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We have studied the early development of the vestibular apparatus and its central connections in the tammar wallaby (Macropus eugenii) in order to determine whether the vestibular system anatomy is sufficiently mature at birth to assist in climbing to the pouch. Structural development was studied with the aid of hematoxylin and eosin stained sections and immunoreactivity for GAP-43, whereas the development of vestibular system connections was examined by carbocyanine dye tracing. At the time of birth, the otocyst has distinct utricle, saccule and semicircular canals with immature sensory regions receiving innervation by GAP-43 immunoreactive fibers. Vestibular nerve fibers can be traced into the brainstem to the developing vestibular nuclei, which are not yet cytoarchitectonically distinct. The vestibular nuclei do not contribute direct projections to the lower cervical spinal cord at birth; most bulbospinal projections in the newborn appear to be derived bilaterally from the gigantocellular, lateral paragigantocellular reticular and ventral medullary nuclei. A substantial bilateral projection to the vestibular ganglion and apparatus from the region of the gigantocellular and lateral paragigantocellular nuclei was seen at birth, but not in subsequent ages. This is similar to a projection seen in newborn Ameridelphians. By postnatal day (P) 5, the vestibular apparatus had extensive projections to all vestibular nuclei and neurons projecting in the lateral vestibulospinal tract could be identified in the lateral vestibular nucleus. Cytoarchitectonic differentiation of the vestibular nuclei proceeded over the next 3 to 4 weeks with the emergence of discrete parvicellular and magnocellular components of the medial vestibular nucleus by P19. GAP-43 immunoreactivity stayed high in the lateral vestibulospinal tract for several months after birth, suggesting that the development of this tract followed a prolonged timecourse. Our findings indicate that central and peripheral connections of the vestibular ganglion are present at birth, but that there is no direct projection from the vestibular nuclei to the cervical spinal cord until P5. Nevertheless, the possibility remains that an indirect projection between the vestibular nuclei and the medial reticular formation is present at birth and mediates control of the climb., ((c) 2008 S. Karger AG, Basel)

The vestibular apparatus provides information about the position and movements of the head. Craniocervical muscles position the head with respect to the upper part of the neck. Motoneurons innervating these muscles are located in the supraspinal nucleus and ventral horn of the rostral cervical cord. Premotor neurons of craniocervical muscles have been found in the medial two-thirds of the medullary reticular formation: the ventromedial part of the parvocellular reticular formation and the gigantocellular reticular formation. In the present study, projections from vestibular nuclei upon craniocervical premotor neurons were investigated using anterograde and retrograde tracers. Vestibulospinal fibers run bilaterally in the medial vestibulospinal tract and ipsilaterally in the lateral vestibulospinal tract. Vestibuloreticular projections are mainly ipsilateral, and originate from the n. vestibularis lateralis pars ventralis and pars dorsalis, and from the n. vestibularis descendens. Terminal labeling is found in the border zone between the parvocellular and gigantocellular reticular formation. These projections show that in addition to direct bilateral vestibulo-craniocervical projections an indirect vestibular pathway to craniocervical motor nuclei exists. The direct pathway probably is the neural substrate for the vestibulocollic reflex, whereas the vestibular projection upon the reticular formation might influence head orientation during various kinds of activities, such as pecking, preening and so on., (Copyright 2002 S. Karger AG, Basel)

The distribution of immunoreactive gonadotropin-releasing hormone (GnRHir) in relation to endocrine and behavioral pathways is not well established for lizards. To more completely understand the GnRHir distribution and its possible function in a lizard, we investigated the brain of Anolis carolinensis, a species whose visual courtship displays, mating postures and gonadal cycles are well known. Using antisera that recognize multiple GnRH forms, we observed no GnRHir cells or fibers in the forebrain. In the midbrain, however, GnRHir cells occurred along the medial border of the medial longitudinal fasciculus. These cells appeared to project GnRHir fibers to nuclei of cranial nerve III and IV. In the hindbrain, positive fibers were observed in the area of the vestibular nuclei and dorsal funiculus. These hindbrain fibers were followed to their terminals in the cerebellum. The GnRHir midbrain distribution suggests an association of GnRH with eye movements, whereas the hindbrain distribution suggests a GnRH-vestibular association. The lack of GnRHir in the forebrain of Anolis could reflect the minimal role of the vomeronasal system in mediating reproduction in this species. Thus, our study cannot verify hypothalamic GnRH control of the pituitary in A. carolinensis, but it does indicate GnRH control of oculomotor and vestibular nuclei, which might play a role in Anolis reproductive behaviors.

We have shown previously that GAP-43, a growth associated protein characteristically present in growing and regenerating axons, is relatively abundant in the spinal cord of adult opossums. In the present study, we combined the orthograde transport of the fluorescent marker Fluoro-Ruby with immunofluorescence for GAP-43 to determine if any of it is present within descending spinal axons. When Fluoro-Ruby was injected into the red nucleus and midbrain tegmentum, the medial pontine or medullary reticular formation, the medullary raphe or the lateral vestibular nucleus, axons were labeled in the expected areas of the spinal cord, but in most cases none showed evidence for GAP-43. In two of the four cases with rubral injections, however, a few labeled axons within the rubrospinal tract showed GAP-43 immunofluorescence, and in one case with an injection of the gigantocellular reticular nucleus and adjacent raphe, labeled axons within lamina IX immunostained for the protein. Since serotoninergic neurons are present within the gigantocellular reticular nucleus and adjacent raphe, and axons of the same phenotype are abundant within lamina IX, we asked whether serotoninergic axons contain GAP-43. When sections of the spinal cord were immunostained for both serotonin and GAP-43, many axons within lamina IX showed evidence for both substances. Such axons appeared to contact presumptive motoneurons. In cases with Fluoro-Ruby injections of the forelimb motor cortex, labeled axons were present within the pyramidal tract, and some of them showed evidence for GAP-43.

Horseradish peroxidase and Fast Blue were injected into the oculomotor and trochlear nuclei of rabbits so as to study the distribution of vestibular neurons that project to these nuclei. After the oculomotor nucleus was injected, labelled neurons were found in the superior, medial, and descending vestibular nuclei as well as in cell group Y. In the superior nucleus, most of the neurons (510 +/- 46) were ipsilateral to the injection, although contralaterally labelled neurons were also observed (104 +/- 19) more peripherally. In cell group Y, 186 +/- 24 contralaterally labelled neurons were observed, whereas hardly any (8 +/- 3) were found on the ipsilateral side. The largest group of labelled neurons (811 +/- 65) constituted a neuronal band located contralaterally in the medial nucleus and rostral part of the descending nucleus. This band rostromedially continued with the caudal part of the group of internuclear neurons of the abducens nucleus. Only 190 +/- 31 neurons were labelled in the medial and descending nucleus ipsilateral to the injected oculomotor nucleus. After injection of the trochlear nucleus, labelled neurons were found in the ipsilateral superior nucleus and contralateral medial and descending nuclei: a few labelled cells were also observed in the ipsilateral medial and descending nuclei as well as in the contralateral cell group Y.

In this study we have used retrogradely transported horseradish peroxidase (HRP) to investigate whether the nucleus prepositus hypoglossi of a lateral-eyed mammal projects to the oculomotor and trochlear nuclei. After the injection of HRP in the oculomotor nucleus of rabbits, labelled neurons were found bilaterally in the nucleus prepositus hypoglossi, though they were more numerous on the ipsilateral side. The majority of these neurons were labelled in the rostral part of the propositus nucleus and were dispersed predominantly in the lateral and ventral zone of the nucleus. Neurons were also labelled in the prepositus nucleus after injection of HRP in the trochlear nucleus. These data were compared with those for frontal-eyed mammals and birds and suggest that the said projections are less well developed in species that possess panoramic vision and a lesser degree of binocular yoking.

Correlations between progression indices of five categories of nuclei and brain centers indicate that Megachiroptera rely primarily on vision and olfaction while Microchiroptera rely primarily on audition. Among Microchiroptera, some rely more heavily on olfaction than on vision as a secondary sense, and vice-versa. There may have been compensatory effects, in the course of evolution, regarding the choice of the primary and secondary information channels.

Two specialized features are described in the auditory system of Acrobates pygmaeus, a small gliding marsupial. Firstly, the ear canal includes a transverse disk of bone that partly occludes the canal near the eardrum. The resultant narrow-necked chamber above the eardrum appears to attenuate sound across a broad frequency range, except at 27-29 kHz at which a net gain of sound pressure occurs. Secondly, the lateral medulla is hypertrophied at the level of the cochlear nucleus, forming a massive lateral lobe comprised of multipolar cells and granule cells. This lobe has connections with the auditory nerve and the cerebellum. Speculations are advanced about the functions of these structures in gliding behaviour and predator avoidance.

Horseradish peroxidase was injected at various levels of the spinal cord of the hen and the cells in the brain stem, labeled due to retrograde transport of the tracer enzyme, were mapped with particular reference to the vestibular nuclear complex. The Deiters' nuclei project ipsilaterally to the spinal cord. The projection from the nucleus Deiters ventralis reaches down to the lumbosacral spinal cord. The spinal projection from the nucleus Deiters dorsalis is somatotopically organized. Cells localized rostrally within the nucleus project to upper parts of the spinal cord cells localized caudally project to the lumbosacral spinal cord. The medial and the descending nucleus contain labeled cells in two compartments of the nuclei. In each of them a rostral one projects to the upper cervical cord, a caudal one projects to the thoracical spinal cord. In addition, labeled cells are observed in the reticular formation, mainly in the pons and the medulla oblongata, in the red nucleus, in the raphe nuclei and in the periaqueductal grey. Very few labeled cells are observed in the locus ceruleus, the nucleus intercalatus and in the nucleus of the solitary tract. In some cases, the number of labeled cells in the various nuclei in the brain stem projecting to the spinal cord was counted to get an impression of the quantitative importance of the vestibulospinal projection relative to afferents to the spinal cord from the other nuclei. The findings are discussed in the light of what is known of the vestibulospinal projection in mammals.

Electrophysiological and light microscopical studies were made on the dogfish vestibulocerebellum. The cell types, estimates of their numbers, and the organization of this part of the cerebellum are described and compared with the other lobes of the elasmobranch cerebellum. Electrophysiological study of the inputs to this region from the VIIIth nerves indicates that primary afferent fibres from the VIIIth nerve project to the ipsilateral granule cell layer.

Electrophysiological and light microscopical studies were made on the vestibular area of the dogfish hindbrain. Three vestibular nuclei were distinguished: the superior nucleus (VES), the magnocellularis nucleus (VEM), and the ventral nucleus (VEV). The distribution of field potentials evoked in the hindbrain by stimulation of nerve VIII confirms the location and extent of the vestibular nuclei. It also raises the possibility of a direct contralateral projection of vestibular nerve fibres. Unit studies confirm the interpretation of the field potentials and provide evidence of mono- and polysynaptic activation of vestibular nuclear neurons by vestibular afferents.

Microelectrode recording in the thalamus of pigeons subjected to tilt and sinusoidal rotational stimuli around the vertical, longitudinal and transversal axes revealed vestibularly driven units in two thalamic nuclei, the nucleus posteroventralis and the nucleus principalis precommissuralis. Many of these units responded in a complex manner suggesting that inputs from contralateral and ipsilateral cupulae and maculae converged on them. A few units received additional visual or proprioceptive information. The homology relationship with a mammalian vestibular thalamic nucleus is discussed briefly.

The topographic organization of the corticonuclear and corticovestibular fibers from the pyramis and copula pyramidis in the albino rat was investigated by the autoradiographic orthograde tracing technique. Cortical efferent fibers from the medial part of the pyramis project to the caudoventral part of the caudomedial and middle subdivisions of the medial cerebellar nucleus, whereas those arising from the lateral part of the pyramis project to the caudomedial part of the posterior interpositus nucleus and also to the dorsal part of the lateral vestibular nucleus via the juxtarestiform body. Fibers from the medial part of the copula pyramidis project to the medial part of the anterior interpositus nucleus. On the other hand, fibers from the medial portion of the lateral part of the copula pyramidis project to the posterior interpositus nucleus, while those from the lateral portion, including the lateral limit of the copula pyramidis, project to the rostroventral part of the lateral cerebellar nucleus. These results indicate that the cortical efferent projections from the pyramis and the copula pyramidis are clearly oriented mediolaterally in the albino rat.

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.