Distribution of afferent fibers in the brainstem from end organs in the ear and lateral line in the European eel

Sensory nerve fibers from the lateral line system and labyrinth of Anguilla anguilla were labeled with horseradish peroxidase and traced to various targets in the ipsilateral brainstem. The three rami of the anterior lateral line nerve and the supratemporal ramus of the posterior lateral line nerve form overlapping terminal zones in the ventral portion of nucleus medialis. The posterior lateral line nerve on the body is represented exclusively in the dorsal half of the nucleus medialis. Eighth nerve fibers from the otolithic end organs in the inner ear send fibers into dorsal portions of three octavus nuclei: anterior, magnocellular, and descending, and saccular fibers lie most medial and utricular fibers most lateral. Fibers from vestibular organs, especially the semicircular canals and utricle, end densely in ventral portions of these nuclei and in the tangential nucleus. All labyrinthine sense organs send fibers into the region of a Mauthner-like neuron, and all except the saccule terminate in the reticular formation, tangential nucleus, and eminentia granularis of the cerebellum. Primary sensory input to the octavolateralis efferent nucleus comes only from the labyrinth, and fibers from the saccule alone penetrate the region of efferent neuronal somata. Fibers from labyrinthine end organs except the saccule project to the reticular formation where they may contact the dendrites of efferent somata. Fibers from the lateral line and the eighth nerve overlap most extensively at the rostral pole of the nucleus medialis and in the eminentia granularis of the cerebellum.     

Central projections of lagenar primary neurons in the chick

Perception of linear acceleration and head position is the function of the utricle and saccule in mammals. Nonmammalian vertebrates possess a third otolith endorgan, the macula lagena. Different functions have been ascribed to the lagena in arboreal birds, including hearing, equilibrium, homing behavior, and magnetoreception. However, no conclusive evidence on the function of the lagena in birds is currently available. The present study is aimed at providing a neuroanatomical substrate for the function of the lagena in the chicken as an example of terrestrial birds. The afferents from the lagena of chick embryos (E19) to the brainstem and cerebellum were investigated by the sensitive lipophilic tracer Neuro Vue Red in postfixed ears. The results revealed that all the main vestibular nuclei, including the tangential nucleus, received lagenar projections. No lagenar terminals were found in auditory centers, including the cochlear nuclei. In the cerebellum, the labeled terminals were found variably in all of the cerebellar nuclei. In the cerebellar cortex, the labeled fibers were found mostly in the uvula, with fewer afferents in the flocculus and paraflocculus. None was seen in the nodulus. The absence of lagenar afferent projections in auditory nuclei and the presence of a projection pattern in the vestibular nuclei and cerebellum similar to that of the utricle and saccule suggest that the primary role of the lagena in the chick lies in the processing of vestibular information related to linear acceleration and static head position. J. Comp. Neurol. 521:3524‐3540, 2013. © 2013 Wiley Periodicals, Inc.     

Central projections of octavolateralis nerves in the brain of a soniferous damselfish (Abudefduf abdominalis)

Sounds and hydrodynamic stimuli are important cues detected by the octavolateralis system in fishes. The central organization of auditory, mechanosensory, and vestibular projections is known for only a few phylogenetically diverse fishes, and less is known about projections in derived perciforms that use sounds for acoustic communication. We used neuronal labeling to provide a detailed analysis of octavolateralis endorgan projections in a soniferous perciform that does not have accessory morphological structures to enhance hearing. Octavolateralis nerves terminate ipsilaterally within seven medullary octaval nuclei: caudal (CON) and medial (MON) octavolateralis, anterior (AON), descending (DON), magnocellular (MgON), tangential (TON), and posterior (PON) octaval nuclei, and the eminentia granularis (EG). Anterior and posterior lateral line nerves project to the CON and MON, with dense projections to the EG. Semicircular canal nerves project primarily to ventral regions including the TON, ventral DON, intermediate DON (DONi), and MgON. Otolithic, semicircular canal, and anterior lateral line nerves all project to the MgON, which may serve a sensorimotor integration function. The DONi receives primarily segregated projections from all otolithic and semicircular canal nerves, whereas the ventral DON and TON receive principally utricular and semicircular canal afferents. The AON receives dense lateral and ventral projections from the saccule and utricle, and medial and dorsal projections from the lagena. These projection patterns are similar to those reported for non‐sonic perciforms, and indicate the absence of neuroanatomical modifications in first‐order octavolateralis nuclei in species that use acoustic communication. Thus patterns of central projections may be conserved among vocal and non‐vocal perciforms. J. Comp. Neurol. 512:628–650, 2009. © 2008 Wiley‐Liss, Inc.     

Afferent and efferent connections of the primary octaval nuclei in the clearnose skate, Raja eglanteria

Horseradish peroxidase techniques were employed to trace the central projections of afferents from the individual endorgans of the membranous labyrinth and to delineate the efferent projections from the primary octaval nuclei to the spinal cord and midbrain octavolateralis area in the clearnose skate, Raja eglanteria. First-order octaval afferents project ipsilaterally to five primary octaval nuclei, namely: magnocellular, descending, posterior, anterior, and periventricular. Octaval afferents also terminate in the reticular formation, nucleus intermedius (primary mechanoreceptive lateral-line nucleus), and vestibulolateral lobe of the cerebellum. Each primary octaval nucleus receives afferent input from each labyrinthine endorgan, with the possible exception of macula neglecta input to the magnacellular nucleus. Within the anterior, descending, and to a lesser extent posterior and magnocellular nuclei, this input is largely nonoverlapping. Semicircular canal cristae afferents terminate ventrally, saccular and lagenar afferents dorsally, utricular afferents laterally, and macular neglecta afferents course ventrally but terminate largely dorsally within these nuclei. In the vestibulolateral lobe of the cerebellum, cristae afferents project primarily to the pars medialis, whereas macular endorgan afferents terminate in the pars lateralis. Primary afferent input to the reticular formation is predominantly from the horizontal canal crista. The densest projections to nucleus intermedius are from the utriculus and sacculus. Vestibulospinal projections originate primarily from the magnocellular and descending nuclei. Second-order auditory neurons are most likely located in dorsomedial parts of the descending and anterior nuclei. Cells in these nuclei project directly to the auditory area of the midbrain octavolateralis complex, but projections to this area originate predominantly from nuclei C1 and C2, which are possible superior olivary homologues.     

Octavolateral projections and organization in the medulla of a teleost fish, the sleeper goby (Dormitator latifrons)

This study is the first to employ simultaneous labeling with different colored fluorescent dyes and confocal microscopy to investigate the central projections of the octavolateral nerves in any fish. Three-dimensional reconstructions of the hindbrain octavolateral nuclei were made and overlap of octavolateral projections was assessed in a teleost, the sleeper goby (Dormitator latifrons). The octavolateral nerves, which innervate the otolithic organs, semicircular canals, and lateral lines, project to seven hindbrain nuclei in diverse, complex patterns. The medulla is generally organized with auditory regions dorsal to vestibular regions. The intermediate subdivision of the descending octaval nucleus (DON) receives interdigitating projections from the otolithic organs, and the dorsomedial DON likely integrates multiple auditory inputs. Afferents from the three otolithic organs (the utricle, saccule, and lagena) project to the intermediate DON in approximately equal proportion, supporting physiological evidence that suggests auditory roles for all three otolithic organs in the sleeper goby. The anterior octaval nucleus receives partially segregated inputs from the octavolateral organs. The dorsal division of the magnocellular octaval nucleus (MgON) receives highly overlapping otolithic organ and semicircular canal input, and we propose that this region is a major octaval integration center. Regions in the ventral medulla (the tangential octaval nucleus, ventral DON, and ventral MgON) receive mainly utricular and semicircular canal inputs, suggesting vestibular roles. Each semicircular canal nerve projects to distinct regions of the hindbrain, with little overlap in most octaval nuclei. Efferent neurons receive bilateral input and project unilaterally to the octavolateral organs.     

Otolith endorgan input to the Mauthner neuron in the goldfish

The Mauthner (M-) cell of the goldfish, Carassius auratus, triggers the rapid escape response of the fish in response to various stimuli, including visual and auditory. The large size and accessibility of the M-cell make it an ideal model system for the study of synaptic transmission, membrane properties, and sensory-motor gating. Although physiological recordings have suggested that afferents from all three of the inner ear endorgans (the saccule, lagena, and utricle) synapse directly on the ipsilateral M-cell, the specific contacts and anatomical distributions of these inputs along the M-cell lateral dendrite remain unknown. We traced specific branches of the auditory (VIIIth) nerve from the three otolith organs of the fish inner ear to the M-cell. The goldfish sacculus gives rise to the vast majority of inputs that contact a large portion of the M-cell lateral dendrite, and these inputs vary greatly in size. In contrast to the ubiquitous distribution of saccular inputs, those from the lagena are segregated to distal regions of the M-cell and synapse on the distal dorsal branch of the lateral dendrite. Similarly, inputs from the utricle are also segregated to distal regions, synapsing on the ventral branch of the lateral dendrite. These results demonstrate that nerves from all three endorgans contact the M-cell, with input-specific segregation of synapses along the M-cell lateral dendrite.     

Quantitative analyses of postembryonic hair cell addition in the otolithic endorgans of the inner ear of the european hake,merluccius merluccius (gadiformes,teleostei)

Bony fishes add sensory hair cells to the saccule and lagena of the ear for at least several years after hatching. However, it is not known whether hair cell proliferation occurs for the whole lifetime of an animal, whether proliferation occurs in all endorgans of the ear, or whether the rate of proliferation is the same in all of the endorgans. To obtain answers to these questions, the extent of postembryonic hair-cell proliferation was determined in the saccule, lagena, and utricle of the ear in the European hake, Merluccius merlaccius, for fish ranging from 7 to 75 cm in total length (6 months to 9 years of age). Results demonstrated that hair-cell addition continued throughout this period in all three otic endorgans, although endorgan size was proportionally greatest in smaller animals. Of the three endorgans, cell addition was greatest in the saccule. Moreover, far more cells were added to the caudal end of the saccule than to the rostral end. Each saccule of the largest hake had over 900,000 hair cells. It is estimated that each saccule adds approximately 110,000 new hair cells each year (or 302 cells/day) over the life span of the fish studied. A significant number of small ciliary bundles, thought to represent newly proliferated hair cells, was found throughout each endorgan, and the number of such bundles declined as the rate of hair cell proliferation decreased. The results demonstrate that extensive proliferation occurs in all three otolithic endorgans of the ears in a fish and that such proliferation continues for virtually the whole life of the animal. The functional significance of this addition is not known. © 1994 Wiley-Liss, Inc.     

The inner ear of the lungfish Protopterus

The sensory end organs of the inner ear of the lungfish, Protopterus, were examined using scanning and transmission electron microscopy. The utricle has a structure and hair cell orientation pattern that are typical for vertebrates, although the hair cells are unusually large. There are the typical three semicircular canals extending from the utricle, with the typical hair cell orientations, but the lateral canal sensory crista looks like the "hemicrista" of some amphibians and amniotes, lacking a saddle-shaped flare on one wall of the ampulla. Unlike most vertebrates that have the saccule and lagena as two separate pouches ventral to the utricle, the lungfish has a single large ventral pouch that contains a single large pasty otoconial mass. This mass covers two hair cell patches, each like a striola with prominent hair cell ciliary bundles, that are presumed to represent saccular and lagenar maculae. However, these two major sensory patches are not completely separate maculae because they lie within a less densely populated field of smaller hair cells, which forms an extrastriolar region that surrounds and fills the region between the two striolae of higher hair cell density. The more caudal lagenar striola is a vertically elongated stripe with hair cell orientation vectors facing antiparallel on either side of a midline drawn vertically along the macula, resembling the macula lagena of some bony fishes but not of tetrapods. The more rostral saccular striola is a curving band with hair cell orientation vectors facing away from its midline, but because this macula curves in three dimensions, the vectors at the rostral end of this striola are oriented mediolaterally, whereas the vectors on the caudal half of this striola are oriented dorsoventrally. The presence of a macula neglecta was confirmed near the posterior canal as a tiny single patch of a few dozen hair cells with all the cell orientations directed caudally. The ciliary bundles on the cells in the striolar-like regions of all of three otolithic organs average over 80 cilia, a number far greater than for any other fish studied to date. The features of the single sacculolagenar pouch with separate striolar-like regions, the cellular orientation in the otolith organs, and the large cells and ciliary bundles in Protopterus also were observed in specimens of the other extant lungfish genera, Lepidosiren and Neoceratodus.     

Central projections of the lateral line and eighth nerves in the bowfin, Amia calva

The first-order connections of the anterior and posterior lateral line nerves and of the eighth nerve were determined in the bowfin, Amia calva, using experimental degeneration and anterograde HRP transport techniques. The termination sites of these nerves define a dorsal lateralis cell column and a ventral octavus cell column. The anterior and posterior lateralis nerves distribute ipsilaterally to two medullary nuclei-nucleus medialis and nucleus caudalis. Nucleus medialis comprises the rostral two-thirds of the lateralis column and contains large, Purkinje-like cells dorsally and polygonal, granule, and fusiform cells ventrally. Nucleus caudalis is located posterior to nucleus medialis and consists of small, granule cells. Anterior lateralis fibers terminate ventrally to ventromedially in both nucleus medialis and nucleus caudalis. Posterior lateralis fibers terminate dorsally to dorsolaterally within these two nuclei. A sparse anterior lateralis input may also be present on the dendrites of one of the nuclei within the octavus cell column, nucleus magnocellularis. In contrast, the anterior and posterior rami of the eighth nerve each terminate within four medullary nuclei which comprise the octavus cell column: the anterior, magnocellular, descending, and posterior octavus nuclei. An eighth nerve projection to the medial reticular formation is also present. Some fibers of the lateralis and eighth nerves terminate within the ipsilateral eminentia granularis of the cerebellum. Lateralis fibers distribute to approximately the lateral half of this structure with posterior lateral line fibers terminating laterally and anterior lateral line fibers terminating medially. Eighth nerve fibers distribute to the medial half of the eminentia granularis.     

Peripheral configuration and central projections of the lateral line system in Astronotus ocellatus (Cichlidae): a nonelectroreceptive teleost

The lateral line system of Astronotus ocellatus comprises one trunk canal, one tail canal, and three head canals. The sensory receptors on the head are innervated by rami of the dorsal anterior, ventral anterior, and posterior lateral line nerves, and those along the trunk and tail by rami of the posterior lateral line nerve. The peripheral configuration of lateral line canals and nerves was examined in whole mount preparations, the central connections of restricted groups of endorgans studied using HRP and degeneration methods, and the neuronal morphology and cytoarchitecture of the lateralis region investigated with Nissl, silver, and Golgi methods. Neurons of the lateralis cell column are diffusely arrayed and of variable morphology. They are oriented primarily in the transverse plane and, with the exception of a dorsal lamina of large multipolar cells, are not organized into zones. Lateralis fibers bifurcate on entering the brainstem, course in lateral tracts, and give off medially directed collaterals to terminate in the ipsilateral nucleus medialis and nucleus caudalis. In addition, fibers terminate in the eminentia granularis of the cerebellum, but only fibers supplying endorgans in the head canals penetrate the granule cell zone of the cerebellar corpus. Fibers supplying sense organs in adjacent canals overlap in their central endings, whereas fibers of distantly separated receptors do not overlap. The rami supplying trunk and tail canal organs do not project as far rostrally in the central neuropil as do the other rami. Endings of posterior lateral line fibers lie dorsal to those of the anterior lateral line nerves, and some lateralis fibers terminate within the confines of the magnocellular, descending, and posterior nuclei of the octavus column. Although there is spatial order to the lateralis projections, there is no clear somatotopic organization in the lateralis region.     

Location of dye-coupled second order and of efferent vestibular neurons labeled from individual semicircular canal or otolith organs in the frog

Vestibular nerve branches innervating the sensory epithelia of the three semicircular canals or of the three otolith organs of frogs were selectively labeled in-vitro with biocytin. Labeled afferent fibers from the semicircular canals, utricle, and lagena were encountered in each of the four vestibular nuclei and their projections overlapped considerably. Saccular afferent fibers projected to the dorsal (acoustic) nuclei and smaller projections to the vestibular nuclei were regionally restricted. Per semicircular canal or otolith organ about equal numbers (11-14) of medium sized vestibular neurons (between 7.5 and 17 microm in diameter) were dye-coupled to afferent fibers. Most of these dye-coupled vestibular neurons were located in the lateral and descending vestibular nuclei between the VIIIth and IXth nerves. The superior vestibular nucleus was relatively free of dye-coupled vestibular neurons. The location of this subpopulation of central vestibular neurons supports the notion that these neurons are part of a particular vestibulospinal pathway. In addition, from each of the canal and/or otolith organs about 3-4 efferent vestibular neurons were labeled retrogradely. These neurons (between 15 and 26 microm in diameter) were located ventral to the vestibular nuclear complex. The branching of efferent vestibular neurons was shown by the presence of neurons that were double labeled by two different fluorescent dyes applied in the same experiment to the anterior and posterior ramus of the same VIIIth nerve, respectively. The branching of these efferent neuron axons explained the presence of collaterals and terminals in the sensory epithelia of a number of untreated ipsilateral endorgans.     

Differential central projections of vestibular afferents in pigeons

The question of whether a differential distribution of vestibular afferent information to central nuclear neurons is present in pigeons was studied using neural tracer compounds. Discrete tracing of afferent fibers innervating the individual semicircular canal and otolith organs was produced by sectioning individual branches of the vestibular nerve that innervate the different receptor organs and applying crystals of horseradish peroxidase, or a horseradish peroxidase/cholera toxin mixture, or a biocytin compound for neuronal uptake and transport. Afferent fibers and their terminal distributions within the brainstem and cerebellum were visualized subsequently. Discrete areas in the pigeon central nervous system that receive primary vestibular input include the superior, dorsal lateral, ventral lateral, medial, descending, and tangential vestibular nuclei; the A and B groups; the intermediate, medial, and lateral cerebellar nuclei; and the nodulus, the uvula, and the paraflocculus. Generally, the vertical canal afferents projected heavily to medial regions in the superior and descending vestibular nuclei as well as the A group. Vertical canal projections to the medial and lateral vestibular nuclei were observed but were less prominent. Horizontal canal projections to the superior and descending vestibular nuclei were much more centrally located than those of the vertical canals. A more substantial projection to the medial and lateral vestibular nuclei was seen with horizontal canal afferents compared to vertical canal fibers. Afferents innervating the utricle and saccule terminated generally in the lateral regions of all vestibular nuclei in areas that were separate from the projections of the semicircular canals. In addition, utricular fibers projected to regions in the vestibular nuclei that overlapped with the horizontal semicircular canal terminal fields, whereas saccular afferents projected to regions that received vertical canal fiber terminations. Lagenar afferents projected throughout the cochlear nuclei, to the dorsolateral regions of the cerebellar nuclei, and to lateral regions of the superior, lateral, medial, and descending vestibular nuclei. © 1996 Wiley-Liss, Inc.     

Semicircular canal, saccular and utricular function in patients with bilateral vestibulopathy: analysis based on etiology

The diagnosis of bilateral vestibulopathy (BV) is typically established based on bilateral semicircular canal dysfunction. The degree to which both otolith organs—the saccule and utricle—are also impaired in BV is not well-established, particularly with respect to the etiology and severity of BV. The aim of this study was to evaluate semicircular canal, saccular and utricular function in patients with BV due to aminoglycoside ototoxicity and bilateral Menière’s disease, and with different severities of BV. Caloric and head impulse testing were used as measures of canal function. Cervical vestibular-evoked myogenic potentials (cVEMP) and ocular VEMPs (oVEMP) were used as measures of saccular and utricular function, respectively. We enrolled 34 patients with BV and 55 controls in a prospective case–control study. Patients with BV were less likely to have saccular (61 %) or utricular (64 %) dysfunction relative to canal dysfunction (100 %). Utricular function differed significantly between patients by etiologic group: the poorest function was found in patients with BV due to aminoglycoside toxicity, and the best function in Menière’s disease patients. Canal and saccular function did not vary according to etiology. Further, utricular but not saccular function was significantly correlated with canal function. Saccular and utricular function had the strongest association with Dizziness Handicap Inventory scores relative to canal function. These data suggest that when a patient with BV is identified in a clinical context, oVEMP testing is the most sensitive test in distinguishing between aminoglycoside toxicity and bilateral Menière’s disease. Both cVEMP and oVEMP testing may be considered to evaluate the functional impact on the patient.     

Organization of dye-coupled cerebellar granule cells labeled from afferent vestibular and dorsal root fibers in the frog Rana esculenta

Application of neurobiotin to the nerves of individual labyrinthine organs and dorsal root fibers of limb-innervating segments of the frog resulted in labeling of granule cells in the cerebellum showing a significant overlap with a partial segregation in the related areas of termination. In different parts of the cerebellum, various combinations of different canal and otolith organ-related granule cells have been discerned. The difference in the extension of territories of vertical canals vs. horizontal canals may reflect their different involvement in the vestibuloocular and vestibulospinal reflex. Dye-coupled cells related to the lagenar and saccular neurons were localized in more rostral parts of the cerebellum, whereas cells of the utricle were represented only in its caudal half. This separation is supportive of the dual function of the lagena and the saccule. The territories of granule cells related to the cervical and lumbar segments of the spinal cord were almost completely separated along the rostrocaudal axis of cerebellum, whereas their territories were almost entirely overlapping in the mediolateral and ventrodorsal directions. The partial overlap of labyrinthine organ-related and dorsal root fiber-related granule cells are suggestive of a convergence of sensory modalities involved in the sense of balance. We propose that the afferent input of vestibular and proprioceptive fibers mediated by gap junctions to the cerebellar granule cells subserve one of the possible morphological correlates of a very rapid modification of the motor activity in the vestibulocerebellospinal neuronal circuit.     

Peripheral distribution and central projections of the lateral-line nerves in goldfish, Carassius auratus

The lateral-line system of the goldfish, Carassius auratus, is described from the receptor distribution on the body surface to the termination of the primary afferents within the octavolateralis column. Goldfish have a continuous lateral-line canal system which consists of supraorbital, infraorbital, operculomandibular and supratemporal commissural canals on the head and a trunk canal extending the length of the trunk. There is a single neuromast between each 2 canal pores. The total number of canal neuromasts on the head ranges from 74 to 100. On the body the total number of canal neuromasts ranges from 52 to 60. Associated with each canal are populations of superficial neuromasts. There are approximately 1,000 superficial neuromasts on the head which are arranged in clusters or rows oriented perpendicular to the long axis of the associated canal. On the body there are 3-9 superficial neuromasts on each scale. Further, there is a total of 1,000 superficial neuromasts arranged in horizontal lines between the fin rays of the caudal fin. The neuromasts are innervated by branches of four lateral-line nerves. The dorsal anterior lateral-line nerve innervates the supraorbital and infraorbital canals and associated superficial neuromasts. The ventral anterior lateral-line nerve innervates the operculomandibular canal and associated superficial neuromasts, as well as superficial neuromasts on the operculum. The middle lateral-line nerve innervates the postotic canal and a cluster of associated superficial neuromasts. The posterior lateral-line nerve innervates the supratemporal commissural canal and associated superficial neuromasts, as well as the canal and superficial neuromasts on the trunk and caudal fin. Application of horseradish peroxidase to the individual branches of the lateral-line nerves shows that the posterior lateral-line nerve projects to the dorsal portion of the medial and caudal nuclei, whereas the anterior lateral-line branches project to the ventral portion. Lateral-line projections to the magnocellular and descending nuclei of the octaval column were also observed. The projections to the eminentia granularis are also segregated such that the anterior lateral-line nerves project to the anterolateral portion and the posterior lateral-line nerve to the caudal portion. Thus, there appears to be a topographic organization of the lateral-line projections to the lateralis column and eminentia granularis. However, this topography does not appear to be precise as the projections of the branches of the anterior lateral-line nerves overlap extensively, as do the branches of the posterior lateral-line nerves.     

Structure and innervation of the inner ear of the bowfin,Amia calva

The hair cell orientation patterns and the number of innervating nerve cells was determined for the ear of the holostean fish Amia calva. Scanning electron microscopy was used to analyze the sensory cells of the three otolith organs: the saccule, the lagena, and the utricle. The utricular hair cell orientation pattern is similar to that found in most vertebrates, while the lagenar hair cell pattern is similar to that in ostariophysine fishes. The saccule has both horizontally and vertically oriented hair cells, although in the living animal many of the horizontally oriented cells are really oriented lateromedially. The transition region between cells oriented horizontally and vertically suggests that the horizontal cells are ontogenetically derived from vertically oriented cells. The lagenar macula has the largest sensory region, with over 216,000 cells in its main sensory part. The utricular macula contains over 56,400 cells in the main sensory region, and the saccular macula over 8,600 cells. The otic nerve divides into anterior and posterior rami, and ganglion cell counts totaled 2,021 in the anterior ramus and 1,619 in the posterior ramus. There are three populations of ganglion cells in each ramus, on the basis of differences in cell diameter. By a rough estimate, the ratio of sensory to ganglion cells is at least 90:1. By comparing SEM data from Amia, other nonteleosts, and various teleost species, we suggest that the ontogenetic derivations of the horizontal cells in the teleosts and nonteleosts are substantially different and apparently unrelated to each other and that the most primitive ray-finned fishes had only vertically oriented sensory cells while the horizontally oriented cells subsequently evolved at least twice, once in Amia and other nonteleosts, and once in the ancestors of the teleosts.     

Primary afferent projections to the central octavus nuclei in the elasmobranch, Rhinobatos sp., as demonstrated by nerve degeneration

Nerve degeneration studies were performed to determine the central projections of the octavus nerve in the guitarfish. Rhinobatos sp. Most first-order octavus nerve afferents are confined to the ventral longitudinal column of the medulla, which consists of four octavus nuclei: the nucleus octavus anterior (NOA), nucleus octavus magnocellularis (NOM), nucleus octavus descendens (NOD); and nucleus octavus posterior (NOP). Central projections of the octavus nerve outside of the ventral octavus nuclear column are to the vestibulolateral lobe of the cerebellum, the large multipolar cells that lie deep to the cerebellar crest of nucleus intermedius, the reticular formation, and the nucleus octavolateralis efferens. In addition, the cytoarchitectural features of the four octavus nuclei have been analyzed: the NOA consists of large spindle-shaped neurons and medium-sized multipolar neurons; the NOM is a small nucleus consisting of very large multipolar neurons; the NOD contains several types of neurons including large spindle-shaped neurons, large multipolar neurons, and polymorphic small and medium-sized neurons; and the NOP contains small spherical and small multipolar neurons evenly distributed throughout the nucleus. On the basis of the morphology of the neurons, the NOA and NOD appear to have subdivisions suggesting a more complex organization of the octavus nuclear column than previously recognized.     

Otolith End Organ Projections to Auditory Neurons in the Descending Octaval Nucleus of the Goldfish, Carassius auratus : A Confocal Analysis

The distribution of axons from the saccule, lagena, and utricle to descending octaval nucleus neurons that project to the auditory midbrain in the goldfish is reported. We have divided these auditory projection neurons, located in the dorsal portion of the descending octaval nucleus (dDO), into two groups, medial and lateral, each of which contains several neuronal populations based on morphology and location. At most levels of the dDO, there are three medial and three lateral populations; the rostral dDO contains an additional lateral population. The saccule provides input to each of the seven medial and lateral populations but appears to be the exclusive/nearly exclusive source of primary input to the most dorsal cell group of the medial population. Along with the saccule, the lagena and utricle each supply the remaining six medial and lateral populations. Neurons in each of these populations receive input from more than one end organ. One medial and one lateral population include neurons that receive remarkably large contacts from utricular afferents. Overall, the results reveal a more substantial input from the lagena and utricle to the main first-order auditory nucleus in the goldfish than was previously recognized, suggest this nucleus is composed of functionally distinct populations, and relate to functional and evolutionary issues about hearing in early vertebrates.     

Peripheral and central aspects of the acoustic and lateral line system of a bottom dwelling catfish, Ancistrus sp.

The topographical relationship between the swim bladder, the inner ear, and the otic lateral line was studied in the bottom dwelling catfish, Ancistrus sp. In addition, afferent and efferent subcomponents of the eighth and lateral line nerves were labelled with horseradish peroxidase (HRP) or with differently fluorescing dextran amines. The swim bladder of Ancistrus consists of two separate, transversely oriented parts of each of which is connected to the sinus impar of the inner ears via two Weberian ossicles and the perilymphatic sac. The osseous capsula of the ear has two foramina other than the nerve foramina. One is for the sinus impar. The other foramen, which also separates two fluid-filled spaces, exits where the horizontal canal of the ear contacts the otic lateral line. Both the otic and the postotic lateral line canal run deep below the epidermis. Each canal contains a neuromast that is innervated by the middle lateral line nerve. Further caudally, the otic lateral line canal gives rise to the postotic and finally to the truck canal whose nonossified anterior part travels through an ossified chamber that surrounds the swim bladder. Thus the anterior part of each trunk lateral line canal is in contact with a bipartite sound pressure receiver, the swim bladder. Anterior and posterior lateral line afferents terminate ipsilaterally throughout the neuropil of the electroreceptive lateral line nucleus and the mechanoreceptive nuclei medialis and caudalis of the medulla. Middle lateral line afferents terminate between the projection sites of anterior and posterior lateral line afferents. Some primary mechanosensory anterior lateral line nerve fibers continue into the ipsilateral eminentia granularis and the valvula cerebelli. In the electroreceptive lateral line projection, anterior lateral line fibers terminate more medially and posterior fibers more laterally. This somatotopy is not as clear-cut in the mechanosensory lateral line. Afferents of the sacculus and the lagena terminate predominantly in the saccular nucleus. Afferents of the utriculus, the horizontal canal, and the anterior vertical canal terminate in the magnocellular vestibular nucleus and in the medial octavolateral nucleus. The projection sites of the anterior part and the posterior part of the eighth nerve show little overlap. Eighth nerve projections to the valvula cerebelli are less prominent than the projections from the lateral line. Eighth nerve and lateral line nerve efferents arise from a common nucleus, the octavolateralis efferent nucleus. Axons of efferent cells may divide to supply two or more branches of the eighth nerve and some axons supply both lateral line and eighth nerve endorgans.(ABSTRACT TRUNCATED AT 400 WORDS)     

Analysis of magnetic elements in otoliths of the macula lagena in homing pigeons with inductively coupled plasma mass spectrometry

Objective

The macula lagena in birds is located at the apical end of the cochlea and contains many tiny otoliths. The macula lagena is innervated and has neural projections to the brainstem, but its physiological function is still unclear. It remains disputable that it is because otoliths in the lagena are rich in elements Fe and Zn that birds can obtain geomagnetic information for homing. To clarify this issue, we carried out a study to determine whether or not otoliths in the lagena of homing pigeons are richer in magnetic elements than those in the saccule and the utricle.

Methods

The contents of ferromagnetic elements (Fe, Co, Ni) and other metal elements in lagenal otoliths of adult homing pigeons were precisely analyzed with inductively coupled plasma mass spectrometry (ICP-MS) of high sensitivity, and then they were compared with those in saccular and utricular otoliths (all the contents were normalized to Ca).

Results

In adult homing pigeons, the contents of ferromagnetic elements (Fe, Co, Ni) in lagenal otoliths were less than 0.7% (normalized to Ca element) and were the same order in magnitude as those in saccular and utricular otoliths. The content of Fe in lagenal otoliths was not significantly different from that in utricular otoliths and was even lower than that in saccular otoliths. The content of Co in lagenal otoliths was lower than that in saccular otoliths and higher than that in utricular otoliths. The content of Ni in lagenal otoliths was not significantly different from that in saccular otoliths and was higher than that in utricular otoliths. The contents of other metal elements Na, Mg, K, Al, Mn and Pb in lagenal otoliths were not significantly different from those in utricular and saccular otoliths. The contents of metal elements Zn, Ba and Cu in lagenal otoliths were lower than those in saccular otoliths.

Conclusion

The contents of magnetic elements in lagenal otoliths of homing pigeons are not much higher than those in utricular and saccular otoliths, which does not support the hypothesis that birds depend on high contents of Fe and Zn in lagenal otoliths for sensation of geomagnetic information. Similarities in morphology, element ingredient and element content between lagenal otoliths and utricular otoliths suggest that the two types of otolithic organs may play similar roles in sensing gravitational and acceleration signals.