The Central Distribution of Primary Afferents from the External Eyelids, Conjunctiva, and Cornea in the Rabbit, Studied Using WGA-HRP and B-HRP as Transganglionic Tracers

We have analyzed the afferent limb of the eyeblink and nictitating membrane response of the rabbit by tracing the central distribution of primary afferents from the periorbital skin, conjunctiva, and cornea using horseradish peroxidase agglutinated to wheat germ (WGA-HRP) or conjugated to choleragenoid (B-HRP) as transganglionic tracers. Afferents in the periorbital skin and conjunctiva distribute most heavily to pars caudalis of the spinal trigeminal nucleus (Vc) and to the dorsal horn of spinal segment C1 (dhC1). These afferents terminate predominantly in laminae IIo and IIi and more weakly to the adjacent laminae I and III. There are much weaker projections to spinal segment C2, rostral Vc, and adjacent reticular formation (laminae IV and V) and to the lateral part of pars interpolaris of the spinal trigeminal nucleus (Vi). No conjunctival primary afferents were seen in the rostral divisions of the trigeminal system. Weak afferent inputs from the periorbital skin are present ventrally in pars oralis of the spinal trigeminal nucleus (Vo) and in the principal trigeminal nucleus (Vp). Corneal afferents distribute most densely in the ventral part of Vi and in islands of neuropil within the trigeminal tract at the level of Vi. They also project to caudal Vc and the adjacent dhC1 in laminae I, II, and III. There are sparse projections to the ventral and dorsal parts of Vp and to the ventral part of Vo. Reticular areas adjacent to ventral Vi also receive a few corneal afferents. WGA-HRP- and B-HRP-labeled terminals were distributed similarly in most areas, but lamina I of Vc received terminals labeled with WGA-HRP and Vp and Vo received cutaneous afferents labeled with B-HRP only. Since all subdivisions of the trigeminal system receive periocular and corneal afferent inputs, we suggest that all these subdivisions may be involved in reflex eyeblinks in the rabbit.     

Projections to the inferior olive of the cat. II. Comparisons of input from the gracile, cuneate and the spinal trigeminal nuclei

The present experiments compared the projections to the inferior olive of the cat from the gracile, cuneate and spinal trigeminal nuclei. A differential labeling strategy was used for these comparisons. It was found that all three somatic sensory nuclei project to portions of all three major divisions of the contralateral inferior olive. The spinal trigeminal n. may also project less densely to the ipsilateral medial accessory olive. Projections to the dorsal accessory nucleus (DAO) and the medially-adjacent ventral lamella of the principal nucleus are roughly somatotopically organized. Although there is considerable overlap between the projection zones, the gracile n. projects predominantly to lateral DAO, the cuneate n. projects predominantly to medial DAO, and the spinal trigeminal nucleus pars caudalis projects predominantly to the most medial portions of DAO and the ventral lamella of principal olive. Projections to the medial accessory olive, on the other hand, are not as highly organized. Instead, they overlap extensively within a small egg-shaped area in the middle of the caudal half of the nucleus. Whereas all portions of the gracile and cuneate nuclei project to the inferior olive, only the pars caudalis of the spinal trigeminal nucleus appears to do so. These results were compared with the three available olivocerebellar maps as well as with the available behavioral and electrophysiological evidence on cerebellar somatotopic organization. This comparison indicated that the inputs to the cerebellum from the three second-order somatosensory nuclei via the inferior olive appear to be generally consistent with cerebellar somatotopic organization. This consistency is apparent not only with respect to the longitudinally-organized, vermal and paravermal differences in the anterior lobe, but also with respect to the transversely-organized specific somatotopy of the intermediate zone of the anterior lobe and the paramedian lobule.     

Ascending and descending internuclear projections within the trigeminal sensory nuclear complex

The cells of origin of ascending and descending internuclear pathways in the trigeminal sensory nuclear complex were studied by the method of retrograde transport of horseradish peroxidase in the cat. The cells of origin of the ascending internuclear pathways are distributed in all laminae of the caudal part of the spinal trigeminal nucleus (Vc) except for lamina II and the caudal regions of the pars interpolaris of the spinal trigeminal nucleus (Vi). The cells arising from the Vc project to all rostral trigeminal nuclei except the caudal Vi and dorsal part of the principal trigeminal nucleus (Vpd), and neurons of the caudal Vi project to the dorsomedial (Vo.dm) and rostrodorsomedial (Vo.r) divisions of the spinal trigeminal nucleus and the ventral part of the principal trigeminal nucleus (Vpv), although the main ascending fibers from the Vc arise from laminae III-V and project to the rostral Vi and pars oralis. By contrast, the cells of origin of the descending internuclear pathways are distributed in all trigeminal nuclei, with chain-like connections between the neighboring nuclei, while the caudal regions of the Vi and laminae I-II do not receive any descending projections. The main ascending fibers from the paratrigeminal nucleus (or interstitial nucleus) at the caudal level of the Vi project to the parabrachial nucleus. These findings indicate that the internuclear pathways are differentially organized between the ascending and descending projections, and suggest that the internuclear trigeminal connections have a smaller influence on the trigeminothalamic tract cells in the Vpd, caudal Vi, and lamina I.     

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.     

Projections from the spinal and the principal sensory nuclei of the trigeminal nerve to the cerebellar cortex in the cat, as studied by retrograde transport of horseradish peroxidase.

Projections from the spinal (Vsp) and the principle sensory (Vp) nuclei of the trigeminal nerve to the cerebellar cortex were studied by means of retrograde transport of horseradish peroxidase in the cat. Neurons projecting to the simple lobule and the dorsal part of the paramedian lobule (PMD) were located mainly in the dorsal part of the nucleus interpolaris (Vi) and of the caudal one third of the nucleus oralis (Vo) and in the rostralmost part of the Vo. Neurons projecting to the medial part of the posterior folia of crus II (crus IIp) were located in the dorsal to ventral parts of the Vi and of the caudal one third of the Vo and in the rostralmost part of the Vo, while those projecting to the lateral part of crus IIp were confined to the ventral part of the Vi and of the caudal one third of the Vo. Neurons of the Vp also projected to all of these cortical areas. They were relatively confined to the ventral part of this nucleus. Thest trigeminocerebellar projections were exclusively ipsilateral to the cell origin. There were sparse projections from the Vi and Vo to lobules V to VIIIa. In addition, a small group of neurons in the subnucleus magnocellularis of the nucleus caudalis of the Vsp also projected to the above cortical areas. No projections were, however, observed to the anterior portion of the anterior lobe, crus I, the anterior folia of crus II, paraflocculus, flocculus and the ventral part of the PMD. The majority of these cerebellar projection neurons were medium-sized and triangular, fusiform or ovoid in shape. There were small neurons of similar types and large multipolar neurons as well.     

Mystacial vibrissae representation within the trigeminal sensory nuclei of the cat

Somatotopic arrangements of axon terminals of primary afferent fibers innervating follicles of the mystacial vibrissae were examined in the cat by the transganglionic horseradish peroxidase (HRP) method. Forty to 60 hours after injecting HRP into a single or a group of vibrissal follicles, transported HRP was visualized by the tetramethylbenzidine technique. HRP-labeled axon terminals were distributed in the ventral subnucleus of the principal sensory trigeminal nucleus (ventral Vp), in the oral and interpolar spinal trigeminal nuclei (Vo and Vi), and in the caudal spinal trigeminal nucleus (Vc) (layer I, deep part of layer II, layers III-V) with its spinal extension into the dorsal horn of the first cervical cord segment (rostral C1). In cross sections through the caudal parts of the ventral Vp, Vi, and layer IV of the Vc and rostral C1, a single mystacial vibrissa was represented in a one-to-one fashion by a patch of dense terminal arbors of primary afferent fibers. The more dorsally a horizontal row of the mystacial vibrissae was located, the more ventrally was it represented in the ventral Vp, the more ventrolaterally in the Vi, and the more ventrally in layer IV of the Vc and the rostral C1. In addition, the more anteriorly a vibrissa was located in a horizontal row of the mystacial vibrissae, the more medially was it represented in the ventral Vp, the more ventromedially in the Vi, and the more laterally in layer IV of the Vc and rostral C1; the most posteriorly located vibrissae in the horizontal rows of the mystacial vibrissae were represented along the lateral border of the ventral Vp and Vi, and most medially in layer IV of the Vc and rostral C1. Thus, the representation pattern in the ventral Vp was rotated clockwise at about 45 degrees angle in the Vi, and projected as a mirror image in layer IV of the Vc and rostral C1. It was also indicated that the anterior-posterior arrangement of the mystacial vibrissae was represented in a rostral-caudal organization within layer IV of the Vc and rostral C1. It was also indicated that the anterior-posterior arrangement of the mystacial vibrissae was represented in a rostral-caudal organization within layer IV of the Vc and rostral C1. Patchy patterns probably replicating the distribution of the vibrissae on the face of the cat were also revealed by the cytochrome oxidase histochemical staining in cross sections through the caudal parts of the ventral Vp, Vi, and layer IV of the Vc and rostral C1.     

Anatomical connections of the nucleus prepositus of the cat

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

Trigeminal and dorsal column nuclei projections to the anterior pretectal nucleus in the rat

The projections of the trigeminal (V) sensory nuclei (VSN) and the dorsal column nuclei (DCN) to the anterior pretectal nucleus (APT) of the rat were investigated by the use of anterograde and retrograde transport of wheat-germ agglutinin-conjugated horseradish peroxidase (WGA-HRP). Injections of WGA-HRP into the APT retrogradely labeled neurons in the contralateral VSN and DCN. The labeled neurons in the VSN were most concentrated in the rostral V subnucleus interpolaris (Vi), but were also found in caudal V subnucleus oralis (Vo). No labeled neurons were seen in V subnucleus caudalis. In the DCN, retrogradely labeled neurons were observed in rostral portions of both the cuneate (Cu) and gracile (Gr) nuclei. Injections of WGA-HRP into the rostral Vi or caudal Vo resulted in dense anterograde terminal labeling in the ventral two-thirds of the APT; the labeling was maximal in the ventromedial part of the caudal half of the APT and did not extend into its most rostral portion. Labeling resulting from injections of tracer into Cu or Gr was located primarily in the ventral half of the APT, was maximal in the mid-levels of the nucleus and extended into its rostral portions. These results indicate the existence of prominent somatosensory projections to the APT and are consistent with recent findings suggesting a role for the APT in sensorimotor integration.     

Organization of the cerebellum in the pigeon (Columba livia): II. Projections of the cerebellar nuclei.

The projections of the deep cerebellar nuclei in the pigeon have been delineated using autoradiographic and histochemical (WGA-HRP) tracing techniques. A medial (CbM) and lateral (CbL) cerebellar nucleus are recognized and CbM may be further partitioned into internal, intermediate, and intercalate divisions. As in mammals, most extracerebellar projections of CbM travel in the fasciculus uncinatus (FU); the rest travel with those of CbL in the brachium conjunctivum (BC). In the pigeon, both of these pathways are bilaterally but primarily contralaterally projecting systems. FU is a predominantly descending tract, with terminations within (1) the vestibular complex, (2) a column of contiguous medial reticular nuclei from pontine to caudal medullary levels; (3) the plexus of Horsley portion of the parvicellular reticular formation, continuing through the nucleus centralis medullae oblongatae, pars dorsalis, into intermediate layer VII of the cervical spinal cord, down to cervical segment 8-9; (4) the lateral reticular nucleus and the paragigantocellular reticular nucleus; (5) the dorsal lamella of the inferior olive. Rostrally FU terminals are found in the locus ceruleus and dorsal subcerulean nucleus. Minimal FU projections are also seen to the motor trigeminal nucleus and the subnucleus oralis of the descending trigeminal system. A small projection from the intercalate division of CbM travels in BC and projects upon the midbrain central grey, the intercollicular nucleus, the lateral tectal periventricular grey, the stratum cellulare externum and, sparsely, upon the dorsolateral thalamus. The bulk of BC originates from the lateral cerebellar nucleus and consists of a massive ascending and a small descending branch. The ascending system projects upon the red nucleus and the dorsally adjacent interstitial nucleus of Cajal and midbrain central grey, the prerubral fields continuing into the stratum cellulare externum, the nucleus intercalatus thalami, the ventrolateral thalamic nucleus, the medial spiriform nucleus, the nucleus principalis precommissuralis, the nucleus of the basal optic root, the nucleus geniculatus lateralis pars ventralis, the dorsolateral thalamus, including the dorsal intermediate posterior, and the dorsolateral intermediate and anterior nuclei. BC also contains axons from the infracerebellar nucleus, which projects upon the trochlear and the oculomotor nuclei. The descending branch of BC distributes to the papilioform nucleus, the medial pontine nucleus, the gigantocellular and paramedian reticular nuclei, and, minimally, the rostral portions of the medial column and ventral lamella of the inferior olive. Taken in conjunction with data on amphibia and reptiles the present findings suggest that the fundamental ground plan of vertebrate cerebellar organization involves a medial and lateral cerebellar nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)     

The rat's postero-orbital sinus hair: I. Brainstem projections and the effect of infraorbital nerve section at different ages

The central terminations, in the trigeminal nucleus, of afferents from the rat's postero-orbital (PO) sinus hair have been investigated with transganglionic transport of horseradish peroxidase (HRP) and succinic dehydrogenase (SDH) histochemistry. The normal pattern of terminations has been compared with that found after section of an adjacent nerve, the infraorbital (IO) nerve, at three ages: neonatal, 1 week old, and adult. The PO afferent fibres have three separate representations in the brainstem-in trigeminal sensory nucleus principalis (Vp) and rostral subnucleus oralis (Vo), in trigeminal subnucleus interpolaris (Vi), and in caudal trigeminal subnucleus caudalis (Vc) and C₁ dorsal horn. In coronal sections the areas of terminations were seen as oval patches lying ventrolaterally in Vp, Vo, and Vi and ventromedially in Vc and C₁. Following neonatal IO nerve section the terminal areas were approximately doubled in Vp, Vo, and Vi but were unchanged in Vc and C₁. IO nerve section at day 7 also caused a significant, though smaller (1.4× compared with 2.0×), increase in the terminal areas in the rostral three nuclei, without changing Vc and C₁. However, no significant change in area occurred after adult IO nerve section. SDH histochemistry at 3 to 4 weeks of age showed patches of terminals on both normal and lesioned sides consistent with those seen after HRP. Previous studies have reported increased functional representation of surrounding intact skin regions, including the PO sinus hairs, after neonatal but not adult, IO nerve section. The present results show that there are concomitant anatomical changes. Like the functional results, the extent of the anatomical changes are dependent on the maturity of the rat when lesioned.     

The organization of olivo-cerebellar projections in the opossum, Didelphis virginiana, as revealed by the retrograde transport of horseradish peroxidase

The retrograde transport of horseradish peroxidase was utilized to map olivo-cerebellar projections in the Virginia opossum. The spinal cerebellum (anterior lobe, paramedian lobule and pyramis) receives input from several separate regions in the dorsal accessory nucleus, the medial accessory nucleus and portions of the principal nucleus. Evidence is present for a topographical organization whereby specific regions of the olive project to restricted longitudinal zones. The visual-auditory region of the posterior vermis receives input from small areas within the caudal part of the medial accessory nucleus. From a distinctly separate region of the caudal medial accessory nucleus (as well as the principal nucleus), axons project to the uvula. The vestibulo-cerebellum is the recipient of axons from the cap of Kooy and from two spatially separate regions of the medial accessory nucleus. The cerebellar hemisphere (Crus I and II, lobus simplex) is the target of axons from parts of all three olivary nuclei and it is possible that the projections from the different nuclei are targeted upon separate zones. The paraflocculus was found to receive an input from the rostral part of the medial accessory nucleus and from the principal nucleus. The present results suggest that a distinct olivary region may project to several widely separate areas of the cerebellum, and that one cerebellar region may receive input from several areas of the olive. The organization of the olivocerebellar projection is highly complex, but when considered in light of known inputs to the olive, certain patterns emerge.     

Connections of the paraflocculus of the cerebellum with the superior colliculus in the rat brain

The interconnections between the paraflocculus of the cerebellum and the superior colliculus (SC) of the adult male Wistar albino rat were traced utilising the retrograde and anterograde transport of horseradish peroxidase conjugated with wheatgerm agglutinin (WGA-HRP). The study comprises three series of experiments. In the first series designed to trace the connections between the paraflocculus and the pre- and deep cerebellar nuclei, microiontophoretic injections of WGA-HRP filled either the entire paraflocculus or one of its subdivisions (ventral or dorsal paraflocculus). The results showed that, the ventral and dorsal paraflocculus receive afferents from distinct and separate regions of pontine nuclei, nucleus reticularis tegmenti pontis (Rtp) and the inferior olive. The efferents from the ventral and dorsal parafloccular subdivisions project to circumscribed regions of the lateral and posterior interpositus cerebellar nuclei. In the second series of experiments, unilateral iontophoretic WGA-HRP injections were placed at varying depths, through the rostrocaudal extent of SC and the tectal connections with the brainstem, and deep cerebellar nuclei were traced. The SC projects to well circumscribed regions of the pontine nuclei. Rtp and the inferior olive and receives afferents from the lateral and posterior interpositus cerebellar nuclei. In the third series of experiments involving combined injections of WGA-HRP into SC and the ventral paraflocculus, the collicular and ventral parafloccular connections were traced. The ventral paraflocculus receives afferents from regions of pons and Rtp which in turn receive inputs from the SC and projects to the lateral and posterior interpositus cerebellar nuclei which in turn give rise to efferents to the SC. These findings demonstrate that the ventral paraflocculus is linked to the contralateral superior colliculus via (i) a crossed afferent tecto-ponto/Rtp-parafloccular pathway, and (ii) a crossed efferent paraflocculo-nucleo-tectal pathway.     

Gracile,cuneate, and spinal trigeminal projections to inferior olive in rat and monkey

In the cat, somatosensory nuclei send substantial projections to the inferior olive, where they terminate in a somatotopic fashion. Although the organization of the cat inferior olive has been used to interpret data from other species, published data suggest this organization may not occur universally. The present study investigated whether the inferior olive in albino rats and cynomolgus monkeys receives the same brainstem somatosensory inputs, whether these inputs are organized somatotopically and, if so, how the organization compares with that in the cat. Projections from the gracile, cuneate and spinal trigeminal nuclei were labeled with wheat germ agglutinin conjugated to horseradish peroxidase or with biotinylated dextran. The results were compared with data from cats (Berkley and Hand [1978] J. Comp. Neurol. 180:253-264). In the rat and monkey, the gracile, cuneate and spinal trigeminal nuclei all project to the contralateral inferior olive, where each nucleus has a distinct preferred terminal field. As in the cat, projections to the medial accessory olive and caudal dorsal accessory olive did not terminate in a precisely organized fashion. Projections to the rostral dorsal accessory olive, however, formed a clear somatotopic map. These somatotopic maps differed from those in the cat in that input from the trigeminal nucleus was confined rostrally, so that the caudal end only received input from the gracile and cuneate nuclei. These data indicate that similar organizational principles characterize the somatosensory projections to the inferior olives of the three species. Nevertheless, distinct species differences occur with regard to the details of this organization. © 1996 Wiley-Liss, Inc.     

Secondary trigeminocerebellar projections in sheep studied with the horseradish peroxidase tracing method

Secondary trigeminocerebellar connections have been studied with HRP histochemistry in 25 sheep. The results indicate that almost all of the cerebellar cortex except flocculus, ventral paraflocculus and lobules I-IV receives bilateral (mostly ipsilateral) fibers from the trigeminal nuclei. A topographical organization of trigeminocerebellar fibers is present. The mesencephalic tract nucleus projects to the anterior lobe, the simple lobule (HVI), lobules VI, VIII, and the dorsal paraflocculus. The ventral group of the princeps and spinal tract (mainly IDV) nuclei projects to all lobules studied in vermis and hemispheres. More dorsal parts of these nuclei have a more restricted projection field including the vermal lobules VI, VII, and IX and the hemisphere. Cells within and ventral to the motor nucleus of the trigeminal nerve were found labeled after injections into the anterior lobe, the simple lobule, and lobule IX. Labeled cells in the region of the nucleus ovalis and close to the solitary tract project to the simple and paramedian lobule and lobule IX.     

Brain stem nuclei giving fibers to lobules VI and VII of the cerebellar vermis.

The HRP method has been used to identify all the brain stem nuclei, which may project to lobule VI and/or VII of the posterior cerebellar vermis. Three tentative degrees of labeling of the different structures have been assigned: 'massive', 'clear' and 'discrete'. (1) Massive projections have been found to reach lobule VI and VII from the inferior olive and lobule VII only from the nucleus reticularis tegmenti pontis. (2) Clear projections have been found to reach lobule VI only from the pontine nuclei, the nucleus reticularis tegmenti pontis, the nucleus reticularis lateralis and the reticularis paramedianus; lobule VII only from the raphe nuclei, and both VI and VII from the perihypoglossal and vestibular nuclei. (3) Discrete projections have been found to reach lobule VI and VII from the deep cerebellar nuclei; lobule VI only from the nucleus tracti solitarii and nucleus cuneatus externus; lobule VII only from the nucleus lemnisci lateralis pars ventralis, the nuclei parabrachiales and the nucleus subcoeruleus.     

Single axonal characterization of trigeminocerebellar projection patterns in the mouse

The cerebellar projection from the trigeminal nuclear complex is one of the major populations of the cerebellar inputs. Although this projection is essential in cerebellar functional processing and organization, its morphological organization has not been systematically clarified. The present study addressed this issue by lobule-specific retrograde neuronal labeling and single axonal reconstruction with anterograde labeling. The cerebellar projection arose mainly from the interpolaris subdivision of the spinal trigeminal nucleus (Sp5I) and the principal trigeminal sensory nucleus (Pr5). Although crus II, paramedian lobule, lobule IX, and simple lobule were the major targets, paraflocculus, and other lobules received some projections. Reconstructed single trigeminocerebellar axons showed 77.8 mossy fiber terminals on average often in multiple lobules but no nuclear collaterals. More terminals were located in zebrin-negative or lightly-positive compartments than in zebrin-positive compartments. While Pr5 axons predominantly projected to ipsilateral crus II, Sp5I axons projected either predominantly to crus II and paramedian lobule often bilaterally, or predominantly to lobule IX always ipsilaterally. Lobule IX-predominant-type Sp5I neurons specifically expressed Gpr26. Gpr26-tagged neuronal labeling produced a peculiar mossy fiber distribution, which was dense in the dorsolateral lobule IX and extending transversely to the dorsal median apex in lobule IX. The projection to the cerebellar nuclei was observed in collaterals of ascending Sp5I axons that project to the diencephalon. In sum, multiple populations of trigeminocerebellar projections showed divergent projections to cerebellar lobules. The projection was generally complementary with the pontine projection and partly matched with the reported orofacial receptive field arrangement.     

The pontocerebellar projection onto the paramedian lobule in the cat: an experimental study with the use of horseradish peroxidase as a tracer.

Horseradish peroxidase (HRP) was injected into cerebellar cortex of the paramedian lobule in 12 cats, and the ensuing distribution of labeled cells in the pontine nuclei was mapped in some detail. The cells in the pontine gray which give origin to fibers to the paramedian lobule lie together, in part in groups, and in part in columns. The columns are situated both medial and ventrolateral to the peduncle, as well as in the dorsolateral pontine nucleus. The projection is bilateral with a clearcut contralateral preponderance, except in the lateralmost region in the dorsolateral nucleus, which projects mainly ipsilaterally. The column medial to the peduncle projects in a topographical pattern to the paramedian lobule. The dorsal part of this column projects to the rostral folia of the paramedian lobule, while successively more ventral parts in the column project to more caudal paramedian lobules. Within the other columns only a faint sign of a topographical organization is found. The location of the pontine columns projecting onto the paramedian lobule largely corresponds to the pontine terminal areas of fibers from the sensory cerebral cortex (SmI and SmII). The corresponding topography in these parts of the corticopontine and pontocerebellar pathways is suitable for a somatotopical impulse transmission from the sensory cortex to the paramedian lobule, in agreement with the results of physiological investigations. Furthermore, a correlation of the pontine areas projecting onto the paramedian lobule with the terminal areas of pontine afferents shows that the pons may be a relay station in mediating influences from other parts of the cortex (MsI, visual and acoustic), the cerebellar nuclei and the colliculi to the paramedian lobule.     

The trigemino-olivary projection in the cat: contributions of individual subnuclei

Anterograde autoradiographic methods were used to determine the projection of the principal sensory trigeminal nucleus and of each of the three spinal trigeminal subnuclei to the inferior olivary complex in the cat. Our data reveal that the principal sensory trigeminal nucleus does not contribute to the trigemino-olivary pathway. Each spinal trigeminal subnucleus has a unique contribution to this pathway: pars oralis projects sparsely to the border between the dorsal accessory and principal olives (DAO-PO), pars interpolaris projects mostly to the rostral medial DAO, and pars caudalis projects mostly to the rostral medial part of the ventral leaf of PO and slightly to the caudal medial accessory olive. In the light of recent physiological and anatomical findings, our data indicate that information from each spinal trigeminal subnucleus reaches a different segment of the contralateral inferior olivary complex, which in turn distributes differentially to the cerebellar cortex.     

Morphology of central terminations of intra-axonally stained, low-threshold mechanoreceptive primary afferent fibers from oral mucosa and periodontium in the rat

Horseradish peroxidase (HRP) was injected intra-axonally into functionally identified primary afferent fibers within the rat spinal trigeminal tract in order to study the morphology of their central terminations. They were physiologically determined to be large, myelinated afferent fibers from periodontium or oral mucosa by means of electrical and mechanical stimulation of their receptive fields. Twenty-eight axons that innervated the periodontium of incisors and 21 axons that innervated the oral mucosa were stained for distances of 2-5 mm from the injection sites at the levels of the main sensory nucleus (Vms), spinal trigeminal nucleus and rostral cervical spinal cord. The collaterals of these primary afferent fibers formed terminal arbors in the medial or dorsomedial part of the Vms, and the oral and interpolar spinal trigeminal nuclei (Vo and Vi). In the caudal spinal trigeminal nucleus (Vc), the collaterals of one half of the periodontium afferent fibers terminated mainly in lamina V at the rostral and middle levels of Vc. On the other hand, the collaterals of the other half of the periodontium afferent fibers terminated mainly in lamina IV at the rostral level of Vc, and rostrally these terminal areas shifted to the most medial part of Vi. The collaterals of mucosa afferent fibers terminated in lamina V at the rostral level of Vc, and these terminal areas shifted gradually to laminae III and IV as the parent axons traveled more caudally. These shifts were staggered rostrocaudally according to the rostrocaudal locations of the receptive fields. The density of collaterals of periodontium afferent fibers in Vi was significantly larger than that of mucosa afferent fibers. The average size of the varicosities of periodontium afferent fibers was significantly larger than those of mucosa afferent fibers in Vo, Vi and Vc. The average number of varicosities belonging to single collaterals of slowly-adapting periodontium afferent fibers in Vi were significantly larger than those in Vo. In Vi, the average number of varicosities of single collaterals of slowly-adapting periodontium afferent fibers were significantly larger than those of rapidly-adapting periodontium afferent fibers.     

Visual pontocerebellar projections in the macaque

The cerebellum plays an important role in the visual guidance of movement. In order to understand the anatomical basis of visuomotor control, we studied the projection of pontine visual cells onto the cerebellar cortex of monkeys. Wheat germ agglutinin horseradish peroxidase was injected into the dorsolateral pons of two monkeys. Retrogradely labelled cells were mapped in the cerebral cortex and superior colliculus, and orthogradely labelled fibers in the cerebellar cortex. The largest number of retrogradely labelled cells in the cerebral cortex was in a group of medial extrastriate visual areas. The major cerebellar target of these dorsolateral pontine cells is the dorsal paraflocculus. There is a weaker projection to the uvula, paramedian lobe, and Crus II, and a sparse but definite projection to the ventral paraflocculus. There are virtually no projections to the flocculus. There are sparse ipsilateral pontocerebellar projections to these same regions of cerebellar cortex. In nine monkeys, we made small injections of the tracer into the cerebellar cortex and studied the location of retrogradely filled cells in the pontine nuclei and inferior olive. Injections into the dorsal paraflocculus or rostral folia of the uvula retrogradely labelled large numbers of cells in the dorsolateral region of the contralateral pontine nuclei. Labelled cells were found ipsilaterally, but in reduced numbers. Injections outside of these areas in ventral paraflocculus or paramedian lobule labelled far fewer cells in this region of the pons. We conclude that the principal source of cerebral cortical visual information arises from a medial group of extrastriate visual areas and is relayed through cells in the dorsolateral pontine nuclei. The principal target of pontine visual cells is the dorsal paraflocculus. © 1994 Wiley-Liss, Inc.