of subtle NMDA-related mechanisms of segmental nocious processing. The trigeminal complex conveys nociceptive messages to several brainstem and thalamic relays that activate a number of cortical areas responsible for pain sensations and reactions. Cortical processing is sustained by reciprocal interactions with thalamic areas and also by a direct modulation of their pre-thalamic relays. The dysfunction of these multiple modulatory mechanisms probably plays a key role in the pathophysiology of chronic trigeminal pain.     

Termination areas of primary afferent fibers of the trigeminal nerve in the rat

The cobalt-labelling technique was used to investigate the termination areas of trigeminal primary afferent fibers. The familiar somatotopic arrangement of fibers and terminals of the three divisions of the trigeminal nerve was recognized both in the spinal tract and in the nuclear complex of the trigeminus. The spinal tract could be traced as far as the 3rd cervical segment of the spinal cord where fibers crossed to the contralateral side. The different divisions of the nuclear complex could be unambiguously defined on the basis of the pattern of fiber terminations. The nucleus principalis was characterized by the even distribution of terminals in the nucleus. The nucleus spinalis was characterized by small bundles of fibers of intranuclear origin, which broke up the even distribution pattern of terminals. The presence of mesencephalic trigeminal fibers in the nucleus oralis distinguished this nucleus from the nucleus interpolaris. The nucleus caudalis was recognized on the ground of its striated structure. Primary trigeminal afferent fibers were located in the following sites: in the solitary nucleus, in the lateral part of the reticular formation, in the dorsal-column nuclei and in the superior vestibular nuclei. Primary fiber terminations could not be observed in the cerebellum.     

Intersubnuclear connections within the rat trigeminal brainstem complex

Prior intracellular recording and labeling experiments have documented local-circuit and projection neurons in the spinal trigeminal (V) nucleus with axons that arborize in more rostral and caudal spinal trigeminal subnuclei and nucleus principalis. Anterograde tracing studies were therefore carried out to assess the origin, extent, distribution, and morphology of such intersubnuclear axons in the rat trigeminal brainstem nuclear complex (TBNC). Phaseolus vulgaris leucoagglutinin (PHA-L) was used as the anterograde marker because of its high sensitivity and the morphological detail provided. Injections restricted to TBNC subnucleus caudalis resulted in dense terminal labeling in each of the more rostral ipsilateral subnuclei. Subnucleus interpolaris projected ipsilaterally and heavily to magnocellular portions of subnucleus caudalis, as well as subnucleus oralis and nucleus principalis. Nucleus principalis, on the other hand, had only a sparse projection to each of the caudal ipsilateral subnuclei. Intersubnuclear axons most frequently traveled in the deep bundles within the TBNC, the V spinal tract, and the reticular formation. They gave rise to a number of circumscribed, highly branched arbors with many boutons of the terminal and en passant types. Retrograde single- or multiple-labeling experiments assessed the cells giving rise to TBNC intersubnuclear collaterals. Horseradish peroxidase (HRP) and/or fluorescent tracer injections into the thalamus, colliculus, cerebellum, nucleus principalis, and/or subnucleus caudalis revealed large numbers of neurons in subnuclei caudalis, interpolaris, and oralis projecting to the region of nucleus principalis. Cells projecting to more caudal spinal trigeminal regions were most numerous in subnuclei interpolaris and oralis. Some cells in lamina V of subnucleus caudalis and in subnuclei interpolaris and oralis projected to thalamus and/or colliculus, as well as other TBNC subnuclei. Such collateral projections were rare in nucleus principalis and more superficial laminae of subnucleus caudalis. TBNC cells labeled by cerebellar injections were not double-labeled by tracer injections into the thalamus, colliculus, or TBNC. These findings lend generality to currently available data obtained with intracellular recording and HRP labeling methods, and suggest that most intersubnuclear axons originate in TBNC local-circuit neurons, though some originate in cells that project to midbrain and/or diencephalon.     

Intersubnuclear Connections within the Rat Trigeminal Brainstem Complex

Prior intracellular recording and labeling experiments have documented local-circuit and projection neurons in the spinal trigeminal (V) nucleus with axons that arborize in more rostral and caudal spinal trigeminal subnuclei and nucleus principalis. Anterograde tracing studies were therefore carried out to assess the origin, extent, distribution, and morphology of such intersubnuclear axons in the rat trigeminal brainstem nuclear complex (TBNC). Phaseolus vulgaris leucoagglutinin (PHA-L) was used as the anterograde marker because of its high sensitivity and the morphological detail provided. Injections restricted to TBNC subnucleus caudalis resulted in dense terminal labeling in each of the more rostral ipsilateral subnuclei. Subnucleus interpolaris projected ipsilaterally and heavily to magnocellular portions of subnucleus caudalis, as well as subnucleus oralis and nucleus principalis. Nucleus principalis, on the other hand, had only a sparse projection to each of the caudal ipsilateral subnuclei. Intersubnuclear axons most frequently traveled in the deep bundles within the TBNC, the V spinal tract, and the reticular formation. They gave rise to a number of circumscribed, highly branched arbors with many boutons of the terminal and en passant types.

Retrograde single- or multiple-labeling experiments assessed the cells giving rise to TBNC intersubnuclear collaterals. Horseradish peroxidase (HRP) and/or fluorescent tracer injections into the thalamus, colliculus, cerebellum, nucleus principalis, and/or subnucleus caudalis revealed large numbers of neurons in subnuclei caudalis, interpolaris, and oralis projecting to the region of nucleus principalis. Cells projecting to more caudal spinal trigeminal regions were most numerous in subnuclei interpolaris and oralis. Some cells in lamina V of subnucleus caudalis and in subnuclei interpolaris and oralis projected to thalamus and/or colliculus, as well as other TBNC subnuclei. Such collateral projections were rare in nucleus principalis and more superficial laminae of subnucleus caudalis. TBNC cells labeled by cerebellar injections were not double-labeled by tracer injections into the thalamus, colliculus, or TBNC.

These findings lend generality to currently available data obtained with intracellular recording and HRP labeling methods, and suggest that most intersubnuclear axons originate in TBNC local-circuit neurons, though some originate in cells that project to midbrain and/or diencephalon.     

Central representation of dental structures in the kitten, including projections to the mesencephalic trigeminal nucleus

Horseradish peroxidase (HRP) was injected into either a single maxillary or a single mandibular primary (deciduous) cuspid tooth of 8- to 10-week-old kittens. The large apex of the primary cuspid allowed for some leakage of the HRP from the pulpal chamber to the periodontal ligament (PDL). Thus, the injection procedure resulted in the application of HRP to the PDL as well as to the pulpal tissues. The transganglionic transport of HRP resulted in discrete terminal fields within the spinal trigeminal nucleus (STN) and the main sensory nucleus (MSN). These projections were clearly somatotopically organized within the STN, but less so within MSN. Within pars oralis (PO) and pars interpolaris (PI), mandibular cuspid dental structures (MdCDS) were represented in a dorsal position relative to the maxillary cuspid dental structures (MxCDS), whereas within pars caudalis (PC) and the adjacent reticular formation the somatotopic representation was not dorsoventral, but rather mediolateral, with the MdCDS represented more medially than the MxCDS. Areas of overlap between MxCDS and MdCDS were found within MSN and to a lesser degree within the superficial laminae of PC. In addition, the fiber pathway leading to labeled somata in the mesencephalic trigeminal (Mes V) nucleus was clearly identified. The majority of the fibers traced to the Mes V nucleus exited the spinal trigeminal tract at the level of the transition from PO to the MSN and traversed the nuclear region in a position dorsal to and separate from the trigeminal motor tract. As in STN, fibers within the caudal Mes V tract appeared to be somatotopically organized, with the fibers from the MdCDS generally more dorsal than the ones from the MxCDS. Labeled fibers, some with terminal arbors, were also identified in close association with the trigeminal motor tract. The findings show a complex pattern of central representation in the immature feline central nervous system for deciduous dental structures.     

Ulex europaeus Agglutinin-I Binding to Dental Primary Afferent Projections in the Spinal Trigeminal Complex Combined with Double Immunolabeling of Substance P and GABA Elements Using Peroxidase and Colloidal Gold

Ulex europaeus agglutinin I (UEA-I) is a plant lectin with an affinity for L-fucosyl residues in the chains of lactoseries oligosaccharides associated with medium and smaller-diameter dorsal root ganglion neurons and their axonal processes. These enter Lissauer's tract and terminate within the superficial laminae of the spinal cord overlapping projections known to have a nociceptive function. This implies that the surface coatings of neuronal membranes may have a relationship with functional modalities. The present investigation further examined this concept by studying a neuronal projection with a nociceptive function to determine whether fucosyl-lactoseries residues were incorporated in its primary afferent terminals. Transganglionic transport of horseradish peroxidase (HRP) following injection into tooth pulp chambers was employed to demonstrate dental pulp terminals in the trigeminal spinal complex, while peroxidase and fluorescent tags were used concomitantly to stain for UEA-I. Double immunolabeling for substance P (SP) and γ-aminobutyric acid (GABA) using peroxidase and colloidal gold allowed a comparison of the distribution of a known excitatory nociceptive transmitter with that of UEA-I binding in specific subnuclei. Synaptic interrelationships between UEA-I positive dental pulp primary afferent inputs and specific inhibitory terminals were also examined.

SP immunoreactivity occurred in laminae I and outer lamina II (II_o) of subnucleus caudalis (Vc) and in the ventrolateral and lateral marginal region of the caudal half of subnucleus interpolaris (Vi), including the periobex area in which Vi is slightly overlapped on its lateral aspect by cellular elements of Vc. The adjacent interstitial nucleus (IN) also showed an intense immunoreactivity for this peptide antibody. UEA-I binding displayed a similar distribution pattern in both Vc and Vi, but extended into lamina II; and the superficial part of Lamina III in Vc. Dental pulp terminals were found to have a comparable distribution; however, many extended into the dorsal portion of the caudal half of Vi and the ventromedial quadrant of rostral Vi.

Electron-microscopic analysis showed that transganglionically labeled dental pulp terminals contained ovoid, complex membrane-bound vacuoles laden with transported HRP. The preterminal axon and synaptic membranes of those dental pulp terminals located in zones of Vc and Vi displaying an affinity for UEA-I were usually characterized by a patchy, electron-dense coating of the peroxidase tag. SP was demonstrated ultrastructurally with Protein-A colloidal gold (3-nm particles), whereas GABA immunoreactivity was revealed by the avidin—biotin—peroxidase method. This combined approach labeled a variety of simple axodendritic to large complex scalloped dental terminals which contained SP and were shown to have a UEA-I affinity. In addition, many of the larger terminals formed contacts with GABA-ergic dendrites and received inputs from GABA-ergic synapses. These complexes were most concentrated in lamina II_o of Vc and the ventrolateral zone of Vi. Many terminals in laminae II_i; and III with a UEA-I-positive surface coating failed to bind with the antiserum for SP, indicating that other transmitters may colocalize with UEA-I and suggesting that absolute correlations between specific oligosaccharide plasmalemmal coatings and functional modalities should be approached with caution. Further studies employing antisera to different transmitters are currently underway to better define the relationship between transmitter localization and anatomical substrates within this circuitry. These studies should eventually provide additional clues about relationships between functional properties and oligosaccharide coatings of primary afferent projections.     

Collateral projections from trigeminal sensory nuclei to ventrobasal thalamus and cerebellar cortex in rats

The retrograde fluorescent labeling technique reveals that trigeminal projections to the ventroposteromedial nucleus of the thalamus (VPM) of the rat originate from the main sensory nucleus (MSN) of the trigeminal and subnuclei interpolaris (V1) and caudalis (Vc) of the spinal trigeminal nucleus. These projections are predominantly contralateral; however, the presence of a few ipsilateral labeled cells in MSN suggests an uncrossed trigeminothalamic pathway. Trigeminocerebellar fibers projecting to the paramedian lobule (PML) of the cerebellar cortex are located in Vi and caudal subnucleus oralis (Vo). This is principally an ipsilateral pathway, but several bisbenzimide-labeled cells are present in contralateral Vi. The most notable finding occurred after paired injections of Evans Blue into VPM and bisbenzimide into PML, demonstrating neurons in Vi with divergent projections to both structures. The presence of this type of projection was not found in mice (Steindler: J. Comp. Neurol. 237:155-175, 1985) and has not been reported in other species.     

Central Representation of Dental Structures in the Kitten,Including Projections to the Mesencephalic Trigeminal Nucleus

Horseradish peroxidase (HRP) was injected into either a single maxillary or a single mandibular primary (deciduous) cuspid tooth of 8- to 10-week-old kittens. The large apex of the primary cuspid allowed for some leakage of the HRP from the pulpal chamber to the periodontal ligament (PDL). Thus, the injection procedure resulted in the application of HRP to the PDL as well as to the pulpal tissues. The transganglionic transport of HRP resulted in discrete terminal fields within the spinal trigeminal nucleus (STN) and the main sensory nucleus (MSN). These projections were clearly somatotopically organized within the STN, but less so within MSN. Within pars oralis (PO) and pars interpolaris (PI), mandibular cuspid dental structures (MdCDS) were represented in a dorsal position relative to the maxillary cuspid dental structures (MxCDS), whereas within pars caudalis (PC) and the adjacent reticular formation the somatotopic representation was not dorsoventral, but rather mediolateral, with the MdCDS represented more medially than the MxCDS. Areas of overlap between MxCDS and MdCDS were found within MSN and to a lesser degree within the superficial laminae of PC. In addition, the fiber pathway leading to labeled somata in the mesencephalic trigeminal (Mes V) nucleus was clearly identified. The majority of the fibers traced to the Mes V nucleus exited the spinal trigeminal tract at the level of the transition from PO to the MSN and traversed the nuclear region in a position dorsal to and separate from the trigeminal motor tract. As in STN, fibers within the caudal Mes V tract appeared to be somatotopically organized, with the fibers from the MdCDS generally more dorsal than the ones from the MxCDS. Labeled fibers, some with terminal arbors, were also identified in close association with the trigeminal motor tract. The findings show a complex pattern of central representation in the immature feline central nervous system for deciduous dental structures.     

与咬肌运动神经元直接联系的运动前神经元在脑干的分布

目的为了定位向咬肌运动神经元投射的最后一级运动前神经元在脑干内的分布。方法注射麦芽凝集素结合的辣根过氧化物酶(WGA-HRP)至咬肌神经逆行跨突触追踪,然后通过免疫组织化学方法显示了该类神经元。结果这类神经元分布在双侧三叉上核(Vsup)、三叉神经感觉主核背侧部(Vpdm)、小细胞网状结构(PCR)和三叉神经脊束核吻侧亚核背侧部(Vodm),以及对侧三叉神经运动核(Vmo)。数量上,Vsup,特别是注射侧Vsup中,标记的神经元数量最多;其他核团内,双侧标记的神经元的数量无明显差别。结论一侧咬肌运动神经元直接接受脑干双侧多个区域调控。     

Cerebellar afferents to paramedian lobule from the trigeminal complex in Tupaia glis: a horseradish peroxidase (HRP) study

Projections from the trigeminal complex to paramedian lobule (PML) were studied in the tree shrew (Tupaia glis) by means of retrograde transport of horseradish peroxidase (HRP). Neurons which project to both dorsal and ventral folia of PML are located primarily in those areas of the trigeminal nuclear complex interpreted as nucleus interpolaris (Vi) and caudal areas of the nucleus oralis (Vo). The majority of HRP-labeled neurons lie in ventral and ventrolateral regions of Vi/Vo. No HRP-reactive cells are present in the principal (Vp), mesencephalic, or motor nuclei nor in nucleus caudalis or rostral portions of oralis. The majority of trigeminocerebellar (TC) cells are found in ipsilateral Vi; however, sparse numbers of labeled somata are present in this subnucleus on the contralateral side. Within Vi/Vo, small fusiform and medium-and large-sized multipolar neurons contain HRP-reaction product. Large multipolar cells are found primarily in ventrolateral portions of Vi/Vo, while medium and small neurons are scattered throughout the ventral half of the nucleus. Small-sized neurons are also present dorsally within Vi/Vo. Axons of labeled TC cells course laterally through the spinal trigeminal tract, enter medial aspects of the restiform body, and arch dorsally into the cerebellum.     

Cell structure and response properties in the trigeminal subnucleus oralis

Extra- and intracellular recording, electrical stimulation, receptive field mapping, and horseradish peroxidase injection techniques were used to study the structure of functionally identified neurons in trigeminal (V) brainstem subnucleus oralis of the rat. Of 15 heavily labeled cells located within oralis, 4 were local-circuit neurons with receptive fields restricted to either an incisor, guard hairs, one vibrissa, or deep facial tissue (nociceptors). Their morphologies were highly varied, with expansive and spiny dendritic trees and recurrent and intersubnuclear axon collaterals. Oralis local-circuit neurons therefore most closely resembled non-vibrissa-sensitive local-circuit cells in adjacent subnucleus interpolaris. Six other stained cells projected to contralateral thalamus, and two others projected to ipsilateral cerebellum. They typically had intramodality convergent receptive fields (i.e., spanning more than one receptor organ, such as multiple vibrissae or teeth) with widespread dendritic trees, and were therefore indistinguishable from similarly projecting cells in interpolaris. Two other cells projected to the ipsilateral spinal cord, as well as other V brainstem subnuclei. One of these responded to high-threshold mechanical stimulation of teeth; the other was discharged by deflection of one mystacial vibrissa. Their dendrites were very widespread and ended in spiny and bulbous appendages. Local axon collaterals were also extensive. The remaining oralis cell had two axons, one projecting to the thalamus, the other to the spinal cord. Its receptive field expressed convergence from multiple receptor organs, including vibrissae, guard hairs, and skin. Its somadendritic morphology was similar to that of oralis cells projecting only to thalamus. We conclude that, with some exceptions, the extensive dendritic trees, axon branching, convergence, and functional diversity of oralis cells approximate those described previously for functionally equivalent neurons in interpolaris (Jacquin et al., 1989a,b). Such anatomical and physiological properties are rarely seen, however, in nucleus principalis (Jacquin et al., 1988a). The structure and function of three atypical principalis cells with structural and functional characteristics typical of oralis cells are also described. It is argued that such cells are rostrally displaced oralis cells.     

Localization and Central Projections of Primary Afferent Neurons That Innervate the Temporomandibular Joint in Cats

Primary afferent neurons that innervate the temporomandibular joint (TMJ) in cats were labeled by injecting a 2-5% solution of wheatgerm agglutinin bound to horseradish peroxidase into the joint capsule and capsular tissues in 14 cats and processing the brain stem and trigeminal ganglia using the tetramethylbenzidine method described by Mesulam (1978). The perikarya of ganglion cells that innervate the TMJ ranged in diameter from 15 to 109 μm and were primarily located in the posterolateral portion of the trigeminal ganglion. The central processes of these neurons entered the brain stem in middle pons and were distributed to all portions of the sensory trigeminal nuclei. However, the majority of labeled fibers and greatest density of terminal labeling were observed in the dorsal part of the main sensory nucleus and the subnucleus oralis of the spinal trigeminal nucleus. Very few labeled fibers were observed in the spinal tract of the trigeminal nerve below the obex. However, evidence for axon terminals was consistently observed in laminae I, II, and III of the medullary dorsal horn. These findings concur with physiological evidence showing that information from the TMJ influences neurons in rostral (Kawamura et al, 1967) and in caudal (Broton et al, 1985) portions of the trigeminal sensory nuclei.     

Central Projections of Identified Trigeminal Primary Afferents after Molar Pulp Deafferentation in Adult Rats

It is known that removal of the tooth pulp from mandibular molar teeth in adult rats alters the mechanoreceptive field properties of many low-threshold mechanoreceptive neurons in the trigeminal brainstem nuclear complex. The present study investigates one possible way that such deafferentation-induced receptive field changes could occur: altered central projections of uninjured trigeminal low-threshold mechanoreceptive primary afferent fibers. Intra-axonal injection of horseradish peroxidase (n = 22) or neurobiotin (n = 44) into characterized fibers was performed ipsilateral to, and 10–32 days after, removal of the coronal pulp from the left mandibular molars in adult rats. Collaterals were reconstructed, quantified, and compared by means of multivariate analyses of variance to equivalent fibers stained in normal adult rats.

Stained mechanosensitive fibers from experimental animals were rapidly conducting and responded to light mechanical stimulation of one vibrissa, one tooth, oral mucosa, facial hairy skin, or guard hairs. Their central projections were indistinguishable from those of control axons in all four trigeminal subnuclei. The numbers of collaterals, areas subtended by collateral arbors, numbers of boutons per collateral, and arbor circularity did not differ from those of control afferents. Collateral somatotopy was also unaffected.

These data suggest that following pulpotomy, the central collaterals of uninjured trigeminal afferents display normal morphologies and maintain normal somatotopy. Changes in the morphology of low-threshold primary afferents cannot account for the changes that occur in the receptive field properties of trigeminal brainstem neurons after pulp deafferentation.     

The Central Projections of the Monkey Tooth Pulp Afferent Neurons

Transganglionic transport of horseradish peroxidase conjugated to wheatgerm agglutinin (HRP:WGA) entrapped in hypoallergenic polyacrylamide gel was used to study the patterns of termination of primary afferents that innervate the upper and lower tooth pulps within the trigeminal sensory nuclear complex (TSNC) of the monkey. HRP:WGA injections were also made into the lower incisors and molars, in order to examine the topographic arrangement of pulpal afferent projections. HRP-labeled pulpal afferents innervating lower and upper teeth projected ipsilaterally to the rostral subnucleus dorsalis (Vpd) and caudal subnucleus ventralis (Vpv) of the nucleus principalis (Vp); the rostrodorsomedial (Vo.r) and dorsomedial (Vo.dm) subdivisions of the nucleus oralis (Vo); the dorsomedial subdivision of the nucleus interpolaris (Vi); and laminae I—II and/or V of the nucleus caudalis (Vc) at its rostralmost level. The HRP-labeled terminals from upper and lower pulpal afferents formed a rostrocaudal column from the midlevel of Vp to the rostral tip of Vc. The label in Vp and Vo was considerably dense, but the column of terminals was interrupted at the Vpd-Vpv transition. The label in Vi and Vc was much less dense compared to that in the rostral nuclei, and the column of terminals was interrupted frequently. The representation of the upper and lower teeth in TSNC was organized in a somatotopic fashion that varied from one subdivision to the next, though their terminal zones overlapped within Vpd. The upper and lower teeth were represented in Vpv, Vo.r, Vo.dm, Vi, and Vc in a ventrodorsal, dorsoventral, lateromedial, lateromedial, and lateromedial sequence, respectively. Topographic arrangement was also noticed for the projections of pulpal afferents from the lower incisors and molars: The representations of the lower incisors and molars in Vpv, Vo.r, Vo.dm, Vi, and Vc were organized in a lateromedial, dorsoventral, ventrodorsal, ventrodorsal, and lateromedial sequence, respectively. The present results indicating sparse projections from pulpal afferents in the monkey's Vc are in good correspondence with a clinical report that trigeminal tractotomy just rostral to the obex has no significant effect on dental pain perception in patients. Furthermore, the present study indicates that projection patterns of pulpal afferents—which include the termination sites, the density of terminations between nuclei, and topographic arrangement—differ among animal species.