Transganglionic Transport Of Horseradish Peroxidase Research Articles

Trigeminal primary afferent projections to the peribrachial area in the guinea pig

Transganglionic transport of horseradish peroxidase (HRP) was used to investigate trigeminal primary afferent endings in the peribrachial area of the guinea pig. After HRP application to several nerve branches of the three divisions of the trigeminal nerve, a projection to the caudal region of the ipsilateral peribrachial region was found for the infratrochlear, ethmoidal, supraorbital, zygomatic, zygomaticofacial, zygomaticotemporal and infraorbital-superior alveolar nerves. Random and scarce endings were also observed for the mental, inferior alveolar, buccal and mylohyoid nerves. No trigeminal primary projection to the peribrachial area was detected when HRP was applied to the sphenopalatine, lingual and auriculotemporal nerves. With the exception of the ethmoidal endings described in the muskrat, a primary afferent projection to the peribrachial region from the other trigeminal nerves has not hitherto been reported in mammals.

Brainstem projections of different branches of the vestibular nerve: an experimental study by transganglionic transport of horseradish peroxidase in the cat. III. The saccular nerve.

The brainstem projection of the saccular nerve was investigated by transganglionic tracing with horseradish peroxidase in the cat. The labeled fibers were located most caudally and laterally (dorsolaterally) in the vestibular nerve root. This was unique when compared with the locations of the fibers from the nerves of the other vestibular-end organs that were studied by the present authors by the same approach previously. In addition, such a comparison revealed a specific location, from lateral to medial, for the fibers from each of the five divisions of the vestibular nerve. In the present study, labeled fibers from the sacculus, of fine caliber, were found close to the restiform body, both medially and laterally, some even penetrating through this structure. Labeled terminals were present in cell group "y". This was unique, compared with the nerves from the other end organs. Such terminals were also found in the four main vestibular nuclei, except for the medial vestibular nucleus, where no labeled terminals could be detected. No labeled terminals were found in the interstitial nucleus of the vestibular nerve. Together with the findings from our previous studies, this suggests that, in contrast to the ampullar nerves, the nerves from the maculae do not project to this structure. This study confirms, but also extends, findings reported from a previous investigation in the cat, using an experimental degeneration technique.

The distribution of primary afferent terminals from the eyelids of macaque monkeys.

Trigeminal sensory afferents from the eyelids convey two types of information that are important for the blink reflex. Movement of the lashes activates low-threshold mechanoreceptors which evoke protective blinks. Information about eyelid position is also transmitted centrally and is used to adapt the metrics of the blink reflex to changing conditions over time. This study employed transganglionic transport of horseradish peroxidase conjugated to choleratoxin-B subunit or wheat-germ agglutinin to investigate trigeminal afferents supplying the eyelids of macaque monkeys. Ganglion cells labeled from upper- and lower-lid injections were located in the ophthalmic and maxillary portions of the trigeminal ganglion, respectively. In both cases, labeled terminals were observed ipsilateral to the injected eyelid in the principal and spinal trigeminal nuclei. However, only a few labeled terminals were present in the principal nucleus, and very sparse terminal labeling was confined to a few locations along the ventral border of the pars oralis and interpolaris of the spinal trigeminal nucleus. The main concentration of label was found in the pars caudalis at and immediately below the spinomedullary junction. The terminal field from the upper eyelid was located ventrally in the pars caudalis, and that from the lower eyelid was located more dorsally. In both cases, the labeled terminal field was densest within lamina II of the spinal trigeminal nucleus. The heavy concentration of eyelid central terminals at the spinomedullary junction is surprising in light of physiological studies indicating representation of all parts of the face throughout the trigeminal nucleus. The distribution of eyelid afferent terminals in the macaque is caudal to the main concentration of corneal afferent terminals at the pars interpolaris/caudalis border. This may be a basis for differences seen in blinks produced by corneal as opposed to supraorbital stimulation. The presence of a single major site of eyelid primary afferent terminals suggests that sensory input for both eyelid proprioception and blink-reflex activation passes through this segment of the spinal trigeminal nucleus. These results provide a basis for investigation of the central connections of pars caudalis neurons in order to better establish the pathways producing trigeminally evoked blinks and blink adaptation.

Substance P immunoreactive nerve terminals in the dorsolateral nucleus of the tractus solitarius: roles in the baroreceptor reflex

Physiological and light microscopic evidence suggest that substance P (SP) may be a neurotransmitter contained in first-order sensory baroreceptor afferents; however, ultrastructural support for this hypothesis is lacking. We have traced the central projections of the carotid sinus nerve (CSN) in the cat by utilizing the transganglionic transport of horseradish peroxidase (HRP). The dorsolateral subnucleus of the nucleus tractus solitarius (dlNTS) was processed for the histochemical visualization of transganglionically labeled CSN afferents and for the immunocytochemical visualization of SP by dual labeling light and electron microscopic methods. Either HRP or SP was readily identified in single-labeled unmyelinated axons, myelinated axons, and nerve terminals in the dlNTS. SP immunoreactivity was also identified in unmyelinated axons, myelinated axons, and nerve terminals in the dlNTS, which were simultaneously identified as CSN primary afferents. However, only 15% of CSN terminals in the dlNTS were immunoreactive for SP. Therefore, while the ultrastructural data support the hypothesis that SP immunoreactive first-order neurons are involved in the origination of the baroreceptor reflex, they suggest that only a modest part of the total sensory input conveyed from the carotid sinus baroreceptors to the dlNTS is mediated by SP immunoreactive CSN terminals. Five types of axo–axonic synapses were observed in the dlNTS. SP immunoreactive CSN afferents were very rarely involved in these synapses. Furthermore, SP terminals were never observed to form the presynaptic element in an axo–axonic synapse with a CSN afferent. Therefore, SP does not appear to be involved in the modulation of the baroreceptor reflex in the dlNTS.

Somatotopic organization of the facial lobe of the sea catfish Arius felis studied by transganglionic transport of horseradish peroxidase.

To reveal the somatotopical organization of the facial lobe (FL), a primary medullary gustatory nucleus in the sea catfish Arius felis, the central projections of the peripheral rami of the facial nerve innervating taste buds located across the entire body surface and rostral oral regions were traced by means of horseradish peroxidase neurohistochemistry. The maxillary barbel, lateral mandibular barbel, medial mandibular barbel, and trunk-tail branches project to four different longitudinal columns (i.e., lobules) extending rostrocaudally in the FL. The trunk-tail lobule, which is located dorsolateral to the barbel lobules, lies in the anterior two-thirds of the FL. The tail is represented in a more rostral portion of the trunk-tail lobule than the trunk, indicating that the rostrocaudal trunk axis is represented in the trunk-tail lobule in a posteroanterior axis. The pectoral fin branch ends in an intermediate region of the FL, whereas the hyomandibular, ophthalmic, lower lip, upper lip, and palatine branches terminate in discrete regions of the caudal one-third of the FL. These results reveal a sharply defined somatotopical organization of the FL of Arius and support the hypothesis that the number and lengths of the barbel lobules within the FL of catfishes are directly related to the number and relative lengths of the barbels. An additional subcolumn, the intermediate nucleus of the FL (NIF), which develops in the medioventral region of the caudal two-thirds of the FL, receives projections in a diffuse somatotopical fashion from the barbels, lower lip, and palatine branches. Trigeminal fibers of the barbel and lower lip branches project in a somatotopic fashion to the FL. The present findings suggest that the FL of Arius is highly organized somatotopically to detect, by tropotaxis, precise spatial information concerning taste and tactile stimuli in the environment.

Projections of ankle joint afferents to the spinal cord and brainstem of the chicken (Gallus G. domesticus)

The projections of the ankle joint capsule afferents were studied by transganglionic transport of horseradish peroxidase injected directly into the ankle joint. The number and size of the labelled dorsal root ganglion cells were measured from synsacral nerves 2-9. In the dorsal root ganglia, all sizes of sensory neurones were labelled, and the largest number of labelled cells was in ganglia 5-7. The extensive sympathetic innervation of the ankle joint was identified by the large number of cell bodies labelled in the sympathetic ganglia of the paravertebral chain. Labelled afferent fibres projected to the spinal cord from the 2nd to the 8th synsacral nerves, with the rostral projection mainly via Lissauer's tract and the dorsal funiculus. Terminal labelling in the dorsal horn was identified in laminae I-III and VI, with a slight projection to V. Two areas of dense labelling, which did not correspond with the largest number of labelled dorsal root ganglion cells, were identified. A rostral area with the highest density of label was observed at the level of synsacral nerves 3-4 and a second slightly less dense area between synsacral nerves 7-8. In the caudal medulla, diffuse terminal labelling was observed in the nucleus gracilis et cuneatus, nucleus of the tractus solitarius, and the nucleus cuneatus externus. These results are discussed in a comparative context to identify similarities and differences between different primary afferent projections in birds and mammals and to highlight the possible functional significance of the avian articular afferent projection.

Synaptic interactions of substance P immunoreactive nerve terminals in the baro- and chemoreceptor reflexes of the cat

The neurochemical anatomy and synaptic interactions of morphologically identified chemoreceptor or baroreceptor afferents in the nucleus of the solitary tract (NTS) are poorly understood. A substantial body of physiological and light microscopic evidence suggests that substance P (SP) may be a neurotransmitter contained in first order sensory chemo- or baroreceptor afferents, however ultrastructural support for this hypothesis is lacking. In the present report we have traced the central projections of the carotid sinus nerve (CSN) in the cat by utilizing the transganglionic transport of horseradish peroxidase. Medullary tissues including the commissural NTS (cNTS) were processed for the histochemical visualization of transganglionically labeled CSN afferents and for the immunocytochemical detection of SP by dual labeling light and electron microscopic methods. At the light microscopic level, dense bilateral labeling with TMB was found in the tractus solitarius (TS) and cNTS, caudal to the obex. Rostral to the obex, significant ipsilateral TMB labeling was detected in the dorsal, dorso-lateral, and medial subnuclei of the NTS, as well as in the TS. Significant staining of SP immunoreactive processes was detected in most subnuclei of the NTS. The cNTS was examined by electron microscopy. Either HRP or SP were readily identified in single labeled unmyelinated axons, myelinated axons, and nerve terminals in the cNTS. SP immunoreactivity was also identified in unmyelinated axons, myelinated axons, and nerve terminals in the cNTS which were simultaneously identified as CSN primary afferents. These ultrastructural data support the hypothesis that SP immunoreactive first order neurons are involved in the origination of the chemo- and baroreceptor reflexes. Axo-axonic synapses were observed between CSN primary afferent terminals and: (a) unlabeled nerve terminals; (b) other CSN primary afferent terminals; and (c) terminals containing SP. Axo-axonic synapses were also observed between CSN primary afferents which contained SP, and other SP terminals. These observations may mediate the morphological bases for multiple forms of presynaptic inhibition in the cNTS, including those involved in cardiorespiratory integration. In conclusion, our results indicate that SP immunoreactive nerve terminals may be important in both the origination and the modulation of the chemo- and/or baroreceptor reflexes.

Musculotopic organization of muscle afferents in the external cuneate nucleus of the gerbil as revealed by transganglionic transport of horseradish peroxidase.

The projections of muscle afferents from six regions (hand, forearm, arm, thorax, shoulder and neck) to the external cuneate nucleus (ECN) of gerbils were investigated using transganglionic transport of horseradish peroxidase (HRP). These were chiefly ipsilateral projections which terminated in six discrete longitudinal columns in the ECN. A medial cell column confined to the caudal ECN received projections from the hand muscles, while the ventromedial and ventrolateral cell columns in the posterior half of the ECN received projections from the forearm and arm muscles respectively. In the caudal ECN, the thoracic and shoulder muscles were represented by the dorsomedial and dorsolateral cell columns respectively. The last mentioned two columns extended anteriorly into the rostral ECN occupying a dorsal and intermediate portion leaving the ventral portion which represented the neck muscles. The latter also extended posteriorly into the middle 1/3 of ECN occupying a lateral portion. Serial sections and reconstruction studies showed that the ECN was located in the dorsal region of the upper medulla, approximately 0.3-1.8 mm rostral to the obex. The rostrocaudal extent of the six cell columns from hand proximally to neck were measured to be 0.5-0.8, 0.3-1.1, 0.3-1.3, 0.4-1.8, 0.4-1.8 and 0.8-1.8 mm rostral to the obex.

Organization of the nucleus of the solitary tract in the hamster: Acetylcholinesterase, NADH dehydrogenase, and cytochrome oxidase histochemistry

The distribution of acetylcholinesterase (AChE), NADH dehydrogenase (NADHd), and cytochrome oxidase (CO) was determined in the nucleus of the solitary tract (NST) in the golden hamster. Histochemical staining was compared to cytoarchitectonic subdivisions of the NST (Whitehead: J. Comp. Neurol. 276:547-572, 1988) and to terminal fields of primary afferents of the nerves that innervate the tongue. These three histochemical methods resulted in differential staining patterns within the NST that were related to certain subdivisions. Transganglionic transport of horseradish peroxidase (HRP) was used to determine the central projections of the chorda tympani (CT), the lingual branch of the trigeminal (L-V), and the lingual-tonsilar branch of the glossopharyngeal nerves (L-IX). Alternate or the same brain sections were processed to reveal transported HRP, and NADHd or AChE levels. Increased staining of the neuropil with NADHd and AChE was coincident with the dense part of the afferent terminal fields of all three nerves in the NST and the laterally adjacent dorsomedial part of the spinal trigeminal nucleus. CO showed this pattern only for the most rostral part of the CT field. The densest AChE staining coincided with gustatory afferent terminal fields. The histochemical staining facilitated the interpretation of the organization of the NST. For example, at caudal levels of the gustatory NST, it is suggested that taste processing is localized predominantly in the medial part of the rostral central, and somatosensory processing in the rostral lateral subdivision. AChE or NADHd staining should facilitate studies of connections, topography, and neuroplastic changes of the gustatory NST.

The octavolateral systems in the stingray, Dasyatis sabina. I. Primary projections of the octaval and lateral line nerves.

The central projections of the electrosensory, mechanosensory, and octaval nerves of the Atlantic stingray were examined by transganglionic transport of horseradish peroxidase. Particular attention was paid to the relation of the projections to cell plates C1 and C2, and to a newly described cell plate, C3. The electroreceptors in the stingray are found in three groups on the dorsal and ventral sides of the body. The electroreceptors are represented topographically on the ipsilateral dorsal nucleus. Those of the rostral part of the head and pectoral fins are represented on the rostroventral part of the nucleus, and those on the caudal part of the head and the pectoral fin on the dorsocaudal part of the nucleus. Mechanosensory lateral line afferents terminate within the ipsilateral intermediate and caudal nuclei, and the lateral granular mass of the vestibulocerebellum. Anterior lateral line afferents also project to the magnocellular octaval nucleus. A topographic representation of the mechanosensory lateral line periphery is present on the intermediate and caudal nuclei. Mechanoreceptors on the trunk are represented laterally and those on the head medially. The terminal field of the anterior lateral line afferents on the intermediate nucleus surrounds cell plates C1 and C2. The anterior lateral line afferents also project to the medial part of the lateral granular mass, whereas the posterior lateral line afferents project to the lateral portion. Sparse projections of anterior lateral line afferents to the periventricular octaval nucleus were also observed. The octaval nerve afferents terminate largely within the octaval column. Octaval nerve projections were also observed to the reticular formation, the periventricular octaval nucleus, the deep cerebellar nucleus, the vestibulocerebellum, particularly the lower lip and medial granular mass, and the intermediate nucleus. A sparse projection to cell plate C3 was found. The relation between cell plates C1 and C2 and the anterior lateral line afferents suggests that these cell plates are related to processing lateral line information. While the relationship between cell plate C3 and the octaval afferents is not strong, the sparse octaval projection C3 receives suggests that it is relaying octaval information.

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

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

The Central Projections of the Monkey Tooth Pulp Afferent Neurons

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

Pudendal Nerve Topography In The Rat Spinal Cord Projections Studied With The Axonal Tracer Wheat Germ Agglutinin Conjugated-Horseradish Peroxidase

The spinal cord termination of the sensory fibers of the rat pudendal nerve has been determined by using the transganglionic transport of horseradish peroxidase conjugated to wheat germ agglutinin. The gross anatomical examination of the pudendal plexus indicated the existence of dorsal and ventral branches in the sensory division of the pudendal nerve. Either branch or both were sectioned and exposed to a 1% solution of the enzyme. After an average survival period of 48hours, the resulting reaction product was studied both at the level of the dorsal root ganglia and the spinal cord. Labeled dorsal root ganglion cells were observed between LI and S4 (peak in SI and L6). Evidence of transganglionic transport of the tracer was observed from L5 to S2. The ventral branch projected medially in the superficial dorsal horn, while the dorsal branch projected laterally. Our findings are consistent with a segregation of sensibilities for the ventral and dorsal branches that convey urogenital (ventral branch) and anal (dorsal branch) cutaneous sensations at the level of termination of primary afferents.

The ganglionic origins and central projections of primary sensory neurons innervating the upper and lower lips in the rat.

Retrograde and transganglionic transport of horseradish peroxidase (HRP) was used to investigate the neurons innervating the upper and the lower lips and their central projections in the rat. Both the upper and the lower lips were observed to be innervated by a very large number of trigeminal sensory neurons, with their cell bodies located in the maxillary and the mandibular parts of the trigeminal ganglion, respectively. The central projections of neurons innervating the upper lip formed a long continuous column starting rostrally at midlevels of the trigeminal main sensory nucleus (5P) and extending caudally through the C1 dorsal horn, with occasional fibers reaching the C3 segment. The heaviest projections appeared in the middle portions of 5P and nucleus interpolaris (5I), as well as in the rostral part of nucleus caudalis (5C). A small but consistent projection to the solitary tract nucleus, originating from cells in the inferior vagal ganglion, was observed in the upper-lip experiments. The central projections from neurons innervating the lower lip also appeared as a long column located dorsally or dorsomedially to the projections from the upper lip. The most prominent projections from the lower lip were located in the caudal part of 5P, the middle part of 5I, and the caudal two-thirds of 5C. Sparse projections could be traced as far caudally as C4. At 5C and cervical levels, some labeling appeared contralaterally in the same location as on the ipsilateral side.

Somatotopic organization of hindlimb skin sensory inputs to the dorsal horn of hatchling chicks (Gallus g. domesticus)

The somatotopic organization of skin sensory nerve projections to the lumbosacral dorsal horn of hatchling chickens was determined with the aid of transganglionic transport of horseradish peroxidase (HRP) processed with tetramethylbenzidine histochemistry. A total of eight hindlimb nerves were studied, five of which were purely cutaneous. When combined, the innervation fields of these nerves covered most of the hindlimb surface, allowing a nearly complete somatotopic map of the hindlimb to be generated. This report describes a novel pattern of cutaneous nerve projections to the dorsal horn. Unlike other vertebrates, cutaneous nerves of chickens formed two separate, somatotopically organized projections across the mediolateral axis of the dorsal horn; when serially reconstructed and superimposed, these projections produced two nonoverlapping somatotopic maps of the skin surface lying side by side. Each of these separate maps was nearly identical to the other in overall topology. These two separate maps appear to represent distinct modalities of sensory information, as projections composing the medial map were preferentially labeled by choleragenoid-HRP, whereas those composing the lateral map were preferentially labeled by wheat germ agglutinin-HRP. In mammals, these HRP ligands selectively label the central projections of myelinated and unmyelinated cutaneous afferents, respectively. The present study, therefore, strongly supports the cytoarchitectonic findings of Brinkman and Martin (Brain Res. 56:43-62, '73) that lamina III lies medial, rather than ventral, to lamina II in the chicken dorsal horn. Further, the present studies also suggest that laminae II and III of chickens are homologous to the homonymous laminae in the dorsal horn of mammals.

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 (1.4x compared with 2.0x), 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.

Brainstem projections of different branches of the vestibular nerve: An experimental study by transganglionic transport of horseradish peroxidase in the cat

The brainstem projections of the ampullar nerves from the vertical semicircular canals, the anterior (AAN) and the posterior ampullar nerve (PAN), were studied in adult cats using the transganglionic horseradish peroxidase (HRP) method. Each nerve was exposed in three experiments. Two animals in each group had labeling which allowed detailed mapping. From the AAN, terminal-type labeling was found in two separate groups, one laterally and one medially, both in the lateral (LV) and in the superior (SV) vestibular nucleus. In addition, such labeling was found in all parts of the medial vestibular nucleus (MV). Labeled structures were found also in the descending vestibular nucleus, (DV) more densely over its lateral part, except for cell group f, where no labeling was found. From the PAN, terminal-type labeling was found medially and laterally in the LV and in the medial part of the SV. In the MV, such labeling was evenly distributed rostrally but concentrated laterally in caudal parts. In the DV, terminal-type labeling was present rostrally, whereas no labeling was seen caudally. In the interstitial nucleus of the vestibular nerve, terminal-type labeling was observed from the AAN but not from the PAN. No labeled fibers from either of the two ampullar nerves were seen outside the vestibular root and nuclei, except for small-caliber fibers from the SV heading towards the brachium conjunctivum. The findings clearly indicate a specific termination for each of the two ampullar nerves.

Effects of cortical and thalamic lesions upon primary afferent terminations, distributions of projection neurons, and the cytochrome oxidase pattern in the trigeminal brainstem complex

Early postnatal lesions of the primary somatosensory cortex alter the vibrissa-related cytochrome oxidase (CO) pattern in nucleus principalis (PrV) of the rat's trigeminal (V) brainstem complex (Erzurumlu and Ebner, '88: Dev. Brain Res. 44:302-308). At present, the reason for this change is not clear. It may be that the corticotrigeminal projection is necessary for the maintenance of vibrissa-related patterns in PrV. However, it is also possible that the loss of the normal pattern of CO activity reflects a change in the organization of brainstem cells resulting from transneuronal retrograde degeneration. In order to address this question, we made lesions of either the primary somatosensory cortex (S-I) or ventrobasal thalamus (VB) in newborn rats and directly assayed distribution of V primary afferents by transganglionic transport of horseradish peroxidase and V-thalamic neurons by retrograde transport of either fluorogold or true blue. Neonatal cortical and thalamic lesions produced no qualitative change in the distribution of primary afferent terminals in either PrV or V subnucleus interpolaris (SpI) beyond that which could be attributed to shrinkage of the brainstem resulting from retrograde degeneration. Most importantly, the "patchy" pattern of terminations observed in normal rats remained apparent in the brain-damaged animals. The normal distribution of V-thalamic neurons in PrV was disrupted by both cortical and thalamic lesions. These cells are normally patterned in a way that matches the distribution of primary afferent terminals and thus that of the mystacial vibrissae. This was not the case in the neonatally brain-damaged rats. Taken together, these results are consistent with the conclusion that neonatal cortical and thalamic lesions disrupt the normal CO pattern in PrV primarily because of their effects upon the patterning of brainstem cells. The present findings demonstrate further that clustering of primary afferents does not require a normal complement of postsynaptic neurons.

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 (C 1–C 5). 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 C 4–C 5. Contralateral buccal and lingual endings were scattered sparsely from the subnucleus caudalis to C 5, 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 II i–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 C 4 and C 5 lingual endings appeared exclusively in laminae I and II o, 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.

Corpuscular Bodies in the Palate of the Rat

The source of innervation of the corpuscular bodies in the palate and the central projections of the afferent fibres of the entire palate was studied in rats by transganglionic transport of horseradish peroxidase conjugated to wheat germ agglutinin (WGA-HRP) and with substance P (SP) immunohistochemistry. WGA-HRP injected into the incisal papilla was taken up by the nerve fibres that terminated in the corpuscles. Retrogradely labelled neurons were observed in the trigeminal ganglion as well as anterogradely labelled terminals in the dorsolateral part of the spinal trigeminal nucleus and in the lateral part of the nucleus of the solitary tract. No labelling could be found in the geniculate ganglion, the facial nerve and the hypoglossal nucleus. Following WGA-HRP injection in the intermolar area and in the soft palate, labelling was only restricted to the trigeminal ganglion. The lamina propria of the entire palate and the corpuscle-enriched area of the incisal papilla and the soft palate were richly innervated by SP-containing fibres. Numerous SP-containing fibres were also observed in the nerve plexus at the base of the corpuscle. In addition, SP-positive neurons were identified in the trigeminal ganglion and SP-labelled terminals in the sensory trigeminal nuclear complex and in the solitary tract nucleus. On the basis of our morphological observations we conclude that the palatal corpuscular bodies are involved in taste perception which is of trigeminal origin.