Oral and facial representation within the medullary and upper cervical dorsal horns in the cat

Transganglionic transport of HRP was used to study the patterns of termination of somatic afferent fibers innervating oral and facial structures within the trigeminal nucleus caudalis and upper cervical dorsal horn of the cat. In separate animals, the superior alveolar, pterygopalatine, buccal, inferior alveolar, lingual, frontal, corneal, zygomatic, infraorbital, mental, mylohyoid, and auriculotemporal branches of the trigeminal nerve were traced in this experiment. The organization of the primary afferents innervating the oral structures is not uniform across laminae and at different rostrocaudal levels of the nucleus caudalis. The superior alveolar and pterygopalatine nerves mainly terminate in laminae I, II, and V at the level of the rostral one-third of the caudalis. By contrast, the lingual, inferior alveolar, and buccal nerve terminate in laminae I-V of, respectively, the rostral third, the entire length, and caudal two-thirds of the caudalis. In addition, the lingual, buccal, and pterygopalatine nerves terminate in the dorsal and middle parts of the interstitial islands or pockets of lamina I neuropil extending to the rostral levels parallel to the nucleus interpolaris. Mediolaterally, in laminae I, II, and V of the rostral third an extensive overlap of projections was found between the branches from each trigeminal division, and some overlap was observed between projections from the mandibular and maxillary divisions. On the other hand, the projections of primary afferents innervating the facial structures are arranged in a somatotopic fashion in rostrocaudal and mediolateral axes over the laminae (I-IV) through the nucleus caudalis and upper cervical dorsal horn. Fibers from the perioral and perinasal regions terminate most rostrally in caudalis, and fibers from progressively more posterior facial regions terminate at successively lower levels. A mediolateral somatotopic arrangement was observed, with fibers from the ventral parts of face ending in the medial regions and fibers from the progressively more dorsal parts of the face ending in successively more lateral regions of the medullary and upper cervical dorsal horns. Corneal afferent terminals are concentrated in the outer parts of lamina II at the levels of the rostral parts of the caudal two-thirds of the caudalis and the interstitial islands of lamina I. The maxillary division terminates first at the most caudal level of the caudalis, followed by the ophthalmic division descending as far as the C2 segment and the mandibular division reaching the most caudal level of the C2 segment.(ABSTRACT TRUNCATED AT 400 WORDS)     

Corneal sensory pathway in the rat: a horseradish peroxidase tracing study

The methods of transganglionic transport of horseradish peroxidase (HRP) and horseradish peroxidase--wheat germ agglutinin (HRP-WGA) were used to determine the location within the trigeminal ganglion of the primary afferent neurons that innervate the rat central cornea, and the brainstem and spinal cord termination sites of these cells. In each of 18 animals, solutions of HRP or HRP-WGA were applied to the scarified corneal surface and allowed to infiltrate into the corneal epithelium and stroma for 15 minutes. Postmortem examination of the corneal whole mounts from the experimental animals, and of corneas and neural tissues from several control animals, showed that the HRP/HRP-WGA remained confined to the central cornea with no spread into adjacent intra- or extraorbital tissues. HRP-labeled corneal afferent somata were located in the dorsal part of the ophthalmic region of the ipsilateral trigeminal ganglion. The central fibers of the corneal afferent neurons projected very heavily to interstitial nuclei of Cajal in the spinal tract of V at the level of caudal pars interpolaris and rostral pars caudalis, lightly to the pars caudalis/C1 transition zone, and sparsely to the dorsal horn of spinal cord segments C1-C3. The trigeminal main sensory nucleus, pars oralis, the rostral three-fourths of pars interpolaris, and an extensive midregion of pars caudalis were totally devoid of reaction product. Terminal fields in caudal pars caudalis and in the spinal cord dorsal horn were concentrated largely in the outer half of lamina II, with lesser accumulations in lamina I, the deeper half of lamina II, and in lamina III. The present study demonstrates for the first time by means of an anatomical tracing procedure the brainstem termination sites of corneal afferent neurons in the rat. The patchy, discontinuous nature of the corneal afferent projection to the caudal trigeminal brainstem nuclear complex (TBNC), and the total lack of corneal projections to rostral subdivisions of the TBNC, provide an exception to the general rule of trigeminal organization in which most areas of the head and face are represented as continuous columns throughout the rostrocaudal extent of the ipsilateral TBNC.     

Oral and facial representation in the trigeminal principal and rostral spinal nuclei of the cat

Transganglionic transport of horseradish peroxidase (HRP) was used to study the patterns of termination of somatic afferent fibers innervating oral and facial structures within the principal nucleus (Vp), nucleus oralis (Vo), and nucleus interpolaris (Vi). The primary trigeminal afferent fibers that innervate the oral cavity supplied by the pterygopalatine, superior alveolar, lingual, buccal, and inferior alveolar branches, as well as the facial skin supplied by the frontal, corneal, zygomatic, infraorbital, auriculotemporal, mylohyoid, and mental branches, were traced in this experiment. The results show that trigeminal afferent nerves that innervate the oral cavity project mainly to the principal nucleus, the rostrodorsomedial part (Vo.r) and dorsomedial division (Vo.dm) of pars oralis, and the dorsomedial region of pars interpolaris, while an extensive overlap of projections is found in the Vo.r, Vo.dm, and rostral Vi. The central processes of fibers innervating the anterior face (i.e., mental, infraorbital, and frontal nerves) terminate in the ventral division of principalis (Vpv), caudal region pars oralis (Vo.c), and ventrolateral Vi, with the largest numbers of terminals being found in the Vpv and Vi. In contrast, the central projection patterns of the corneal, zygomatic, mylohyoid, and auriculotemporal afferents are different from those of other afferent nerves examined, and present a discrete projection to the trigeminal sensory nuclear complex (TSNC). The corneal, mylohyoid, and auriculotemporal afferents mainly project to the restricted regions of principalis and caudal Vi, while zygomatic afferent nerve fibers project to the caudal third of pars interpolaris. The typical somatotopic organization with the face of the mouth open inverted is represented in the rostrocaudal midlevels of the Vpv and caudal pars interpolaris. The Vpd receives topographical projection from primary afferent nerves that innervate the oral structure only, while this projection was organized in a complicated manner. The relationship between the functional segregation and the cytoarchitectonic differentiation of the TSNC is discussed, particularly with respect to this somatotopic organization, combined with the characteristics of projecting cells in the TSNC.     

Topographic organization of central terminal region of different sensory branches of the rat mandibular nerve

The central projection of primary neurons comprising the auriculotemporal nerve, cutaneous branch of the mylohyoid nerve, inferior alveolar nerve, mental nerve, lingual nerve, and buccal nerve was investigated using transganglionic transport of HRP in young rats. In view of the topographic organization of central projection fields, the nerves were divided into two groups; i.e., those projecting to the dorsolateral margin of the trigeminal nucleus principalis, subnucleus oralis, and interpolaris (the auriculotemporal, mylohyoid, and mental nerves) and those projecting more medially (the inferior alveolar, lingual, and buccal nerves). The former group of nerves projected more caudally than the latter in the medullary and spinal dorsal horn complex rostral to the 3rd cervical segment, in general. Furthermore, the latter group projected to the nucleus of the solitary tract and the supratrigeminal and paratrigeminal nuclei, whereas the other nerves did not. The data indicate the following points: Primary neurons innervating the intraoral structures terminate medial (in trigeminal nucleus principalis and subnucleus oralis) and ventral (in subnucleus interpolaris) to the terminal fields of those innervating the facial skin. Primary neurons innervating the intraoral structures project to the nucleus of the solitary tract and the supra- and paratrigeminal nuclei, whereas those innervating the facial skin do not. Primary neurons innervating the periphery of the face project to the spinal dorsal horn and those innervating the intra/perioral region project to medullary dorsal horn, though this segregation from the medulla to the 3rd cervical segment is relatively loose. Only those trigeminal primary neurons, whose receptive fields extend to or beyond the midline, project to the contralateral dorsal horn from the medulla to the 3rd cervical segment.     

Topographic representation of lower and upper teeth within the trigeminal sensory nuclei of adult cat as demonstrated by the transganglionic transport of horseradish peroxidase

Transganglionic transport of horseradish peroxidase-wheat germ agglutinin conjugate (HRP-WGA) entrapped in hypoallergenic polyacrylamide gel was used to study the patterns of termination of primary afferents that innervate the lower and upper tooth pulps within the trigeminal sensory nuclear complex (TSNC). HRP injections were made into the inferior and superior alveolar nerves in order to compare the central projections of the whole nerve with those from tooth pulps. In addition, the relationship between the distribution of the trigeminothalamic tract cells and the projection sites of the tooth pulp afferents was investigated by injecting HRP into the posterior ventral thalamus. HRP-labeled tooth pulp afferent fibers innervating the lower and upper teeth projected to the subnucleus dorsalis (Vpd) of pars principalis, the rostrodorsomedial part (Vo.r) and nucleus dorsomedialis (Vo.dm) of pars oralis, the medial regions of pars interpolaris, and laminae I, II, and V of pars caudalis. Terminal fields of the lower tooth pulp afferents formed a rostrocaudally running, uninterrupted column from the midlevel of Vpd to the caudal tip of caudalis. In contrast, the column of termination of upper tooth pulp afferents was discontinuous at the Vpd/Vo.r transition, and ended at the more rostral level of the caudalis than that of the lower tooth pulp afferents. The representation of the lower and upper teeth in the TSNC was organized in a somatotopic fashion which varied from one subdivision to the next, although terminal zones of the inferior and superior alveolar nerves overlapped within the Vo.r, Vo.dm, and dorsomedial part of rostral pars interpolaris. The lower and upper teeth were represented in the Vpd, Vo.r, Vo.dm, medial region of pars interpolaris, and laminae I, II, and V, in a ventrodorsal or caudorostral, dorsoventral, lateromedial, dorsoventral, and mediolateral or dorsomedial-ventrolateral sequence, respectively. The smaller, more focal terminal areas of the teeth contrasted sharply with more extensive terminal fields of the alveolar nerves. The HRP injections within the thalamus indicated that neurons in Vpd, the caudal pars interpolaris, and laminae I/V of caudalis, which are subdivisions of TSNC that receive pulpal projections, sent their axons to the ipsilateral and contralateral posterior ventral thalamus.(ABSTRACT TRUNCATED AT 400 WORDS)     

Contralateral projections of trigeminal mandibular primary afferents in the guinea pig as seen by transganglionic transport of horseradish peroxidase

Transganglionic transport of horseradish peroxidase (HRP) was used to investigate contralateral projections of trigeminal mandibular fibers in the guinea pig. After application of HRP to the buccal, lingual, auriculotemporal, mylohyoid, mental and inferior alveolar nerves, crossing fibers and contralateral endings were found in the caudal region of the nucleus of the solitary tract (most of these belonging to the buccal and lingual nerves), the dorsomedial region of the subnucleus caudalis of the trigeminal sensory nuclear complex (TSNC), and the dorsal horns of the first 5 cervical spinal cord segments (C1-C5). The greatest numbers of crossing fibers in the medullary and cervical dorsal horn segments belonged to the mental and mylohyoid nerves, though these nerves did not project contralaterally to C4-C5. Contralateral buccal and lingual endings were scattered sparsely from the subnucleus caudalis to C5, and only very few contralateral auriculotemporal terminals were observed. Though laminae I-V of the dorsomedial region of the medullary and cervical dorsal horns all exhibited contralateral endings of the mental and mylohyoid nerves, most such endings were found in laminae IIi-III, followed by lamina IV, which suggests their involvement in the reception of mechanical stimuli and in the sensory motor reflexes of the orofacial region. The contralateral buccal and lingual terminals were distributed somatotopically in the first 5 cervical cord segments, with the lingual endings rostral to the buccal terminals within each segment. In C4 and C5 lingual endings appeared exclusively in laminae I and IIo, suggesting that like the ipsilateral lingual projections at this level, which also terminate in these laminae, they may be involved in pain and temperature sensation.     

The central projections of tooth pulp afferent neurons in the rat as determined by the transganglionic transport of horseradish peroxidase

Transganglionic transport of horseradish peroxidase (HRP) or horseradish peroxidase-wheat germ agglutinin conjugate (HRP-WGA) was used to map in detail the central projections of trigeminal primary afferent neurons that innervate the dental pulp organ of the rat. In each of ten animals, 0.5-2.0 microliters of enzyme solution was injected into the pulp chamber of the first maxillary molar tooth. Postmortem examination of the decalcified teeth in all cases showed that the HRP/HRP-WGA remained confined to the pulp chamber and pulp roots, with no spread of enzyme into periapical tissues. HRP-labeled tooth pulp afferent fibers projected to all four rostrocaudal subdivisions of the ipsilateral trigeminal brainstem nuclear complex (TBNC) and to the upper cervical spinal cord. The labeled terminal fields formed a column that stretched relatively uninterrupted from just caudal to the rostromedial tip of the trigeminal principal sensory nucleus to at least the C2 segment of the spinal cord. The density of the afferent projection varied markedly from one rostrocaudal level of the TBNC to the next but was heaviest in an area encompassing the caudal one-half of the principal sensory nucleus and the rostral two-thirds of pars oralis. Fibers projected only lightly to pars caudalis, where they terminated preferentially in laminae I, IIa, and the junctional zone between laminae IV and V. HRP-labeled terminals in C1 and C2 were located almost exclusively in laminae I. In the dorsoventral axis, the terminal fields in the TBNC were located in a surprisingly dorsal part of the complex, well within what has been shown by others to be largely an area of termination for mandibular division fibers. Most fibers ended in medial parts of the TBNC, with the exception of two modestly labeled terminal fields located in the lateral aspects of rostral pars oralis and rostral pars caudalis. No labeled fibers terminated in the contralateral TBNC or contralateral cervical spinal cord.     

Distribution of gastrin‐releasing peptide in the rat trigeminal and spinal somatosensory systems

Gastrin‐releasing peptide (GRP) has recently been identified as an itch‐specific neuropeptide in the spinal sensory system in mice, but there are no reports of the expression and distribution of GRP in the trigeminal sensory system in mammals. We characterized and compared GRP‐immunoreactive (ir) neurons in the trigeminal ganglion (TG) with those in the rat spinal dorsal root ganglion (DRG). GRP immunoreactivity was expressed in 12% of TG and 6% of DRG neurons and was restricted to the small‐ and medium‐sized type cells. In both the TG and DRG, many GRP‐ir neurons also expressed substance P and calcitonin gene‐related peptide, but not isolectin B₄. The different proportions of GRP and transient receptor potential vanilloid 1 double‐positive neurons in the TG and DRG imply that itch sensations via the TG and DRG pathways are transmitted through distinct mechanisms. The distribution of the axon terminals of GRP‐ir primary afferents and their synaptic connectivity with the rat trigeminal sensory nuclei and spinal dorsal horn were investigated by using light and electron microscopic histochemistry. Although GRP‐ir fibers were rarely observed in the trigeminal sensory nucleus principalis, oralis, and interpolaris, they were predominant in the superficial layers of the trigeminal sensory nucleus caudalis (Vc), similar to the spinal dorsal horn. Ultrastructural analysis revealed that GRP‐ir terminals contained clear microvesicles and large dense‐cored vesicles, and formed asymmetric synaptic contacts with a few dendrites in the Vc and spinal dorsal horn. These results suggest that GRP‐dependent orofacial and spinal pruriceptive inputs are processed mainly in the superficial laminae of the Vc and spinal dorsal horn. J. Comp. Neurol. 522:1858–1873, 2014. © 2013 Wiley Periodicals, Inc.     

Cells of origin of the trigeminohypothalamic tract in the rat

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

An HRP study of the central projections of primary trigeminal neurons which innervate tooth pulps in the cat

Trigeminal ganglia and brain stem of adult cats were studied following HRP injections into tooth pulps or after exposure of the cut end of the inferior alveolar nerve to HRP. Ipsilateral ganglion cells within a wide range of sizes were labeled in both experimental situations, whereas no labeled cells were observed in the contralateral ganglion in any animal. Labeled central branches of tooth pulp and inferior alveolar neurons were observed in all subdivisions of the ipsilateral trigeminal sensory complex. Terminal labeling in the tooth pulp experiments was confined to the dorsomedial parts of the main sensory nucleus and subnuclei oralis and interpolaris. Caudal to the obex terminal labeling was restricted to the medial halves of laminae I, IIa and V of the medullary dorsal horn. In the inferior alveolar nerve experiments dense terminal labeling was observed in the dorsal parts of the main sensory nucleus and subnuclei oralis and interpolaris. Caudal to the obex terminal labeling was located throughout laminae I to V in contrast to the tooth pulp experiments. Neither of the two experimental situations offers any evidence for a bilateral or contralateral brain stem projection of primary trigeminal neurons.     

The central projection of masticatory afferent fibers to the trigeminal sensory nuclear complex and upper cervical spinal cord

Retrograde and anterograde transport of horseradish peroxidase-wheat germ agglutinin (HRP-WGA) conjugate was used to study the organization of primary afferent neurons innervating the masticatory muscles. HRP applied to the nerves of jaw-closing muscles--the deep temporal (DT), masseter (Ma), and medial pterygoid (MP)--labeled cells in the trigeminal ganglion and the mesencephalic trigeminal nucleus (Vmes), whereas HRP applied to nerves of the jaw-opening muscles--anterior digastric (AD) and mylohyoid (My)--labeled cells only in the trigeminal ganglion. Cell bodies innervating the jaw-closing muscles were found with greater frequency in the intermediate region of the mandibular subdivision, while somata supplying the jaw-opening muscles were predominant posterolaterally. The distribution of their somatic sizes was unimodal and limited to a subpopulation of smaller cells. Projections of the muscle afferents of ganglionic origin to the trigeminal sensory nuclear complex (TSNC) were confined primarily to the caudal half of pars interpolaris (Vi), and the medullary and upper cervical dorsal horns. In the Vi, Ma, MP, AD, and My nerves terminated in the lateral-most part of the nucleus with an extensive overlap in projections, save for the DT nerve, which projected to the interstitial nucleus or paratrigeminal nucleus. In the medullary and upper cervical dorsal horns, the main terminal fields of individual branches were confined to laminae I/V, but the density of the terminals in lamina V was very sparse. The rostrocaudal extent of the terminal field in lamina I differed among the muscle afferents of origin, whereas in the mediolateral or dorsoventral axis, a remarkable overlap in projections was noted between or among muscle afferents. The terminals of DT afferents were most broadly extended from the rostral level of the pars caudalis to the C3 segment, whereas the MP nerve showed limited projection to the middle one-third of the pars caudalis. Terminal fields of the Ma, AD, and My nerves appeared in the caudal two-thirds of the pars caudalis including the first two cervical segments, the caudal half of the pars caudalis and the C1 segment, and in the caudal part of the pars caudalis including the rostral C1 segment, respectively. This rostrocaudal arrangement in the projections of muscle nerves, which corresponds to the anteroposterior length of the muscles and their positions, indicates that representation of the masticatory muscles in lamina I reflects an onion-skin organization. These results suggest that primary muscle afferent neurons of ganglionic origin primarily mediate muscle pain.(ABSTRACT TRUNCATED AT 400 WORDS)     

Central projections and somatotopic organisation of trigeminal primary afferents in pigeon (Columba livia)

Injections of cholera toxin B-chain conjugated to horseradish peroxidase into individual peripheral branches of the trigeminal nerve or into the trigeminal ganglion showed that an ascending trigeminal tract (TTA) terminated in distinct ventral and dorsal divisions of the principal sensory nucleus (PrVv and PrVd, respectively), and a descending tract (TTD) terminated within pars oralis, pars interpolaris, and pars caudalis divisions of the nucleus of TTD (nTTD) and within the dorsal horn of the first six cervical spinal segments. In PrVD, mandibular, ophthalmic, and maxillary projections were predominantly located dorsally, ventrally, and medially, respectively. In nTTD, mandibular projections lay dorsomedially, ophthalmic projections lay ventrolaterally, and maxillary projections lay in between. At caudal medullary and spinal levels, mandibular projections were situated medially, ophthalmic projections were situated laterally, and maxillary projections were situated centrally. The terminations within the dorsal horn were most dense in laminae III and IV and were least dense in lamina II, with laminae III-IV also receiving topographically organised contralateral projections. Extratrigeminal projections were mainly to the external cuneate nucleus by way of a lateral descending trigeminal tract (ITTD; Dubbeldam and Karten [1978] J. Comp. Neurol. 180:661–678) and to the region of the tract of Lissauer and lamina I of the dorsal horn. Other projections were to a region medial to the apex of pars interpolaris, to the nuclei ventrolateralis anterior (Vla) and presulcalis anterior (Pas) of the solitary complex, and sparsely to the lateral reticular formation (plexus of Horsley) ventral to TTD. No projections were seen to the trigeminal motor nuclei or to the cerebellum. © 1996 Wiley-Liss, Inc.     

Central projections of primary sensory neurons innervating different parts of the vibrissae follicles and intervibrissal skin on the mystacial pad of the rat.

The cell bodies and central projections of neurons innervating the vibrissae follicles and adjacent skin in the rat were investigated by retrograde and transganglionic transport of HRP. The cell bodies of neurons innervating the vibrissa follicle via the deep vibrissa nerve (DVN) were the largest, followed by those innervating the follicle via the superficial vibrissa nerve (SVN). The smallest cell bodies were those innervating the intervibrissal skin. The DVN neurons terminated centrally as an almost uninterrupted column through the trigeminal sensory nuclear complex. The DVN projections to nucleus caudalis and C1 dorsal horn were entirely restricted to laminae III, IV, and V. Besides the projections to lamina V, the DVN projections were strictly localized somatotopically at all levels replicating the peripheral organization of the vibrissae. The SVNs projected sparsely to midlevels of the main sensory nucleus but not to nuclei oralis and interpolaris. The main SVN projections appeared in laminae I-III of nucleus caudalis. In addition, a small projection to lamina V was observed. The projections to laminae II and III were organized mediolaterally in a similar way as the DVN projections; those to laminae I and V were less restricted. The intervibrissal skin neurons projected sparsely to the caudal main sensory nucleus and to the border between nuclei oralis and interpolaris. The projections to nucleus caudalis were restricted to laminae I-III and V and were organized in a similar way as the SVN projections.     

The central projections of trigeminal primary afferent neurons in the cat as determined by the tranganglionic transport of horseradish peroxidase

The central projections of five peripheral branches of the trigeminal nerve were investigated by the method of transganglionic transport of horseradish peroxidase (HRP). In separate animals, the corneal, supraorbital, infraorbital, mental, or inferior alveolar branches were transected and soaked in concentrated solutions of HRP. Forty-eight to 72 hours after surgery, the brainstem, upper cervical spinal cord, and trigeminal ganglia were perfusion-fixed and processed according to the tetramethylbenzidine technique. The results show that trigeminal primary afferent neurons which innervate the cornea project mainly to the levels of caudal pars interpolaris and caudal pars caudalis. In contrast, trigeminal primary afferent neurons whose peripheral processes course through the supraorbital, infraorbital, or mental nerves project most heavily to the trigeminal main sensory nucleus, pars interpolaris, and the rostrocaudal middle three-fifths of pars caudalis. Trigeminal primary afferent neurons which give origin to the inferior alveolar nerve project heavily and in approximately equal numbers of all rostrocaudal levels of the trigeminal brainstem nuclear complex (TBNC). A small number of fibers from each of the latter four cell populations project directly to the contralateral C1-C2 dorsal horn. A small number of fibers from each cell population studied end in the reticular formation immediately adjacent to the spinal nucleus of V. It is concluded that the cornea and facial skin regions of the cat are represented nonuniformly along the rostrocaudal length of the TBNC.     

Development and plasticity in hamster trigeminal primary afferent projections

At birth (gestational day 16), the hamster infraorbital nerve projects to the appropriate portion of the brainstem, though the projection lacks adult-like internal organization (patchiness). Infraorbital nerve damage at this time does not produce appreciable transganglionic atrophy in the central projections of the infraorbital nerve, but it does result in a failure to develop normal infraorbital primary afferent patches. Such damage also produces a more widespread central projection of spared mandibular afferents into regions occupied by 'regenerate' infraorbital terminals (J. Comp. Neurol., 235 (1985) 129-143). In the present study, transganglionic transport techniques were again used to show that, by postnatal day 5 (gestational day 21), rostrocaudally continuous aggregates of horseradish peroxidase-labelled infraorbital terminals are visible throughout the trigeminal brainstem nuclear complex. This aggregation pattern is nearly adult-like and isomorphic with the distribution of the mystacial vibrissae on the face. A similar infraorbital lesion performed on postnatal day 5, however, markedly decreased the density of the adult central projection of the infraorbital nerve to subnuclei principalis, oralis, interpolaris, and the magnocellular laminae of caudalis. The projection to superficial laminae of caudalis and the cervical dorsal horn was maintained. A postnatal-day-5 infraorbital lesion also failed to produce a more widespread central projection from spared mandibular primary afferents. These data suggest a relationship between the postnatal maturity of trigeminal primary afferents and the response of damaged and undamaged trigeminal afferents to infraorbital nerve transection in hamster. The similarity in the central primary afferent response to lesions at equivalent gestational times (postnatal days 5 and 0, respectively) in hamster and rat, suggests that this plasticity gradient may be a general characteristic of mammalian trigeminal primary afferents.     

Central projections and trigeminal ganglion location of corneal afferent neurons in the monkey, Macaca fascicularis

The method of transganglionic transport of horseradish peroxidase-wheat germ agglutinin conjugate (HRP-WGA) was used to determine the location within the monkey trigeminal ganglion of the primary afferent neurons that innervate the cornea, and the brainstem and spinal cord termination sites of these cells. In each of four animals. Gelfoam pledgets were saturated with 2% HRP-WGA in saline and applied to the scratched surface of the central cornea for 30 minutes. Postmortem examination of the corneal whole mounts revealed that the tracer solution remained confined to approximately the central one-fourth of the cornea with no spread into the peripheral cornea or limbus. Seventy-two to 96 hours after tracer application, 126-242 labeled cell bodies were observed in the medial region of the ipsilateral trigeminal ganglion. The majority of neurons were concentrated in an area of the ganglion that lay directly caudal to the entering fibers of the ophthalmic nerve, but smaller numbers of cells lay somewhat more laterally, near the region where the ophthalmic and maxillary nerves come together. A very small number of neurons in one animal innervated the cornea by sending their fibers into the maxillary nerve. HRP-WGA-labeled terminal fields were present to some extent in all four major rostrocaudal subdivisions of the ipsilateral trigeminal brainstem nuclear complex (TBNC), but the size of the terminal fields and the intensity of labeling differed markedly from one level of the TBNC to the next. Labeled fibers projected heavily to the transitional zone between caudal pars interpolaris and rostral pars caudalis (i.e., the "periobex" region of the TBNC) and moderately to the trigeminal main sensory nucleus, pars oralis, and caudal pars caudalis at the level of the pyramidal decussation. Remaining areas of the TBNC, including rostral pars interpolaris and the midlevel of pars caudalis, received few, if any, corneal afferent projections. Occasional labeled fibers were observed in the dorsal horn of C1 and in the rostral half of C2. It is hoped that data generated in the current investigation of nonhuman primates will contribute to a better understanding of the neural substrates that subserve corneal sensation and the blink reflex in humans.     

Differential terminal distribution of single large cutaneous afferent fibers in the spinal trigeminal nucleus and in the cervical spinal dorsal horn

Large myelinated cutaneous afferent fibers innervating the face were intra-axonally stained with HRP. The terminal arborizations of collaterals given off from single axons were confined to the outer part of nucleus subnuclei oralis and interpolaris, to layers IV and V in the subnucleus caudalis, and mainly to layers III and IV in the cervical dorsal horn.     

Representation of upper and lower primary teeth in the trigeminal sensory nuclear complex in the young dog

Transganglionic transport of horseradish peroxidase-wheat germ agglutinin conjugate entrapped in polyacrylamide gel was used to study the patterns of termination of primary afferents that innervate the upper and lower primary tooth pulps within the trigeminal sensory nuclear complex of the young dog. The lower and upper primary tooth pulp afferents projected to the subnucleus dorsalis of the principal nucleus, the rostrodorsomedial part and subnucleus dorsomedialios (Vo.dm) of the parts oralis, the nucleus of the intermediate plexus (Vi.ip) of the pars interpolaris, and laminae I, II and V of the caudalis. The lower and upper primary teeth were topographically represented in the Vo.dm, rostrocaudal mid-levels of Vi.ip and in laminae I/V of the caudal levels of the pars caudalis, whereas an extensive overlapped projection was seen in other subdivisions.     

Trigeminal primary afferents project bilaterally to dorsal horn and ipsilaterally to cerebellum, reticular formation, and cuneate, solitary, supratrigeminal and vagal nuclei

Horseradish peroxidase (HRP) applied to the transected mandibular division of the trigeminal ganglion was transported anterogradely to previously well known primary afferent terminal zones in the dorsal brainstem trigeminal nuclear complex of the rat, and retrogradely to cell bodies in the trigeminal motor, supratrigeminal and mesencephalic nuclei. Primary trigeminal afferents were also visible in the ipsilateral cerebellar cortex and paraflocculus, and the dentate, cuneate, solitary, supratrigeminal, and dorsal motor vagal nuclei, parvicellular reticular formation, area postrema, and C1–C6 dorsal horn. The contralateral medulla and cervical dorsal horn were also innervated by primary afferents which crossed in the posterior commissure. These projections were also labeled when HRP was applied to individual sensory branches of the mandibular nerve.     

Distribution and central projections of primary afferent neurons that innervate the masseter muscle and mandibular periodontium: a double-label study

A double-label strategy was used to determine the distribution and central projections of primary afferent neurons that innervate the periodontium and muscles of mastication in cats. Central injections of either Fast Blue (FB) or a mixture of wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP) and HRP were made into one of three cytoarchitectonically distinct regions of the spinal trigeminal nucleus. These regions included the subnucleus oralis (Vo), the subnucleus interpolaris (Vi), and the medullary dorsal horn (MDH). In each case, injections were also made into the periodontium of the ipsilateral mandibular teeth or into the ipsilateral masseter muscle. FB injections preceded the peroxidase injections by at least 48 hours and total survival time ranged from 72 to 96 hours. Animals were perfused with phosphate-buffered paraformaldehyde (4%; pH 7.2). Serial frozen sections were made through the brainstem and trigeminal ganglion. Tetramethylbenzidine was used as a chromagen to demonstrate HRP and sections were viewed with brightfield and epifluorescent illumination. Cells containing peripherally injected tracer were observed in the lateral portion of the ganglion and in the mesencephalic nucleus (Vmes). Double-labeled ganglion cells were observed in most cats that received periodontal injections in combination with central injections in the dorsal part of spinal trigeminal nucleus regardless of the rostrocaudal level of the central injection. In the animals that received intramuscular injections, double-labeled ganglion cells were observed only in the animals that received central injections caudal to the Vo. Double-labeled Vmes perikarya were observed in cats that received either intramuscular or periodontal injections in combination with central injections into the MDH and Vo but not in animals that received injections into the Vi. These results demonstrate that ganglion cell periodontal afferents project to the three major rostrocaudal subdivisions of the spinal trigeminal nucleus while ganglion cell muscle afferents have more limited central projections to caudal regions of the nucleus. Masseter and periodontal Vmes afferents also project ot the spinal trigeminal nucleus--specifically, to the Vo and MDH. These findings are consistent with physiological observations regarding the role of periodontal and masseteric afferents in oral and facial reflexes and somesthetic mechanisms.