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.     

Cell loss in lumbar dorsal root ganglia and transganglionic degeneration after sciatic nerve resection in the rat

The effects of sciatic nerve resection on lumbar dorsal root ganglion cells and their central branches have been studied in the adult rat. A quantitative analysis of the lumbar dorsal root ganglia indicated a 15–30% cell loss on the operated side. Argyrophilia indicating transganglionic degeneration was observed in Fink-Heimer stained sections from the lumbar spinal cord and the brainstem. The areas of degeneration argyrophilia were mainly located in the medial part of the ipsilateral L2–L6 dorsal horn laminae I–IV, the tract of Lissauer, the dorsal funiculus and the gracile nucleus. A few degenerating fibers could also be observed in the ipsilateral dorsal horn laminae V and VI, and in the ipsilateral ventral horn as well as in the contralateral dorsal and the gracile nucleus. The results confirm and extend previous findings at other levels and in other species. This suggests that cell loss and transganglionic degeneration may be general phenomena affecting a substantial proportion of primary sensory neurons following peripheral nerve injury.     

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

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

Central distribution of primary afferent fibers in the Arnold's nerve (the auricular branch of the vagus nerve): A transganglionic HRP study in the cat

Central projections of the Arnold's nerve (the auricular branch of the vagus nerve; ABV) of the cat were examined by the transganglionic HRP method. After applying HRP to the central cut end of the ABV, HRP-labeled neuronal somata were seen in the superior ganglion of the vagus nerve. Main terminal labeling was seen ipsilaterally in the solitary nucleus, in the lateral portions of the ventral division of the principal sensory trigeminal nucleus, in the marginal regions of the interpolar subnucleus of the spinal trigeminal nucleus, in the marginal and magnocellular zones of the caudal subnucleus of the spinal trigeminal nucleus, in the ventrolateral portions of the cuneate nucleus, and in the dorsal horn of the C1–C3 cord segments. In the solitary nucleus, labeled terminals were seen in the interstitial, dorsal, dorsolateral and commissural subnuclei; some of these terminals may be connected monosynaptically with solitary nucleus neurons which send their axons to the somatomotor and/or visceromotor centers in the brainstem and spinal cord.     

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

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

Segmental distribution and central projectionsof renal afferent fibers in the cat studied by transganglionic transport of horseradish peroxidase

The segmental and central distributions of renal nerve afferents in adult cats and kittens were studied by using retrograde and transganglionic transport of horseradish peroxidase (HRP). Transport of HRP from the central cut ends of the left renal nerves labeled afferent axons in the ipsilateral minor splanchnic nerves and sensory perikarya in the dorsal root ganglia from T12 to L4. The majority of labeled cells (85%) were located between L1 and L3. A few neurons in the contralateral dorsal root ganglia were also labeled. Labeled cells were not confined to any particular region within a dorsal root ganglion. Some examples of bifurcation of the peripheral and central processes within the ganglion were noted. A small number of preganglionic neurons, concentrated in the intermediolateral nucleus, were also identified in some experiments. In addition, many sympathetic postganglionic neurons were labeled in the renal nerve ganglia, the superior mesenteric ganglion, and the ipsilateral paravertebral ganglia from T12 to L3 Transganglionic transport of HRP labeled renal afferent projections to the spinal cord of kittens from T1 1 to L6, with the greatest concentrations between Ll and L3. These afferents extended rostrocaudally in Lissauer's tract and sent collaterals into lamina I. In the transverse plane, a major lateral projection and a minor medial projection were observed along the outer and inner margins of the dorsal horn, respectively. From the lateral projection many fibers extended medially in laminae V and VI forming dorsal and ventral bundles around Clarke's nucleus. The dorsal bundle was joined by collaterals from the medial afferent projection and crossed to the contralateral side. The ventral bundle extended into lamina VII along the lateroventral border of Clarke's nucleus. Some afferents in the lateral projection could be followed ventrally into the dorsolateral portion of lamina VII in the vicinity of the intermediolateral nucleus. In the contralateral spinal cord, labeled afferent fibers were mainly seen in laminae V and VI These results provide the first anatomical evidence for sites of central termination of renal afferent axons. Renal inputs to regions (laminae I, V, and VI) containing spinoreticular and spinothajamic tract neurons may be important in the mediation of supraspinal cardiovascular reflexes as well as in the transmission of activity from nociceptors in the kidney. In addition, the identification of a bilateral renal afferent projection in close proximity to the thoracolumbar autonomic nuclei is consistent with the demonstration in physiological experiments of a spinal pathway for the renorenal sympathetic reflexes.     

Brain stem terminations of the trigeminal and upper spinal ganglia innervation of the cerebrovascular system: WGA-HRP transganglionic study

The central projections of the nerve fibers innervating the middle cerebral and basilar arteries were investigated by transganglionic tracing of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) in the rat. WGA-HRP was applied to the exposed basilar and/or middle cerebral arteries. Sections of the brain, trigeminal and upper spinal ganglia were reacted with tetramethylbenzidine for detection of the tracer. The results demonstrate that trigeminal neurons that innervate the middle cerebral artery project to the trigeminal main sensory nucleus, pars oralis, and the dorsocaudal two-fifths of pars interpolaris of the trigeminal brain stem nuclear complex. Terminals were also visible in the ipsilateral nucleus motorius dorsalis nervi vagi (dmnX) and in the lateral nucleus tractus solitarius (nTs) bilaterally at the level of the obex. The ventral periaqueductal gray, including the dorsal raphe and C2 dorsal horn, were also innervated by nerve fibers from the middle cerebral artery. Ipsilateral trigeminal rhizotomy prior to WGA-HRP application over the middle cerebral artery impeded the visualization of nerve terminations throughout the brain stem. Pretreatment with capsaicin reduced the density of labeled neurons and terminals within the trigeminal ganglion and the brain stem, respectively, following WGA-HRP application over the middle cerebral artery. Basilar artery fibers terminate in the C2 dorsal horn, the cuneate nuclei, dmnX, and nTs bilaterally. A few projections were also labeled in the ventral periaqueductal gray. Unilateral upper two spinal dorsal rhizotomy prior to WGA-HRP application over the exposed basilar artery resulted in terminal labeling within the C2 dorsal horn, the cuneate nucleus, dmnX, and nTs contralateral to the rhizotomy, whereas the ipsilateral side was devoid of any labeling. Bilateral superior cervical ganglionectomy prior to WGA-HRP administration to the middle cerebral and basilar arteries did not alter the visualization of nerve terminations throughout the brain stem.     

Sexual dimorphism in sympathetic preganglionic neurons of the rat hypogastric nerve

Horseradish peroxidase (HRP) applied to one hypogastric nerve labelled sensory neurons in T11-L3 dorsal root ganglia (DRG) bilaterally and preganglionic neurons (PGN) in the spinal cord segments T13-L3. An average of 130 small DRG neurons were labelled per animal (male or female). These were concentrated in the L1 + L2 DRGs (92%). About 75% were located ipsilateral to the site of HRP application. Central projections from DRG neurons were noted throughout Lissauer's tract and in the marginal zones (medial and lateral) near the borders of Lissauer's tract. A short projection was also seen extending to the dorsolateral funiculus. More than 90% of the preganglionic neurons were located in segments L1 + L2. Most of these were found in the dorsal commissural nucleus (75%) and most of the remainder were located bilaterally in the intermediolateral columns. Somewhat more intermediolateral neurons were labelled on the ipsilateral side than on the contralateral side. There were a few intercalating neurons and a very few funicular cells. An average of 415 PGNs were labelled in the male animals and 110 in the females, demonstrating a strong sexual dimorphism. No dimorphism was found in the sensory components.     

Central projections of cervical primary afferent fibers in the guinea pig: an HRP and WGA/HRP tracer study

The central course and the projections of the first and the second cervical dorsal root ganglia and of suboccipital muscle primary afferent fibers in the guinea pig were studied by means of anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA/HRP) or aqueous solution of horseradish peroxidase (HRP). Injections of WGA/HRP into the second cervical dorsal root ganglion produced labeling in the dorsal and ventral horns. Within the spinal cord, the largest amount of HRP reaction product was found within the lateral third of the substantia gelatinosa and within the central cervical nucleus. The main area of termination in the medulla was the external cuneate nucleus. However, HRP reaction product was also found within the medial and inferior vestibular nuclei, cell group x, the perihypoglossal nuclei, the nucleus of the solitary tract, and the nucleus of the spinal trigeminal tract. Descending fibers could be detected as caudal as spinal segment T5. Injections of WGA/HRP into the first cervical dorsal root ganglion produced heavy terminal label within the central cervical nucleus but not within the substantia gelatinosa. Again, the external cuneate nucleus was the main area of termination within the medulla. Label could not be observed within the vestibular nuclear complex or within the spinal trigeminal nucleus. Injections of aqueous HRP into the suboccipital muscles produced heavy transganglionic label within the central cervical nucleus, whereas the substantia gelatinosa totally lacked terminal label. Ascending proprioceptive fibers reached the external cuneate nucleus and group x. Scanty projections could be detected within the vestibular nuclei as well as within the perihypoglossal nuclei except for the nucleus prepositus hypoglossi. Label was absent in the spinal trigeminal nucleus.     

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

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

Spinal cord projections from hindlimb muscle nerves in the rat studied by transganglionic transport of horseradish peroxidase, wheat germ agglutinin conjugated horseradish peroxidase, or horseradish peroxidase with dimethylsulfoxide

The spinal cord projections of four different groups of hindlimb muscle nerve branches--the medial and lateral gastrocnemius nerves, muscle branches of the deep peroneal nerve, muscle branches of the femoral nerve, and a nerve to the hamstring muscles--were studied with transganglionic transport of horseradish peroxidase (HRP) in the rat. The influence of varying the postoperative survival (3, 6, and 10 days) and of using wheat germ agglutinin-HRP conjugate (WGA-HRP), or HRP with dimethylsulfoxide (DMSO) instead of free HRP was studied for the gastrocnemius nerves. After 3 days' survival following application of HRP to the gastrocnemius nerves, fine granular labeling was found mainly in lamina V in L4-5, and coarse granular labeling was found in Clarke's column as far caudally as L2, and in laminae VI and VII predominantly in Th12-L2. After 6 or 10 days' survival, the fine labeling in lamina V was sparse or absent, whereas the coarse labeling appeared to remain or to be only slightly reduced in Clarke's column and in laminae VI and VII. No labeling suggestive of terminals was observed in laminae I-III from the gastrocnemius nerves. Except for sparse labeling in lamina I in some of the cases and some minor differences rostrocaudally, the spinal distribution of labeling was similar to that from the other nerves investigated. The distribution of labeling obtained after application of WGA-HRP or HRP with DMSO to the gastrocnemius nerves was very similar to that obtained with free HRP after 3 days' survival. The results indicate that the spinal cord projections of hindlimb muscle nerves in the rat distribute mainly in the deep part of the dorsal horn and in the intermediate zone. Furthermore, the lack of labeling suggestive of terminals in laminae I-III from the gastrocnemius nerves suggests, in conflict with earlier findings in the cat, that primary afferent fibers from muscles do not necessarily terminate in these laminae in the rat. The results suggest, furthermore, that fine granular labeling found in lamina V represents fine-calibered afferent fibers. Finally, the similar spinal projection patterns of the different muscle nerves investigated suggest either a less developed or an essentially different somatotopic organization for muscle afferents compared to cutaneous afferents, as revealed in earlier studies.     

Projections from the red nucleus and surrounding areas to the brainstem and spinal cord in the cat. An HRP and autoradiographical tracing study

HRP injections at the C2, T1 and S1 spinal levels and in the medullary lateral tegmental field revealed that the contralaterally projecting rubro-bulbospinal neurons are located not only in the caudal but also to a certain extent in the rostral red nucleus (RN). These RN projections are somatotopically organized. Neurons projecting to the sacral cord are located in the ventrolateral RN, those projecting to the upper part of the spinal cord lie in the dorsomedial RN and those projecting to the medullary lateral tegmentum were found in the dorsal portions of the RN. These last neurons are smaller than many of the other RN neurons. The HRP results also revealed that the RN does not project to the caudal raphe nuclei. The autoradiographical results confirmed the HRP findings. They further indicated that the contralateral RN projections to the caudal brainstem precerebellar nuclei (nucleus corporis pontobulbaris, lateral reticular nucleus, lateral cuneate nucleus) and the dorsal column nuclei are also somatotopically organized. This was also true for the RN projections to the dorsomedial and intermediate facial subnuclei and the caudal pontine and medullary lateral tegmental field. These areas receive afferents from mainly the dorsal portions of the RN. Regarding the RN projections to the spinal cord, the autoradiographical tracing results revealed somatotopically organized contralateral RN projections to laminae V, VI and VII. Moreover, a small but distinct RN projection to a dorsolaterally located group of motoneurons at the C8-T1 level was demonstrated. Ipsilaterally a minor projection to the cervical and upper thoracic lateral intermediate zone was observed. Finally, strong ipsilateral projections from the rostral mesencephalon to the inferior olive were seen. These projections were derived from various rostral mesencephalic areas, including the nucleus of Darkschewitsch, the nucleus accessorius medialis of Bechterew, the interstitial nucleus of Cajal and the area of the rostral interstitial nucleus of the medial longitudinal fasciculus. In the cat it was difficult to define which of the mesencephalic areas projecting to the inferior olive represented the parvocellular RN. A new subdivision of the RN is proposed based on its projections and not on the size of its cells. In this concept the first group is formed by the RN neurons projecting contralaterally to the caudal brainstem and spinal cord. The second group consists of RN neurons projecting to the inferior olive.(ABSTRACT TRUNCATED AT 400 WORDS)     

The termination of forelimb nerves in the feline cuneate nucleus demonstrated by the transganglionic transport method

The projection of forelimb nerves to the cuneate nucleus was studied in the cat by the transganglionic transport method. The cut ends of the median, ulnar, musculocutaneous, medial cutaneous, lateral brachial and antebrachial cutaneous nerves and of the superficial and deep branches of the radial nerve were exposed to horseradish peroxidase (HRP) or to HRP conjugated to wheat germ agglutinin.Nerves innervating the skin terminated in a somatotopical pattern on the cell clusters in the middle region of the cuneate nucleus. Afferents from the paw occupied the largest area and were situated dorsally, with the ulnar part represented medially and the radial part laterally. The palmar side of the digits seemed to be represented superficial to the dorsal side. The projection from the arm was split into a ventromedial and a ventrolateral area. Superimposed on this somatotopy, a reversed termination pattern was also present. Thus medially projecting nerves also had a small separate projection to the lateral part of the nucleus and vice versa. The rostral region of the nucleus was organized in a similar way except for the rostral pole where the somatotopy was lost. The caudal region differed from the middle one in that it appeared to lack representation of the upper and lower arm.The deep branch of the radial nerve terminated in the middle-ventral ‘reticular’ region of the cuneate nucleus, with a sparse projection also to the ventral parts of the rostral and caudal regions, including the base of the dorsal horn. Also the musculocutaneous, median and ulnar nerves, but not the pure cutaneous nerves, had projections to these areas, indicating a modality segregation in the cuneate nucleus. The rostral pole of the nucleus, however, appeared to constitute an area of overlap between projections from deep and superficial receptors.     

Tracing of afferent and efferent pathways in the left inferior cardiac nerve of the cat using retrograde and transganglionic transport of horseradish peroxidase

Retrograde and transganglionic transport of horseradish peroxidase (HRP) was used to trace afferent and efferent pathways in the left inferior cardiac nerve of the cat. Cardiac efferent and afferent neurons were located, respectively, in the stellate ganglion (average cell count per experiment: 2679) and in the ipsilateral dorsal root ganglia (DRG) from C8 to T9 (average cell count per experiment: 213). Labeled cardiac afferent projections to the spinal cord were most dense in segments T2–T6 where they were located in Lissauer's tract and in lamina 1 on the lateral border of the dorsal horn. Labeled affrent axons extended ventrally through lamina 1 into lamina 5 and the dorsolateral region of lamina 7 in proximity to the intermediolateral nucleus. A weak projection was noted on the medial side of the dorsal horn. These sites of termination are similar to projections by other sympathetic afferent pathways (i.e. renal, hypogastric and splanchnic nerves) to the lower thoracic and lumbar spinal cord, indicating that visceral afferents may have a uniform pattern of termination at various segmental levels. This pattern of termination in regions of the gray matter containing spinothalamic tract neurons and neurons involved in autonomic mechanisms is consistent with the known functions of sympathetic afferent pathways in nociception and in the initiation of autonomic reflexes.     

Representation of the forepaw in the feline cuneate nucleus: a transganglionic transport study

The projection of forelimb nerves innervating the paw to the cuneate nucleus was studied in the cat by the transganglionic transport method. Exposure of a single digital nerve to the tracer (a conjugate of horseradish peroxidase to wheat-germ agglutinin) resulted in a longitudinal sequence of labeled patches throughout the extent of the nucleus. In the middle region the labeled patches coincided with the location of the cell clusters that are characteristic of this part of the nucleus. A very precise somatotopic termination pattern was found in the middle region of the nucleus. Afferent fibers from the palm were represented superficially close to the dorsal rim. The digits were represented in a mediolateral sequence, with the first digit in the dorsolateral part of the nucleus and the fifth digit in the dorsomedial part. The ventral surfaces of the digits were represented superficial to the dorsal surfaces. The dorsum of the paw was represented close to the center of the nucleus. A similar somatotopic organization, but much less detailed, was found in the rostral and caudal regions of the cuneate nucleus. These dissimilarities in somatotopic detail between the different cytoarchitectonic regions of the cuneate nucleus probably reflect differences in function between these regions.     

Brainstem terminations of extraocular muscle primary afferent neurons in the monkey

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

The central projection of muscle afferent fibres to the lower medulla and upper spinal cord: an anatomical study in the cat with the transganglionic transport method

The projection of muscle afferent fibres to the medulla oblongata and upper spinal cord was studied in the cat by using transganglionic transport of wheat germ agglutinin-horseradish peroxidase conjugate. The results demonstrate a precise, musculotopic termination pattern in the external cuneate nucleus; thus, fibres from the intrinsic muscles of the paw terminate medially; those from forearm, arm, and shoulder muscles terminate progressively more laterally; and those from neck and thoracic muscles terminate in the ventrolateral and dorsolateral parts, respectively. Muscle afferent fibres to the main cuneate nucleus terminate in the ventral "reticular" region of the nucleus, with a sparse projection also to the ventral part of the rostral and caudal regions, including the base of the dorsal horn. Fibres from the neck muscles terminate slightly more laterally in the ventral region than do those from the limb muscles, but otherwise, and thus contrary to the case in the external cuneate nucleus, no topographic organization was detected. In the spinal cord, projection was found to laminae I and V, and from the musculature of the back of the neck to the central cervical nucleus.     

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.     

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

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