Morphology and physiology of cells in slice preparations of the dorsal cochlear nucleus of mice

Horseradish peroxidase (HRP) was injected into cells from which intracellular recordings were made in slices of the dorsal cochlear nucleus (DCN) in order to correlate physiology with morphology. In general, the morphology of cells labeled intracellularly with HRP corresponded to those made with Golgi impregnations in mice and other mammals. The following cells were labeled: one granule cell, four cartwheel cells, eight fusiform cells, two other cells in the fusiform cell layer, and two tuberculoventral association cells in the deep layers of the DCN. The axon of the granule cell runs parallel to isofrequency laminae with collaterals branching perpendicularly and running along the tonotopic axis. The cartwheel cells have dendrites in the molecular layer that are densely covered with spines. The axon of one cell terminates just dorsally to the cell body. Fusiform cells have the characteristic spiny, apical and smooth, basal dendrites. The basal dendrites are conspicuously oriented parallel to isofrequency laminae. Axons of the fusiform cells exit through the dorsal acoustic stria without branching. The two tuberculoventral association cells in the deep DCN have axons that terminate both in the deep DCN, within the same isofrequency lamina that contains the cell body, and in the ventral cochlear nucleus (VCN). Intracellular recordings from 11 of these cells show that they cannot be distinguished on the basis of their responses to intracellularly injected current. All cell types fired large action potentials that were followed by a fast and a slower undershoot, distinguishing them from cells of the VCN but not from one another. Most cells responded to shocks of the auditory nerve root with early EPSPs and later IPSPs. The latencies of EPSPs show that some were monosynaptic and others polysynaptic. That there was no systematic relationship between the latencies of EPSPs and the cell types from which they were recorded shows that shocks to the nerve root may have activated more than just the large, myelinated, auditory nerve fibers.     

Synaptic relationships of Golgi type II cells in the medial geniculate body of the cat.

A combined analysis with the Golgi and silver-degeneration methods and electron microscopy in the ventral nucleus of the medial geniculate body has confirmed that the Golgi type II neuron forms dendro-dendritic synapses with the principal neuron in terminal aggregates called synaptic nests. Both types of neurons receive synaptic contacts from the afferent axons that ascend from the posterior colliculus and from those that descend from the auditory cortex. Only the principal neuron projects to the auditory cortex. The Golgi type II cells that receive endings from afferent axons send presynaptic processes to principal cells that are also contacted by the very same afferent axons. The axons of Golgi type II cells project to synaptic nests other than those supplied by the dendrites of the parent cell and link the Golgi type II cells with each other. On the surface of the Golgi type II cell there is a segregation of the different types of synaptic endings and a consistent sequence in their synaptic relationships. The endings of colliculogeniculate and Golgi type II axons predominate on the distal dendrites in the synaptic nests. Corticogeniculate endings congregate more on the soma and proximal dendrites. In the synaptic nests the Golgi type II dendrites are presynaptic to the principal cell dendrites, whereas both kinds of dendrites are postsynaptic to the very same axons, which project either from the posterior colliculus or from Golgi II cells...     

Differential expression of the metabotropic glutamate receptor mGluR1α by neurons and axons in the cochlear nucleus: In situ hybridization and immunohistochemistry

mGluR1α is a metabotropic glutamate receptor involved in synaptic modifiability. A differential expression in specific neuronal types could reflect their different connections and response properties in central auditory processing. Using in situ hybridization and immunohistochemistry, we studied mGluR1α receptor expression throughout the cochlear nucleus. Robust labeling occurred in the dorsal cochlear nucleus and small cell shell, with less in the ventral cochlear nucleus. Among the most intensely labeled were the granule cells of the small cell shell. In the dorsal cochlear nucleus, most cell types expressed message and receptor protein, except granule cells. High levels of receptor were expressed by corn cells and cartwheel cells. The terminal dendrites and synaptic spines of cartwheel and fusiform cells contained receptor protein in the molecular layer, where they could synapse with parallel fibers. Fusiform dendrites also expressed mRNA for mGluR1α. The basal dendrites of fusiform cells contained receptor protein in the region where they receive cochlear nerve synapses. Immunostaining of terminal axons was prominent in the molecular layer and the small cell shell, where they were associated with synaptic nests, structures thought to provide long-term changes in excitability. Differential expression levels may reflect different functional requirements of specific cell types, including inhibitory interneurons, like corn cells and cartwheel cells, and excitatory interneurons, like granule cells in the small cell shell, which may participate in local circuits involved in modulatory or gating functions, such as stimulus enhancement or suppression. In presynaptic axons, mGluR1α may relate to the long-term signaling requirements of their modulatory functions. Synapse 28:251–270, 1998. © 1998 Wiley-Liss, Inc.     

Cochlear nerve projections to the small cell shell of the cochlear nucleus: The neuroanatomy of extremely thin sensory axons

Labeling cochlear nerve fibers in the inner ear of chinchillas with biotinylated dextran polyamine was used to trace the thin fibers (Type II), which likely innervate outer hair cells. These axons, 0.1–0.5 μm in diameter, were distinguished from the thicker Type I, fibers innervating inner hair cells, and traced to small‐cell clusters in the cochlear nucleus. This study provided two major new insights into the outer hair cell connections in the cochlear nucleus and the potential significance of very thin axons and synaptic nests, which are widespread in the CNS. 1) EM serial reconstructions of labeled and unlabeled material revealed that Type II axons rarely formed synapses with conventional features (vesicles gathered at junctions). Rather, their endings contained arrays of endoplasmic reticulum and small spherical vesicles without junctions. 2) Type II axons projected predominantly to synaptic nests, where they contacted other endings and dendrites of local interneurons (small stellate and mitt cells, but not granule cells). Synaptic nests lacked intrinsic glia and, presumably, their high‐affinity amino acid transporters. As functional units, nests and their Type II inputs from outer hair cells may contribute to an analog processing mode, which is slower, more diffuse, longer‐lasting, and potentially more plastic than the digital processors addressed by inner hair cells. Synapse 33:83–117, 1999. © 1999 Wiley‐Liss, Inc.     

Dorsal cochlear nucleus projections to the inferior colliculus in the cat: A light and electron microscopic study

In the Present report retrograde and anterograde labeling techniques are used to study the Projections of the dorsal cochlear nucleus (DCN) to the inferior colliculus in the cat. Horseradish peroxidase (HRP) or wheat germ agglutinin (WGA-HRP) injections into the inferior colliculus produce large numbers of labeled neurons in the DCN on the opposite side. Labeled cells with projections to the colliculus are identified as fusiform and giant cells and are organized into rostrocaudal bands. The axons of these DCN neurons are labeled by anterograde transport of ³H-leucine and/or proline and studied in light and electron microscopic autoradiographs. Axons from the DCN terminate within the central nucleus of the inferior colliculus in densely labeled, rostrocaudally oriented bands. Less heavily labeled extensions of these bands are found in the deepest layer of the dorsal cortex, and light labeling is found adjacent to the bands in the central nucleus and in the ventrolateral nucleus. Cells in the dorsomedial DCN project to the most ventromedial part of the central nucleus while progressively more ventrolateral cells in the DCN project to more dorsolateral parts of the central nucleus. This present evidence suggests that the DCN sends afferents to only two of the four subdivisions of the central nucleus. Within these regions, the axons from the DCN form terminal boutons or boutons de passage characterized by medium-sized, round synaptic vesicles. The labeled endings nearly always make asymmetric synaptic contacts on the dendrites of disc-shaped and stellate cells in the central nucleus. A few axosomatic contacts are found on one particular cell type, possibly the stellate variety. The results support the hypothesis that each subdivision of the central nucleus receives afferents from a different set of cell types in the auditory nuclei of the lower brainstem. The banding patterns of the efferent cells in the cochlear nucleus and the axons within the central nucleus suggest that these inputs are congruent to the fibrodendritic layers of the central nucleus and may contribute to tonotopic organization in the central nucleus. Finally, the results suggest that each of the two major classes of cells in the central nucleus receives different patterns of inputs from the DCN. These morphological differences could contribute to different electrophysiological responses to the sound stimuli by these cells.     

Horizontal cell axons and axon terminals in goldfish retina

Retinas of ordinary and black moor varieties of goldfish (Carassius auratus) were prepared by the Golgi method, mounted flat or sectioned vertically, and studied in the light microscope. Three types of horizontal cells whose dendrites contact only cones, and one type whose dendrites contact only rods, were observed. The cone horizontal cells (Cajal's “external horizontal cells”) all have slender axons which descend gradually to the inner nuclear layer and terminate there in long, fusiform expansions (Cajal's “internal horizontal cells”). The thin and thick portions of the axons, as well as the perikarya of the horizontal cells, bear small numbers of straight, horizontally-directed, knobby filamentous appendages which may be sites of synaptic contact. The cone horizontal cell axons in goldfish, unlike those in higher vertebrates, do not terminate in contact with synaptic endings of photoreceptor cells, but in proximity to cells and processes deep in the inner nuclear layer. Axons have not yet been demonstrated on rod horizontal cells in goldfish.     

Cartwheel neurons of the dorsal cochlear nucleus: a Golgi-electron microscopic study in rat

Cartwheel neurons in rat dorsal cochlear nucleus (DCN) were studied by Golgi impregnation-electron microscopy. Usually situated in layers 1-2, cartwheel neurons (10-14 micrometers in mean cell body diameter) have dendritic trees predominantly in layer 1. The dendrites branch at wide angles. Most primary dendrites are short, nontapering, and bear only a few sessile spines. Secondary and tertiary dendrites are short, curved, and spine-laden. The perikaryon forms symmetric synapses with at least two kinds of boutons containing pleomorphic vesicles. The euchromatic nucleus is indented and has an eccentric nucleolus. The cytoplasm shows several small Nissl bodies, a conspicuous Golgi apparatus, and numerous subsurface and cytoplasmic cisterns of endoplasmic reticulum with a narrow lumen, joined by mitochondria in single or multiple assemblies. In primary dendrites mitochondria are situated peripherally, while in distal branches they become ubiquitous and relatively more numerous. Dendritic shafts usually form symmetric synapses with boutons that contain pleomorphic vesicles. The majority of the dendritic spines are provided with a vesiculo-saccular spine apparatus. All dendritic spines have asymmetric synapses. Most of these are formed with varicosities of thin, unmyelinated fibers (presumably axons of granule cells) running parallel to the long axis of the DCN or radially. These varicosities contain round, clear synaptic vesicles. On the initial axon segment few symmetric synapses are present. The axon acquires a thin myelin sheath after a short trajectory. Cartwheel neurons outnumber all other neurons in layers 1-2 (with the exception of granule cells), and presumably correspond to type C cells with thinly myelinated axons described by Lorente de Nó. The axons of these neurons provide a dense plexus in the superficial layers without leaving the DCN. The possible functional role of cartwheel neurons is discussed.     

Distribution and light microscopic features of granule cells in the cochlear nuclei of cat, rat, and mouse

In the present study the cytology and the topography of the cochlear granule cell domain (a comprehensive term introduced here for all granule cell-containing regions of the cochlear nuclear complex) have been studied light microscopically in Nissl, Bielschowsky, and Golgi-Del Rio-Hortega material of cats, rats, and mice; in Golig rapid material of 0-14-day-old kittens; and in sections of 6-week-old kittens following HRP injections in the superficial dorsal cochlear nucleus (DCN). The domain has been parcellated in seven subdivisions which, in spite of some species' differences, are easily identifiable in all of the included animals. The cochlear granule cells are considered as a particular class of neuron, which is slightly different from, but nevertheless principally similar to the cerebellar granule cells in both shape and mode of neuronal connections. The digitiform terminals of the cochlear granule cells differentiate after the first two weeks of extrauterine life. In several respects these cells show larger variation among species than do the cerebellar granules, the similarity between the two classes of granule cells being most conspicuous in the rodent. The silver, Golgi rapid, and HRP material suggest that all, or at least the majority, of the granule cell axons project to the molecular layer of the DCN, forming parallel fibers similar to those of the cerebellar cortex. Also, the cochlear parallel fibers traverse the spiny apical dendrites of principal neurons (the pyramidal cells) and the smoother dendrites of molecular layer stellate cells.     

The facial motor nucleus of the opossum: Synaptic endings on dendrites

The diameters of dendrites of large, medium and small neurons (Falls and King, '76) were measured from Golgi impregnations of the opossum facial motor nucleus in order to classify dendritic profiles sectioned in the transverse plane in electron micrographs. Three categories of dendrites are described: (1) proximal (4–7 μ in diameter); (2) intermediate (2–4 μ in diameter) and (3) distal (0.5–2 μ in diamter). The distribution of axodendritic synaptic endings was determined, recognizing that the neuronal source of individual dendritic profiles when seen in the transverse plane of section cannot be absolutely determined in view of the overlap in size of the dendrites issuing from the three types of neurons. Presynaptic terminals were categorized according to vesicle shape (spherical, pleomorphic or ellipsoidal), vesicle size, terminal size, junctional characteristics and post synaptic distribution. The vesicle size is expressed as a mean area (nm²) and was determined by using a cybergraphic tablet and a PDP-12 computer system. In any given plane of section, synaptic terminals cover most of the membrane of proximal dendrites and decrease in number as intermediate and distal dendrites are encountered. In Golgi impregnations four classes of afferent fibers which ramify among the dendrites of facial neurons can be distinguished. As yet, their sources have not been identified. Possible sites of origin for presynaptic profiles are discussed in the context of previous light microscopic findings.     

The cytoarchitecture of the dorsal cochlear nucleus in the 3-month- and 26-month-old C57BL/6 mouse: A golgi impregnation study

The cytoarchitecture of the dorsal cochlear nucleus (DCN) was compared in 3- and 26-month-old C57BL/6 mice. The effects of genetically controlled progressive hearing loss present in the CNS in this mouse strain were analyzed with Nissl-stained and Golgi-impregnated material. The DCN was divided into the superficial molecular, an intermediate fusiform-granule, and the deep polymorphic layers. The molecular layer (ML) consisted of many fibers and a few small ovoid to spherical, fusiform, and granule cells. The fusiform-granule layer (FL) contained large fusiform and many granule cells. Most FL fusiform cells were oriented with their long axes perpendicular to the DCN surface and were present as small aggregations or individually. Cartwheel cells were adjacent to the FL fusiform cells. The deep polymorphic layer (PL) contained spherical, fusiform, granule, and multipolar neurons. The granule cells formed a dorsal cap of the DCN. From this cap, sheets of granule cells separated the DCN from the posterior ventral cochlear nucleus (PVCN) and from the brainstem. The internal organization, neuronal location, orientation, and morphology were similar in both age groups. The granule cells had four to five primary dendrites, varicosities, and few to no dendritic appendages. The FL fusiform cells displayed different dendritic morphology in the two ages. One or two elaborate primary ML apical dendrites in the 3-month-old mice were covered with spikelike dendritic spines. The basal one or two PL dendrites were less elaborate and had few dendrite spines. In contrast, FL fusiform neurons in 26-month-old mice had regular dendritic varicosities and fewer spines which were short and stumpy. Basal dendrites had varicosities and interruptions. Cartwheel neurons in 3-month-old mice had elaborate ML dendritic trees covered with dendritic spines. In 26-month-old mice the dendrites had many varicosities and fewer short blunted dendritic spines. Large multipolar neurons in older mice had thinner dendrities with more varicosities than were in the 3-mcnth group. In both age groups multipolar cells had few dendritic spines limited distally. Small and large spherical cells had two to five primary dendrites with varicosities, little higher-order branching, and spines. Fusiform cells had one or two primary dendrites, little secondary branching, and few to no spines. Minor degenerative changes were noted in spherical and fusiform cells in the two age groups. These included dendritic varicosities, interruptions, and some irregularities of somata surface. Degenerative changes present in the cochlea had significant effects on a limited population of DCN neurons. Finally, the neusronal morphology and architecture of the DCN in C57BL/6 mouse is similar to other mammalian species.     

Projections from auditory cortex to the cochlear nucleus in rats: Synapses on granule cell dendrites

Previous work has demonstrated that layer V pyramidal cells of primary auditory cortex project directly to the cochlear nucleus. The postsynaptic targets of these centrifugal projections, however, are not known. For the present study, biotinylated dextran amine, an anterograde tracer, was injected into the auditory cortex of rats, and labeled terminals were examined with light and electron microscopy. Labeled corticobulbar axons and terminals in the cochlear nucleus are found almost exclusively in the granule cell domain, and the terminals appear as boutons (1–2 μm in diameter) or as small mossy fiber endings (2–5 μm in diameter). These cortical endings contain round synaptic vesicles and form asymmetric synapses on hairy dendritic profiles, from which thin (0.1 μm in diameter), nonsynaptic “hairs” protrude deep into the labeled endings. These postsynaptic dendrites, which are typical of granule cells, surround and receive synapses from large, unlabeled mossy fiber endings containing round synaptic vesicles and are also postsynaptic to unlabeled axon terminals containing pleomorphic synaptic vesicles. No labeled fibers were observed synapsing on profiles that did not fit the characteristics of granule cell dendrites. We describe a circuit in the auditory system by which ascending information in the cochlear nucleus can be modified directly by descending cortical influences. © 1996 Wiley-Liss, Inc.     

Morphology and physiology of cells in slice preparations of the posteroventral cochlear nucleus of mice

In an effort to understand what integrative tasks are performed in the cochlear nuclei, the present study was undertaken to describe neuronal circuits in the posteroventral cochlear nucleus (PVCN) anatomically and physiologically. The cochlear nuclear complex receives auditory information from the cochlea through the auditory nerve. Within the cochlear nuclei, signals travel along several parallel and interconnected pathways. From the cochlear nuclei, transformed versions of the signals are passed to higher auditory centers in the brainstem. We have recorded electrophysiological responses from cells that were subsequently visualized with horseradish peroxidase (HRP). Responses to shocks to the auditory nerve root and to intracellularly injected current pulses were recorded and correlated with morphology. Two types of stellate cells and octopus cells were distinguished. T stellate cells project out of the cochlear nuclei through the Trapezoid body; D stellate cells do not. The axons of D stellate cells extend Dorsalward to the dorsal cochlear nucleus (DCN) but have not been traced out of the nucleus. Both T and D stellate cells have terminal collaterals in the multipolar cell region of the PVCN and in the DCN. The endings of one T stellate cell formed a narrow band rostrocaudally in the fusiform cell layer of the DCN that resembled an isofrequency band. The endings of one D stellate cell lay closely apposed to multipolar cells in the deep layer of the DCN. The dendrites of T stellate cells are often aligned along the path of auditory nerve fibers and end in tufts, whereas those of D stellate cells extend radially in the plane of the lateral surface of the PVCN toward granule cell areas and branch sparingly. Octopus cells have dendrites oriented perpendicularly to the path of auditory nerve fibers. Their axons were cut medially in the slices; none had collateral branches. Both T and D stellate cells were monosynaptically excited to threshold by shocks to the nerve root, indicating that they could participate in local circuits that we measure physiologically. T stellate cells have action potentials that peak at about 0 mV and are followed by single undershoots. The D stellate cell that was best impaled fired overshooting action potentials that were followed by double undershoots. Octopus cells were monosynaptically excited to threshold by shocks to the auditory nerve.     

Electron microscopic features of physiologically characterized, HRP-labeled fusiform cells in the cat dorsal cochlear nucleus

We report on the anatomy and physiology of three fusiform cells in the dorsal cochlear nucleus (DCN) of the cat. The extra- and intracellular responses of these cells to pure tones showed features typical of the cell type. Peristimulus time histograms (PSTHs) were usually of the pauser or buildup configuration with chopping behavior noted in certain instances. Intracellular records during stimulus presentations revealed sustained depolarizations for the duration of the tone followed by a prolonged after-hyperpolarization (AHP). On rare occasions, a hyperpolarization corresponding to the pause region of the PSTH was noted. Occasionally, a stimulus-induced depolarization would be maintained after stimulus offset. Rebound excitation was also observed after the AHP. Morphologically, all three cells showed the standard fusiform cell features at the light microscopic level. The cell body gave rise to apical and basal dendritic trees. The apical tree branched frequently and displayed numerous spines distally. The basal tree had fewer branches and fewer, more irregular appendages. The axon originated from the cell body and gave rise to one or more collaterals before leaving the nucleus via the dorsal acoustic stria (DAS). At the electron microscopic (EM) level, the axon collaterals may terminate on a variety of cell types in the DCN, including fusiform cells. Their vesicles are round and the terminals closely resemble many unlabeled terminals seen on the cell body and apical and basal dendrites of our labeled fusiform cells. Terminals containing round vesicles, believed to be eighth nerve terminals, were found, with one exception, only on the basal dendrites. The spine-laden, distal apical dendrites received primarily terminals containing round vesicles, presumed to originate from the unmyelinated axons of granule cells. The cell body and unmyelinated initial segment received mostly terminals containing pleomorphic and flat vesicles, which also made up a large percentage of the dendritic input. Some relevant correlations, between the distribution of synaptic terminals and the observed physiology, may be possible.     

Differentiation of the giant and fusiform cells in the dorsal cochlear nucleus of the hamster

Two categories of large neurons--fusiform cells and giant cells--can be distinguished in the dorsal cochlear nucleus of the hamster. In the adult, these neurons are located in separate laminae in the nucleus and have distinct dendritic morphology. However, the two cell types are not distinguishable in the newborn hamster. At birth the large cells in the dorsal cochlear nucleus are clustered into one group and are alike morphologically. On postnatal day 5, laminae are still not apparent, but the neurons have begun to acquire their adult shapes. By day 15 laminae have formed, and the cells appear mature with the one exception that the apical dendrites of the fusiform cells have not acquired the spines which will cover their surface in the adult. The appearance of laminae coincides with the growth of axons and dendrites into a interstitial zone between the layers of cell bodies. Dendritic growth occurs during the time of axonal ingrowth and establishment of contacts between the axons and dendrites. The growth of the apical dendrites of fusiform cells, which are not contacted by these fibers, lags behind. These results demonstrate that afferent ingrowth and the differentiation of dendrites in the dorsal cochlear nucleus are temporally related. The synchronous development may serve to ensure a specific synaptic arrangement between the axons and their target dendrites.     

Fine structure of the cell clusters in the cochlear nerve root: Stellate,granule, and mitt cells offer insights into the synaptic organization of local circuit neurons

The small cell shell of the cochlear nucleus contains a complex integrative machinery which can be used to study the roles of interneurons in sensory processing. The cell clusters in the cochlear nerve root of the chinchilla provide the simplest example of this structure. Reported here are the neuronal architecture and synaptic organization of the three principal cell types and the three distinctive neuropil structures that could be characterized with the Nissl and Golgi methods and electron microscopy. Granule cells were characterized by several dendrites with claw-like terminals that received synaptic contacts from multiple excitatory mossy fiber rosettes. Given their relatively large number and their prolific parallel fiber synapses, the granule cells provide a suitable substrate for a tangential spread of excitatory activity, which could build to considerable proportions. The mitt cells had a thickened, single dendrite, its terminal branches arranged in a shape reminiscent of a baseball catcher's mitt. The dendritic mitt enclosed an enormous, convoluted mossy fiber rosette forming many excitatory synapses on just one cell. This could provide for a discrete, comparatively fast input-output relay of signals. Small stellate cells had longer, radiating dendrites that engaged the synaptic nests. These nests were strung in long strands, containing heterogeneous synapses from putative excitatory and inhibitory inputs. Given the prevalence of the synaptic nests, the small stellate cells appear to have the greatest integrative capacity. They provide the main output of the synaptic nests. © 1996 Wiley-Liss, Inc.     

Morphology and synaptic connections of ultrafine primary axons in lamina I of the spinal dorsal horn: candidates for the terminal axonal arbors of primary neurons with unmyelinated (C) axons

Neurons in Rexed's lamina I have the bulk of their dendritic arbors confined within this lamina. This study examines the morphology and synaptic connections of primary axons which generate axonal endings in lamina I of the spinal dorsal horn and are in position to deliver their inputs directly to lamina I neurons. Primary axons were made visible for light and electron microscopical study by applying horseradish peroxidase (HRP) to the severed central stumps of cervical and lumbar dorsal roots and allowing sufficient time for the orthograde movement of the HRP into the terminal axonal arbors. Golgi preparations provided supplementary light microscopical views of these axons. Lamina I receives the terminal arborization of two very different kinds of primary axons. One of these generates many ultrafine endings along unbranched, long rostrocaudally oriented, strand-like collaterals which arise from thin parent branches in Lissauer's tract. In view of these thin parent branches, most ultrafine primary axons are considered to be unmyelinated (C) primary axons. The second kind of primary axon generates large caliber endings on branched collaterals. These arise from relatively thick parent branches in Lissauer's tract which, on the basis of their size, are considered to be myelinated (A delta) primary axons. The scalloped endings of both primary axons lie in the interior of glomeruli where they form axodendritic synapses on small dendritic shafts and spines. It is at these synapses that these two kinds of primary axons are thought to transfer nociceptive and thermal inputs directly to the dendritic arbors of lamina I neurons. Transmitter release at these axodendritic synapses in response to primary inputs can be modified, probably diminished or inhibited, by synaptic events within the glomeruli from at least three sources. Synaptic vesicle-containing dendrites form dendroaxonic synapses on primary endings and two kinds of axons form axoaxonic synapses either on primary endings or on the intervaricose segments of the primary axons.     

Projections to the inferior colliculus from the anteroventral cochlear nucleus in the cat: possible substrates for binaural interaction

The projections to the inferior colliculus of the cat are shown in autoradiographs after injections of 3H-amino acids into the anteroventral cochlear nucleus and anterograde axonal transport. Labeled bands of axons are seen in the central nucleus of the inferior colliculus, parallel to the fibrodendritic laminae, and in layers 3 and 4 of the dorsal cortex. A bilateral projection to the lateral, low-frequency part of the inferior colliculus is observed. In contrast, the more ventromedial, mid- and high-frequency parts receive only a contralateral input. The projections from the cochlear nucleus to both the contralateral midbrain and bilaterally to the superior olivary complex appear to be tonotopically organized. After HRP injections in the inferior colliculus, small numbers of stellate neurons are labeled in the lateral and ventral low-frequency parts of the anteroventral cochlear nucleus on the ipsilateral side. EM autoradiographs show labeled axonal endings from both sides of the anteroventral cochlear nuclei are present in the same proportion in pars lateralis. Axonal endings from either cochlear nucleus have small, round synaptic vesicles and make asymmetric synaptic contacts on dendrites. Axons from the contralateral side also make axosomatic contacts on large disc-shaped or stellate cells. Neurons from the ipsilateral anteroventral cochlear nucleus apparently make more synaptic endings per cell as compared to neurons from the contralateral side. Together, bilateral inputs from the anteroventral cochlear nucleus can account for a third of the endings with round synaptic vesicles in pars lateralis of the central nucleus. Morphological similarities among the ascending inputs to the inferior colliculus are discussed. Both direct circuits from the cochlear nucleus to the inferior colliculus and indirect circuits via the superior olivary complex or lateral lemniscus may display banding patterns, nucleotopic organization, or differential synaptic organization. The direct inputs from the anteroventral cochlear nucleus to the colliculus may influence binaural interactions which occur in the superior olivary complex. In addition, direct inputs may create new binaural responses in the inferior colliculus that are independent of lower centers.     

GABA neurons in the superficial layers of the rat dorsal cochlear nucleus: light and electron microscopic immunocytochemistry

This article is an application of light and electron microscopic immunocytochemistry to the study of the neuronal circuit of the superficial layers in the rat dorsal cochlear nucleus (DCN). An antiserum against the intrinsic marker glutamate decarboxylase (GAD) is used to identify and map axon terminals and neurons that use gamma aminobutyric acid (GABA) as a neurotransmitter. It is demonstrated that layers 1 and 2 of the DCN contain a very high density of GABAergic boutons, matched only by the granule cell domains of the ventral cochlear nucleus, especially the superficial granule cell domain. These two layers also contain much higher concentrations of GABAergic cell bodies than all other magnocellular regions of the cochlear nuclear complex. Cartwheel and stellate neurons, and probably also Golgi cells, previously characterized in Golgi and electron microscopic investigations, appear immunostained and, therefore, are presumably inhibitory. The synaptic relations between parallel fibers, the axons of granule cells, and cartwheel and stellate neurons are confirmed. The present study also supports the conclusion that stellate cells are coupled to one another by gap junctions. Also scattered in layer 1 are large, GABAergic neurons that occur with irregular frequency and presumably represent displaced Purkinje cells, previously identified with a Purkinje-cell-specific marker. Granule neurons and pyramidal neurons remain unstained, even after topical injection of colchicine, which enhances immunostaining of the other glutamate-decarboxylase-positive cells, and therefore must use transmitters different from GABA. The possible analogies between the spiny cartwheel and the aspiny stellate cells of the DCN and the cerebellar Purkinje and stellate/basket cells are discussed in the light of data from Golgi, electron microscopy, and transmitter imunocytochemistry.     

Immunocytochemical localization of GABA in the cochlear nucleus of the guinea pig

The immunocytochemical distribution of gamma-aminobutyric acid (GABA) was determined in the cochlear nucleus of the guinea pig using affinity-purified antibodies made against GABA conjugated to bovine serum albumin. Light microscopic immunocytochemistry shows immunoreactive puncta, which appear to be GABA-positive presynaptic terminals, distributed throughout the cochlear nucleus. In the ventral cochlear nucleus, these puncta are often found around unlabeled neuronal cell bodies. While occasional labeled small cells are found in the ventral cochlear nucleus, most GABA-immunoreactive cell bodies are present in the superficial layers of the dorsal cochlear nucleus. Based on size and shape, immunoreactive cells in the dorsal cochlear nucleus are divided into 3 classes: medium round cells with diameters averaging 16 microns, small round cells with average diameters of 9 microns and small flattened cells with major and minor diameters averaging 11 and 6 microns, respectively. Labeled fusiform and granule cells are not seen. A similar distribution of label was seen using antibodies against glutamic acid decarboxylase. Electron microscopic immunocytochemistry of the anteroventral cochlear nucleus shows GABA immunoreactive boutons containing oval/pleomorphic synaptic vesicles on cell bodies and dendrites. Other major classes of terminals, including those with small round, large round and flattened synaptic vesicles are unlabeled.