Afferent connections of the oculomotor nucleus in the chick

Horseradish peroxidase was injected into the oculomotor nucleus of the chick in order to locate and characterize the neurons projecting to this nucleus. In the rostral mesencephalon, 120-180 neurons were labelled in the medial area of the ipsilateral nucleus campi Foreli; 190-220 in the interstitial nucleus of Cajal (most of them contralateral); and smaller numbers bilaterally in the medial mesencephalic reticular formation, the nucleus of the basal optic root complex, and the central grey matter. More caudally, numerous neurons were labelled in the contralateral abducens nucleus and the vestibular complex and a few in the nucleus reticularis pontis caudalis. Labelled neurons appeared ipsilaterally in the caudal region of the nucleus vestibularis superior and in the rostral tip of the nucleus descendens just lateral to the tractus lamino-olivaris. In the contralateral vestibular complex, a group of labelled cells observed in the dorsolateral area may be homologous to the mammalian cell group Y. At the level of the contralateral abducens nucleus, the most numerous group of cells (625-700) projecting to the oculomotor nucleus formed a lateromedial fringe that affected the nucleus tangentialis, the rostral tip of the nucleus descendens, and the ventrolateral region of the nucleus medialis. Only a few labelled neurons were seen in the contralateral nucleus vestibularis superior, the ipsilateral cell group A, and the ipsilateral nucleus vestibularis medialis.     

Projections of the vestibular and cerebellar nuclei in Rana pipiens

The efferent projections and cytoarchitecture of the vestibulocerebellar region were examined to determine the nuclear boundaries and potential homologies. The anterior portion of the vestibular complex projects to the ipsilateral oculomotor and trochlear nuclei and is the major source of commissural fibers. Neurons in the rostromedial portions of the complex project to the contralateral trochlear nucleus. Large neurons in the ventrolateral portion of the complex give rise to a bilateral vestibulospinal pathway. Medium-sized neurons in the neuropil and small neurons in the central gray giving rise to bilateral projections to the spinal cord and oculomotor nuclei as well as commissural and ipsilateral cerebellar efferents. Projections from the nucleus of the cerebellum reach the contralateral spinal cord and cerebellar nucleus and there is also a bilateral projection to the ventral rhombencephalic and mesencephalic basal plates. The medial portion of the nucleus gives rise to commissural, ipsilateral mesencephalic and contralateral spinal projections. The lateral portion of the nucleus projects to the contralateral ventral mesencephalon. On the whole, the results of this investigation substantiate the division of the anuran vestibular complex in anurans into nuclei which may be homologous to the superior nucleus and nucleus of Deiters in mammals. The case for distinct descending and medial nuclei is less compelling. Further, it appears possible to divide the nucleus of the cerebellum into medial and lateral components whose connectivity is similar to that of reptiles and to a lesser extent mammals.     

Anatomical and physiological characteristics of vestibular neurons mediating the vertical vestibulo-ocular reflexes of the squirrel monkey

The morphology of 35 vestibular neurons whose firing rate was related to vertical eye movements was studied by injection of horseradish peroxidase intracellularly into physiologically identified vestibular axons in alert squirrel monkeys. The intracellularly injected cells were readily classified into four main groups. One group of cells, down position-vestibular-pause neurons (down PVPs; N = 12), increased their firing rate during downward eye positions, paused during saccades, and were located in the medial vestibular nucleus (MV) and the adjacent ventrolateral vestibular nucleus (VLV). They had axons that crossed the midline and ascended in the medial longitudinal fasciculus (MLF) to terminate in the trochlear nucleus, the lateral aspect of the caudal oculomotor nucleus, and the dorsal aspect of the rostral oculomotor nucleus. A second group of cells (N = 15) were also located in the MV and VLV, but increased their firing rate during upward eye positions, and paused during saccades. These cells had axons that crossed the midline and ascended in the contralateral MLF to terminate in the medial aspect of the oculomotor nucleus. A third group of cells (N = 4) were located in the superior vestibular nucleus, generated bursts of spikes during upward saccades, and increased their tonic firing rate during upward eye positions. These cells had axons that ascended laterally to the ipsilateral MLF to terminate in regions of the trochlear and oculomotor nuclei similar to those in which down PVPs terminated. A fourth group of cells (N = 4), located in the VLV, had axons that projected to the spinal cord, although they had firing rates that were significantly correlated with vertical eye position. Electrical stimulation of the vestibular nerve evoked spikes at monosynaptic latencies in each of the above classes of cells, six of which were injected with horseradish peroxidase. Each group of cells had collateral projections to other areas of the brainstem. Some of the neurons that projected to the contralateral trochlear and oculomotor nuclei had collaterals that crossed the midline to terminate in the oculomotor nucleus ipsilateral to the soma, and some gave rise to small collaterals that terminated in the abducens nucleus. Other areas of the brainstem that received collateral inputs from neurons projecting to oculomotor and trochlear nuclei included the interstitial nucleus of Cajal, the caudal part of the dorsal raphe nucleus, the nucleus raphe obscurus, Roller's nucleus, the intermediate and caudal interstitial nuclei of the MLF, and the nucleus prepositus.     

A comparative study of the tangential-oculomotor projection.

Most of the neurons of the tangential vestibular nucleus of birds project to the controlateral oculomotor complex, but it is not known whether there is a homologous projection in mammals. In this study, horseradish peroxidase (HRP) was injected into the oculomotor nucleus of chicks and rabbits, and the distributions of labelled neurons in the target region of the vestibular complex in the two species were compared. In chicks, a large number of labelled neurons formed a continuous band of neurons located in the contralateral tangential, descending and medial nuclei. In rabbits a similar band of labelled neurons was found in the contralateral descending and medial vestibular nuclei, but most of the neurons were caudal to the incoming vestibular nerve fibers, and only a few rostral neurons were located among these fibers. Our results suggest that the tangential nucleus neurons projecting to the oculomotor nucleus may be homologous to the most lateral neurons of the neuronal band of rabbits.     

Organization of the cerebellum in the pigeon (Columba livia): III. Corticovestibular connections with eye and neck premotor areas.

The connections of the cerebellar cortex with vestibular premotor neurons of the oculomotor and collimotor systems in the pigeon were delineated in experiments using WGA-HRP as an anterograde and retrograde tracer. Putative premotor neuron pools were identified by injections into the oculomotor (mIII) and trochlear nuclei (mIV) and into the most rostral portion of the cervical neck motor nucleus, nucleus supraspinalis (SSp). The retrograde data indicate that ipsilateral projections upon oculomotor neurons arise from the medial portions of the superior (VeS) and tangential (Ta) nuclei. Contralateral projections originate from the infracerebellar nucleus, the interstitial vestibular region including the main (lateral) portion of the tangential nucleus, and from the descending and medial vestibular nuclei (VeD, VeM). These projections were confirmed in anterograde studies that also defined the connections of these vestibular premotor regions with specific subnuclear divisions of the pigeon's "oculomotor" nuclei (mIII, mIV, mVI). The organization of projections from the vestibular nuclei to the pigeon's extraocular motoneurons is similar to that reported in mammals. Projections upon neck premotor neurons arise primarily from neurons in the interstitial region of the vestibular nuclear complex. After injections in SSp, retrogradely labeled neurons were found, contralaterally, in the lateral part of the tangential and superior vestibular nuclei and in the dorsolateral vestibular nucleus (VDL). Ipsilateral labeling was seen in the medial interstitial region (VeM, VeD, and medial Ta). These projections were confirmed in anterograde experiments. With the exception of VDL, vestibular nuclei projecting to neck motoneurons also project to extraocular motoneurons. Thus the infracerebellar nucleus projects exclusively, and the superior vestibular nucleus predominantly, upon oculomotor (mIII, mIV) nuclei; VDL projects predominantly upon the neck motor nucleus, whereas the interstitial vestibular regions (medial Ta, rostral VeD, intermediate VeM) project upon both collimotor and oculomotor neurons. The pattern of retrograde labeling seen in the cerebellar cortex after injections into vestibular premotor nuclei was used to define the projections of specific cerebellar cortical zones upon vestibular eye and neck premotor neurons. Corticovestibular projections upon these regions arise from the auricle and lateral unfoliated cortex, the posterior lobe components of cortical zones B and E, and from the vestibulocerebellum. Each of these cortical zones projects upon components of the vestibular nuclear complex, which are premotor to either oculomotor nuclei or collimotor nuclei. The hodological findings are related to the functional organization of the oculomotor and collimotor systems in the pigeon and compared with the mammalian data.     

The vestibular nuclei in the domestic hen (Gallus domesticus) III. Ascending projections to the mesencephalic eye motor nuclei

Following injections of horseradish peroxidase in the oculomotor and the trochlear nuclei in the hen, the occurrence of labeled cells was plotted in the vestibular nuclei. The majority of labeled cells was localized in the superior, the medial, and the tangential nucleus. Within the superior nucleus the cells were found mainly caudally, extending medially and ventrally in central areas. In the medial nucleus labeled cells were localized exclusively in its rostral half, mainly in ventrolateral regions. Most, if not all, cells in the nucleus tangentialis project rostrally. In addition, rostrally projecting vestibular cells were found in the cell group A and the rostrolateral part of the descending nucleus. The projection to the oculomotor nuclear complex is from the superior nucleus and the cell group A bilateral but chiefly ipsilateral, from the medial nucleus bilateral, from the tangential nucleus and the rostral pole of the descending nucleus chiefly contralateral. Massive labeling was found in the abducens nucleus, somewhat less in the reticular formation, mainly in the lateral regions of the medial part at the level of the abducens and facial nuclei. Labeled cells were, in addition, found in the deep layers of the optic tectum, and scattered cells in the nucleus raphe. The findings are discussed in the light of what is known of the organization of the vestibular nuclei in the hen and the rostral projection of the vestibular nuclei in mammals.     

Studies of suspected neurotransmitters in the vestibuloocular pathways.

Isolated fresh cat trochlear and oculomotor nuclei, which contain the axon terminals of inhibitory neurons whose cell bodies are in the superior vestibular nucleus (SVN), actively synthesize and store [3H]GABA, [14C]acetylcholine, [3H]dopamine and [3H]tyramine from labeled precursors of these compounds. Twelve to 14 days following lesions of the ipsilateral superior vestibular nucleus or its efferent pathway to the oculomotor and trochlear nuclei, at a time when there is extensive degeneration of superior vestibular nucleus axon terminals in these nuclei, the synthesis and storage of GABA in the ipsilateral trochlear nucleus is markedly reduced compared to that in the contralateral trochlear nucleus; the synthesis of acetylcholine, dopamine and tyramine is not measurably affected. The oculomotor nuclei, which unlike the trochlear nuclei receive a heavy bilateral projection from the SVN, show no asymmetric decrease after SVN lesions in their ability to synthesize any of the compounds tested. The data support the identity of GABA as an inhibitory transmitter in the superior vestibular nucleus-trochlear nucleus pathway.     

A study of some of the ascending and descending vestibular pathways in the pigeon (Columba livia) using anterograde transneuronal autoradiography

A mixture of tritiated proline and fucose was injected into the endolymph of one of the membranous labyrinths of each of 5 white king pigeons (Columba livia). The membranous labyrinth was resealed and the animals were allowed to survive for 15 days. Brains and upper parts of the spinal cords were sectioned and processed by standard autoradiographic procedures. Clear labeling was noted in structures usually associated with both the ascending auditory pathways and the ascending and descending vestibular pathways. Vestibular structures ipsilateral to the injected labyrinth which contained heavy labeling were Scarpa's ganglion and all 6 vestibular nuclei. No labeling was noted in the contralateral Scarpa's ganglion and sparce, if any, labeling was noted in the contralateral vestibular nuclei. Contralateral structures associated with ascending vestibulo-ocular pathways which contained heavy labeling were the medial longitudinal fasciculus, abducens nucleus, trochlear nucleus, and two parts of the oculomotor nucleus--the dorsolateral part and the ventromedial part. Less heavily labeled ipsilateral vestibulo-ocular-related structures included the medial longitudinal fasciculus, abducens nucleus and the ventrolateral edge of the trochlear nucleus. The dorsomedial part of the oculomotor nucleus was heavily labeled on the side ipsilateral to the injected labyrinth. Slight, if any, labeling was noted in either the ipsilateral or contralateral brachium conjunctivum or regions corresponding to the mammalian ascending tract of Deiters. The medullary core of most folia but primarily the medullary core and granular areas of folia IX and X of the cerebellum were labeled.(ABSTRACT TRUNCATED AT 250 WORDS)     

Afferent connections of the lateral vestibular nucleus in the cat

Localization of labelled neurons (sources of projections to the lateral vestibular nucleus) in the brain was studied by means of microiontophoretic injections of horseradish peroxidase. Bilateral projections of the midbrain structures to all parts of the nucleus mentioned (field of Forel, interstitial nucleus of Cajal, oculomotor nerve nucleus and red nucleus) were found. There have been also shown: bilateral projections from more caudally localized structures of the superior, medial and inferior (descending) vestibular nuclei, group "Y" of the vestibular nuclear complex, facial nucleus and hypoglossi, nucleus prepositus nervi hypoglossi and spinal trigeminal nucleus; ipsilateral projections from crus IIa of lobulus ansiformus of the cerebellar hemisphere; contralateral projections from lateral reticular nucleus of medulla oblongata and Deiters' nucleus. Trajectories of the labelled fibre system projecting to Deiters' nucleus are described.     

Transneuronal transport in the vestibular and auditory systems of the squirrel monkey and the arctic ground squirrel. I. Vestibular system

Transneuronal transport of [3H]proline, [3H]fucose, and [3H]leucine in various combinations from pledgets implanted in the ampulla of a single semicircular duct was studied in the squirrel monkey and arctic ground squirrel after long survival periods. Tritiated amino acids implanted in any single ampulla resulted in labeling of nearly all vestibular and auditory receptors, nearly all cells of the vestibular and spiral ganglia and central transport via nearly all root fibers of both nerves. Primary vestibular fibers were distributed to the vestibular nuclei (VN) and specific parts of the cerebellum in the pattern previously described. Transneuronal transport of [3H]proline by vestibular neurons was present in all known secondary pathways, except those projecting to thalamic nuclei. Observations were similar in both species, except for small differences in commissural vestibular projections. Major commissural transport was to all parts of the opposite medial vestibular nucleus (MVN) and to peripheral parts of the superior vestibular nucleus (SVN), but some transport was present in all contralateral VN, including ventral cell group y. Descending transneuronal transport was evident in vestibulospinal tract (VST) ipsilaterally and in the medial longitudinal fasciculus (MLF) bilaterally. Both [3H]proline and [3]fucose were transported transneuronally to the ipsilateral abducens nucleus (AN); with long survivals [3H]proline was transported peripherally via the ipsilateral abducens nerve root. Ascending transport in the MLF was bilateral, asymmetric and greatest contralaterally. Fibers entered the contralateral MLF near the AN and the lateral wing of the ipsilateral MLF rostral to most of the VN. Terminals in the trochlear nuclei (TN) were bilateral and greatest contralaterally. In the monkey terminals in ipsilateral oculomotor complex (OMC) were distributed uniformly in all subdivisions, except for the medial rectus subdivision (MRS), where terminals were more numerous. The greatest density of terminals was present contralaterally in the superior rectus subdivision (SRS) of the OMC; only sparse terminals were present in the MRS on that side. Transport in the ipsilateral abducens nerve roots in the monkey and the virtual absence of transport to the MRS of the contralateral OMC suggested transneuronal transport to abducens motor neurons, but not to internuclear neurons (AIN). The AIN project only to the MRS of the contralateral OMC and do not appear to receive vestibular input. Comparable observations were made in the AN, TN and OMC of the ground squirrel, although the representation of the extraocular muscles in the OMC is unknown.(ABSTRACT TRUNCATED AT 400 WORDS)     

Brain stem projections to the facial nucleus of the rat

Horseradish peroxidase was injected into the medial and lateral columns of the facial nucleus of the rat. Following medial injections, cells were labelled by retrograde transport in the ipsilateral spinal trigeminal nucleus (caudalis) both medial vestibular nuclei, contralateral midbrain reticular formation and nucleus of the lateral lemniscus. The periaqueductal grey, interstitial nucleus and nucleus of Darkschewitch were also labelled ipsilaterally. Injections into the lateral column of the facial nucleus labelled the spinal trigeminal nucleus (oralis) and parabrachial nuclei ipsilaterally and the Darkschewitch and red nuclei contralaterally.     

Retrograde transport of [3H]glycine from the cochlear nucleus to the superior olive in the guinea pig

The origins of descending glycinergic projections to the guinea pig cochlear nucleus were investigated using retrograde labelling techniques. To identify the cell groups that provide descending projections to the cochlear nucleus, horseradish peroxidase, a nonspecific retrograde neuronal marker, was injected into the cochlear nucleus. After 24 or 48 hours, labelled cell bodies were evident bilaterally in all of the periolivary nuclei that surround the lateral and medial superior olive. The largest numbers of labelled neurons were located in the ventral nucleus of the trapezoid body bilaterally and in the lateral nucleus of the trapezoid body and dorsal periolivary nucleus ipsilaterally. Labelled cells were also present in the inferior colliculus bilaterally and in the contralateral cochlear nucleus. [3H]Glycine was employed as a retrograde tracer to identify the cell groups providing descending glycinergic projections to the cochlear nucleus. Three to 48 hours after injection of 19, 190, or 380 microM [3H]glycine into the cochlear nucleus, retrogradely labelled cell bodies were observed ipsilaterally in all of the periolivary nuclei. No labelled neurons were found in the inferior colliculus. After injections of the highest concentration of [3H]glycine, labelled cells were also found contralaterally in the ventral and lateral nuclei of the trapezoid body and also in the contralateral cochlear nucleus. We conclude that descending glycinergic projections to the cochlear nucleus originate mostly in ipsilateral periolivary cell groups. Minor glycinergic projections originate from the contralateral cochlear nucleus and also from the contralateral ventral and lateral nuclei of the trapezoid body.     

Afferent and efferent connections of the oculomotor region of the fastigial nucleus in the macaque monkey.

Afferent and efferent connections of the fastigial oculomotor region (FOR) were studied in macaque monkeys by using axonal transport of wheat germ agglutinin conjugated horseradish peroxidase (WGA-HRP). When injected HRP is confined to the FOR, retrogradely labeled cells appear in lobules VIc and VII of the ipsilateral vermis and in group b of the contralateral medial accessory olive (MAO). In reference to the maps of topographical organization, the extent of the effective site in the fastigial nucleus (FN) could be assessed from the distributions of labeled Purkinje cells (P cells) in the vermis and labeled olivary neurons in the MAO. In contrast to the unilateral nature of the P-cell and climbing-fiber projections, those from the other brainstem regions to the FOR were bilateral. Following the injection of HRP into the FOR, the largest number of retrogradely labeled cells appeared in the pontine nuclei. Although the number of labeled cells was greater on the contralateral side in both the peduncular and dorsomedial pontine nuclei (DMPN), the number of each side was virtually identical in the dorsolateral pontine nucleus (DLPN). In the nucleus reticularis tegmenti pontis (NRTP), labeled cells were located only in its medial and dorsolateral portions bilaterally. In the vestibular complex, labeled cells appeared in the superior (SVN), medial (MVN), and inferior vestibular nuclei (IVN) bilaterally. The lateral vestibular nucleus (LVN), including y group and the ventrolateral vestibular nucleus, were free of labeled cells. Labeled cells appeared also in the perihypoglossal nucleus (PHN) bilaterally. In the pontine raphe (PR) and paramedian pontine reticular formation (PPRF), labeled cells appeared bilaterally in the caudal third of the area between the oculomotor and abducens nuclei. Labeled cells appeared also in the mesencephalic and medullary reticular formation. Tracing of anterogradely labeled axons demonstrated that most fibers from the FOR decussated within the cerebellum and entered the brainstem via the contralateral uncinate fasciculus. Some crossed fibers ascended with the contralateral brachium conjunctivum and terminated in the midbrain tegmentum. A small contingent of fibers advanced further to the thalamus. In the mesodiencephalic junction, labeled terminals were found contralaterally in the rostral interstitial nucleus of medial longitudinal fasciculus (riMLF) and a medial portion of FOrel's H Field. They appeared also in the central mesencephalic reticular formation (cMRF), the periaqueductal gray (PAG), the posterior commissure nucleus, and the superior colliculus. The oculomotor and trochlear nuclei, the red nucleus, and the interstitial nucleus of Cajal were free of labeled terminals.(ABSTRACT TRUNCATED AT 400 WORDS)     

Auditory brainstem of the ferret: sources of projections to the inferior colliculus

The organization of the auditory brainstem in adult, darkly pigmented ferrets was studied by using the retrograde transport of the lectin wheat germ agglutinin-horseradish peroxidase injected into one inferior colliculus. Retrogradely labelled neurons were found bilaterally in every nucleus of the auditory brainstem. The greatest number of labelled neurons was found in the cochlear nuclei contralateral to the injection site, the ipsilateral medial superior olivary nucleus, both lateral superior olivary nuclei, the ipsilateral ventral nucleus of the lateral lemniscus, both dorsal nuclei of the lateral lemniscus, and the contralateral inferior colliculus. Quantitative assessment of the projections from the cochlear nuclei showed that the number of contralaterally projecting neurons exceeded the number of ipsilaterally projecting neurons by about 50 to one. This ratio remained relatively stable over a wide range of volumes of injected lectin, whereas the absolute number of labelled neurons on each side varied by at least twofold for a constant volume of lectin. These results provide basic data on the ferret auditory system and demonstrate quantitatively some properties of the projections between the cochlear nucleus and the inferior colliculus.     

Cerebellar efferents in the lizard Varanus exanthematicus. II. Projections of the cerebellar nuclei

The projections of the cerebellar nuclei have been studied in the lizard Varanus exanthematicus with various experimental anatomical techniques. In anterograde degeneration experiments (lesions of the cerebellar peduncle) both ascending and decending contralateral projections were found. Ascending fibers which could be traced from the cerebellar commissure ventralward decussated at the level of the trochlear and oculomotor nuclei. These fibers coursed rostralward to the mesodiencephalic junction. With anterograde tracing techniques (3H-leucine and HRP) this tract was found to terminate in the nucleus ruber and the interstitial nucleus of the fasciculus longitudinalis medialis. Moreover, retrograde tracer studies (HRP, "Fast Blue") showed that this tract appeared to arise mainly in the lateral cerebellar nucleus. With both anterograde degeneration and tracing techniques (3H-leucine and HRP) a bundle of fibers could be followed, which decussates in the basal part of the cerebellum and passes dorsally around the contralateral medial cerebellar nucleus to the lateral side of the brainstem. This contralaterally descending projection system was found, lateral to the vestibular nuclear complex, and as far caudally as the descending vestibular nucleus, to terminate on various vestibular nuclei. Horseradish peroxidase studies showed that this contralaterally descending projection system originates mainly in the medial cerebellar nucleus, but ipsilaterally descending projections were also found. With the fluorescent double labeling technique ("Fast Blue" and "Nuclear Yellow") the projections of the cerebellar nuclei described above were confirmed. Furthermore, double labeling revealed neurons in both cerebellar nuclei (especially the medial nucleus) that project to both the mesencephalon and the cervical spinal cord. The present results indicate that the efferent connections of the cerebellar nuclei in the lizard Varanus exanthematicus are organized as two main projections, an ascending projection comparable to the mammalian brachium conjunctivum arising in the lateral cerebellar nucleus, and a descending projection comparable to the mammalian hook bundle (fasciculus uncinatus), originating mainly in the medial cerebellar nucleus. Such projections are common for terrestrial vertebrates.     

Extra- and intracellular HRP analysis of the organization of extraocular motoneurons and internuclear neurons in the guinea pig and rabbit

The distribution of extraocular motoneurons and abducens and oculomotor internuclear neurons was determined in guinea pigs by injecting horseradish peroxidase (HRP) into individual extraocular muscles, the abducens nucleus, the oculomotor nucleus, and the cerebellum. Motoneurons in the oculomotor nucleus innervated the ipsilateral inferior rectus, inferior oblique, medial rectus, and the contralateral superior rectus and levator palpebrae muscles. Most motoneurons of the trochlear nucleus projected to the contralateral superior oblique muscle although a small number innervated the ipsilateral superior oblique. The abducens and accessory abducens nuclei innervated the ipsilateral rectus and retractor bulbi muscles, respectively. The somata of abducens internuclear neurons formed a cap around the lateral and ventral aspects of the abducens nucleus. The axons of these internuclear neurons terminated in the medial rectus subdivision of the contralateral oculomotor nucleus. At least two classes of guinea pig oculomotor internuclear interneurons exist. One group, located primarily ventral to the oculomotor nucleus, innervated the abducens nucleus and surrounding regions. The second group, lying mainly in the dorsal midline area of the oculomotor nucleus, projected to the cerebellum. Intracellular staining with HRP demonstrated similar soma-dendritic organization for oculomotor and trochlear motoneurons of both guinea pigs and rabbits. Dendrites of oculomotor motoneurons radiated symmetrically from the soma to cover approximately one-third of the entire nucleus, and each motoneuron sent at least one dendrite into the central gray overlying the oculomotor nucleus. In both species, a small percentage of oculomotor motoneurons possessed axon collaterals that terminated both within and outside of the nucleus. The dendrites of trochlear motoneurons extended into the medial longitudinal fasciculus and the reticular formation lateral to the nucleus. Our data on the topography of motoneurons and internuclear neurons in the guinea pig and soma-dendritic organization of motoneurons in the guinea pig and rabbit show that these species share common organizational and morphological features. In addition, comparison of these data with those from other mammals reveals that dendritic complexity (number of dendrites per motoneuron) of extraocular motoneurons exhibits a systematic increase with animal size.     

Evidence for glycine as an inhibitory neurotransmitter of vestibular, reticular, and prepositus hypoglossi neurons that project to the cat abducens nucleus

The localization and distribution of brain-stem afferent neurons to the cat abducens nucleus has been examined by high-affinity uptake and retrograde transport of 3H-glycine. Injections of 3H-glycine selectively labeled (by autoradiography) only neurons located predominantly in the ipsilateral medial vestibular and contralateral prepositus hypoglossi nuclei, and in the contralateral dorsomedial reticular formation, the latter corresponding to the location of inhibitory burst neurons. The specificity of uptake and retrograde transport of 3H-glycine was indicated by the absence of labeling of the dorsomedial medullary reticular neurons ipsilateral and in close proximity to the injection site, where local uptake by diffusion could have occurred. The selectivity of uptake and transport was demonstrated by the absence of retrograde labeling following injections of 3H-GABA or 3H-leucine into the abducens nucleus. The immunohistochemical localization of glycine and GABA revealed a differential distribution of the 2 inhibitory neurotransmitter candidates in the extraocular motor nuclei. Glycine-immunoreactive staining of synaptic endings in the abducens nucleus was dense with a widespread soma-dendritic distribution but was sparse in the trochlear and oculomotor nuclei. By contrast, GABA-immunoreactive staining within the oculomotor and trochlear nuclei was associated with synaptic endings that were particularly prominent on the somata of motoneurons. GABA-immunoreactive staining in the abducens nucleus, however, was sparse. These differences between glycine- and GABA-immunoreactive staining in the extraocular motor nuclei were correlated with differences in the immunoreactivity of axons in the descending (glycine) and ascending (GABA) limbs of the medial longitudinal fasciculus. Glycine-immunoreactive neurons, furthermore, were observed in the same locations as neurons that were labeled autoradiographically by retrograde transport of 3H-glycine from the abducens nucleus. Electrophysiological recordings from abducens motoneurons and internuclear neurons revealed a marked reduction in the slow positivity of the orthodromic extracellular potential elicited by ipsilateral vestibular nerve stimulation following systemic administration of strychnine, an antagonist of glycine. Intracellular recordings demonstrated that the vestibular-evoked disynaptic inhibitory postsynaptic potentials in abducens neurons were effectively blocked by strychnine but were unaffected by picrotoxin, an antagonist of GABA.(ABSTRACT TRUNCATED AT 400 WORDS)     

Rhombomeric organization of vestibular pathways in larval frogs

Rhombencephalic subnuclei and projection pathways related to vestibular function were mapped in larval ranid frogs. The retention of overt postembryonic rhombomeres (r) allowed direct visualization of the locations of neurons retrogradely labeled with fluorescent dextran amines from the midbrain oculomotor complex, cerebellum, vestibular nuclei, and spinal cord. Oculomotor projecting vestibular neurons were mainly located in bilateral r1/2, ipsilateral r3, and contralateral r5-8, and spinal projecting vestibular neurons mainly in ipsilateral r4 and contralateral r5. Vestibular commissural neurons were located in r1-3 and r5-7 and were largely excluded from r4. Cerebellar projecting neurons included contralateral inferior olivary neurons in r8 and vestibular neurons in bilateral r6/7 and contralateral r1/2. Mapping these results onto adult anuran vestibular organization indicates that the superior vestibular nucleus derives from larval r1/2, the lateral vestibular nucleus from r3/4, and the major portions of the medial and descending vestibular nuclei from r5-8. The lateral vestibulospinal tract projects from an origin in r4, whereas a possible ascending tract of Deiters arises in r3. Rhombomere 5 contains a nuclear group that appears homologous to the tangential nucleus of fish, reptiles, and birds and thus likely serves gravistatic and linear vestibulomotor reflexes. Comparisons between frogs and other vertebrates suggest that vestibular neurons performing similar computational roles during head movements originate from the same segmental locations in different species.     

Internuclear neurons in the ocular motor system of frogs.

Medial and lateral rectus motoneurons of frogs were localized after retrograde labeling with horseradish peroxidase (HRP) injected in the medial rectus muscle or applied on the cut end of the abducens nerve. Coordinates of these cell columns were used as target areas for the injection of small amounts of HRP (20-60 nl) and [3H]leucine (25-40 nl) and as search areas for retrogradely and anterogradely labeled internuclear neurons (INT) in in vivo and in vitro experiments. HRP injection in the medial rectus subdivision of the oculomotor nucleus (n = 6) resulted in retrograde labeling of cell bodies in the contralateral principal abducens nucleus. On the average about 16 cells per animal were found. Somatic diameters were about 13.5 +/- 2.8 microns (n = 32). The number and the size of these abducens internuclear neurons (AbINT) are smaller than those of lateral rectus motoneurons (n = 75; diameter: 19 +/- 3.2 microns). A crossed projection of AbINT to medial rectus motoneurons in the contralateral oculomotor nucleus is further supported by autoradiographic results. Following injection of [3H]leucine into the abducens nucleus, a high density of silver grains was visible within the contralateral oculomotor nucleus, mainly in the caudal part of the oculomotor nucleus, where medial rectus motoneurons are located. Injection of [3H]leucine in vivo (n = 4) and in vitro (n = 3) resulted in a similar high density of silver grains within the contralateral oculomotor nucleus, but the background level of silver grains was significantly higher after in vitro (264 +/- 38/2,500 microns2) than after in vivo injections (195 +/- 17/2,500 microns2). HRP injection in the principal abducens nucleus (n = 9) resulted in retrograde labeling of cell bodies in the medial rectus subdivisions of the bilateral oculomotor nuclei. Ipsilateral projections predominated, with about 10 (+/- 8) labeled cells over contralateral projections (about 3 +/- 2). Average diameters of these oculomotor internuclear neurons (OcINT) were again smaller (10.8 +/- 2 microns; n = 18) than those of medial rectus motoneurons (14.4 +/- 3 microns; n = 52). In addition, retrogradely labeled cells were consistently encountered in the bilateral vestibular nuclei, the cerebellar nuclei, the dorsal brainstem caudal to the abducens nuclei, and ipsilaterally in the pretectum. Most of the vestibular neurons were located in the rostral part of the vestibular nuclear complex. These neurons might constitute part of the three-neuronal arc of the vestibulo-ocular reflex in the frog. Labeled cells in the pretectum were restricted to the ipsilateral posterior thalamic nucleus (P).(ABSTRACT TRUNCATED AT 400 WORDS)     

The motor nuclei and sensory neurons of the IIIrd, IVth, and VIth cranial nerves in the monitor lizard, Varanus exanthematicus

The motor nuclei of the oculomotor, trochlear, and abducens nerves of the reptile Varanus exanthematicus and the neurons that subserve the sensory innervation of the extraocular muscles were identified and localized by retrograde and anterograde transport of horseradish peroxidase (HRP). The highly differentiated oculomotor nuclear complex, located dorsomedially in the tegmentum of the midbrain, consists of the accessory oculomotor nucleus and the dorsomedial, dorsolateral, intermediate, and ventral subnuclei. The accessory oculomotor nucleus projects ipsilaterally to the ciliary ganglion. The dorsomedial, dorsolateral, and intermediate subnuclei distribute their axons to the ipsilateral orbit, whereas the ventral subnucleus, which innervates the superior rectus muscle, has a bilateral, though predominantly contralateral projection. The trochlear nucleus, which rostrally overlaps the oculomotor nuclear complex, is for the greater part a comma-shaped cell group situated lateral, dorsal, and medial to the medial longitudinal fasciculus. Following HRP application to the trochlear nerve, almost all retrogradely labeled cells were found in the contralateral nucleus. The nuclear complex of the abducens nerve consists of the principal and accessory abducens nuclei, both of which project ipsilaterally. The principal abducens nucleus is located just beneath the fourth ventricle laterally adjacent to the medial longitudinal fasciculus and innervates the posterior rectus muscle. The accessory abducens nucleus has a ventrolateral position in the brainstem in close approximation to the ophthalmic fibers of the descending trigeminal tract. It innervates the retractor bulbi and bursalis muscles. The fibers arising in the accessory abducens muscles form a loop in or just beneath the principal abducens nucleus before they join the abducens nerve root. The afferent fibers conveying sensory information from the extraocular muscles course in the oculomotor nerve and have their perikarya in the ipsilateral trigeminal ganglion, almost exclusively in its ophthalmic portion.