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PubMed1982

The neuronal architecture of the anteroventral cochlear nucleus of the cat in the region of the cochlear nerve root: electron microscopy.

We have studied the posterior division of the anteroventral cochlear nucleus, where the cochlear nerve root enters the brain, in the cat. In Nissl preparations, this region contains two types of neuronal cell bodies: globular and multipolar. The two types can be identified in the electron-microscope by comparing Nissl substance and rough endoplasmic reticulum. Globular cell bodies receive many synaptic terminals, which cover 85% of the surface. In contrast, multipolar cell bodies are almost entirely wrapped by thin glial sheets--synaptic terminals contact less than 15% of the surface and tend to cluster at the bases of dendrites. Synaptic terminals are of three kinds, types 1, 2, and 3, which contain large round, small round-to-oval, and small flattened synaptic vesicles, respectively. Terminals of all three kinds synapse on both types of cell bodies. However, only globular cell bodies receive the largest type 1 terminals, which correspond to end-bulbs, seen in Golgi impregnations to arise from cochlear nerve axons. Cochlear ablation leads to degeneration of type 1, but not type 2 or 3 terminals. We conclude that neurons with globular cell bodies receive heavy somatic input from the cochlear nerve, as well as from other sources. Neurons with multipolar cell bodies receive very little input to their perikarya--giving their dendrites a more important role in determining their response properties. We suggest a morphological basis for correlating individual kinds of neurons with certain electrophysiological response types.

Animals↗
PubMed1998

Localization of neurotrophin-3-like immunoreactivity in the rat cochlear nucleus.

Immunohistochemistry as well as immunohistofluorescence were used to investigate the distribution of the neurotrophin-3 (NT3) in the adult rat cochlear nucleus. We found a widespread distribution of NT3 immunolabeled neurons throughout the three divisions of this nucleus. NT3-like immunoreactivity was clearly population-specific, with some cell groups heavily (various small neurons and granule cells) or moderately (large neurons of the ventral cochlear nucleus) stained, while others remained negative (a major fraction of medium and large neurons of the dorsal cochlear nucleus). Double-labeling experiments were performed using antibody against the glial fibrillary acid protein, a classic marker for mature astrocytes. This colocalization study revealed that NT3 immunoreactivity was also present in a subpopulation of astrocytes, particularly in the glia limitans and their projections. Numerous small cells also colocalized NT3 together with the glial marker in the granule cell domain and in the molecular cell layer of the dorsal cochlear nucleus. These results suggest that NT3 may exist in widespread populations of adult cochlear nucleus neurons as well as in glial cells. This abundant distribution of NT3-like immunoreactivity implies that this neurotrophin may have an important role in the continued maintenance of mature cochlear nucleus and makes it an attractive candidate for playing a role in regulation or stabilization of neuronal circuits in this nucleus.

Animals↗
PubMed1985

Responses of cochlear nucleus units to electrical stimulation through a cochlear prosthesis: channel interaction.

The responses of 39 single units in the ventral cochlear nucleus of acute anesthetized guinea-pigs were studied with continuous electrical stimuli presented through a dual-channel implant in the scala tympani. Implants had four electrodes placed along the axis of the cochlea with 1 mm separations. With a specific pair (either apical or basal) of stimulating electrodes, about half of the units responded when current was flowing apically, while the rest responded to current in the opposite direction. No obvious relation existed between the effective polarity of the basal and apical pairs of electrodes. Two response types were observed while stimulating through both pairs simultaneously. Seventy-nine percent of the units responded to the sum of the current waveforms presented through the two pairs. Twenty-one percent responded to the difference between the waveforms. Both types of responses were observed for suprathreshold as well as for some intensities of stimulation that alone were subthreshold. The type of response was not dependent on the absolute threshold or threshold difference between the two pairs. For equal peak-intensities of stimuli, two-channel stimulation evoked larger responses than single-channel stimulation, provided the two channels were in their effective polarities. Responses to dual-channel stimulation were consistently larger than the summed responses to the two individual single-channel stimuli. The observed responses to the dual-channel stimulation indicate that the adequate stimulus was determined by the linear combination of fields produced by the individual channels in the vicinity of the stimulating electrodes before the auditory nerve is stimulated, and that excitation takes place in a spatially restricted area of the auditory nerve.

Animals↗
PubMed1991

Projections from the posteroventral cochlear nucleus to the superior olivary complex in guinea pig: light and EM observations with the PHA-L method.

The efferent neural projections from posteroventral cochlear nucleus to the superior olivary complex in guinea pig were examined with the Phaseolus vulgaris-leucoagglutinin anterograde tract-tracing method. Light microscopic analysis demonstrated that the posteroventral cochlear nucleus of guinea pig bilaterally projects to the superior para-olivary nucleus and the rostral, medioventral, and lateroventral peri-olivary regions. Ipsilaterally, the posteroventral cochlear nucleus projects to the lateral superior olive, the caudal peri-olivary region, and areas immediately surrounding the capsule of the lateral superior olive. Contralaterally, the posteroventral cochlear nucleus projects to the medial nucleus of the trapezoid body. All of these projection axons travel in the trapezoid body and their terminals make, primarily, en passant endings upon their targets. Exclusively within the contralateral medial nucleus of the trapezoid body, some neurons terminate also in calyceal endings. The assumption that immunolabeled structures observed with light microscopy made actual synaptic contact in their respective target areas was confirmed with electron microscopy. With postembedding immunocytochemical procedures applied to thin sections, the electron microscope revealed labeled synaptic vesicles and pre- and postsynaptic membrane specializations. The projection pattern from posteroventral cochlear nucleus was found to be topographically organized in three distinct regions. The more dorsally located neurons of the posteroventral cochlear nucleus terminated dorsomedially in the ipsilateral lateral superior olive, ventromedially in the contralateral superior para-olivary nucleus, and medially in the contralateral medioventral peri-olivary region. The more ventrally located neurons of the posteroventral cochlear nucleus terminated dorsolaterally in the ipsilateral lateral superior olive, dorsolaterally in the contralateral superior para-olivary nucleus, and laterally in the contralateral medioventral peri-olivary region.

Animals↗
PubMed2003

Activity-dependent regulation of the subcellular localization of neuronal calcium sensor-1 in the avian cochlear nucleus.

Neurons in the avian cochlear nucleus, nucleus magnocellularis (NM), are highly sensitive to manipulations of afferent input, and removal of afferent activity through cochlear ablation results in the death of approximately 20-40% of ipsilateral NM neurons. The intracellular cascades that determine whether an individual NM neuron will die or survive are not fully understood. One early event observed in NM following deafferentation is a rapid rise in intracellular calcium concentration. In most cellular systems, the activity of calcium-binding proteins is believed to accommodate calcium influx. The calcium-binding protein, neuronal calcium sensor-1 (NCS-1), is an intracellular neuronal calcium sensor belonging to the EF-hand superfamily. NCS-1 has been implicated in calcium-dependent regulation of signaling cascades. To evaluate NCS-1 action in NM neurons, the localization of NCS-1 protein was examined. Double-label immunofluorescence experiments revealed that NCS-1 expression is evident in both the presynaptic nerve terminal and postsynaptic NM neuron. The postsynaptic expression of NCS-1 typically appears to be closely associated with the cell membrane. This close proximity of NCS-1 to the postsynaptic membrane could allow NCS-1 to function as a modulator of postsynaptic signaling events. Following deafferentation, NM neurons were more likely to show diffuse cytoplasmic NCS-1 labeling. This increase in the number of cells showing diffuse cytoplasmic labeling was observed 12 and 24 h following cochlea ablation, but was not observed 4 days following surgery. This activity-dependent regulation of NCS-1 subcellular localization suggests it may be associated with, or influenced by, processes important for the survival of NM neurons.

Action Potentials↗
PubMed1987

Effect of olivocochlear bundle transection on choline acetyltransferase activity in the rat cochlear nucleus.

Using a microdissection and quantitative microassay approach, choline acetyltransferase activities were mapped in the cochlear nuclei of rats having complete transections of the olivocochlear bundle on one side in the brain stem. In rats in which the trapezoid body was not significantly damaged by the lesion, consistent reductions of choline acetyltransferase activity in subregions of the lesion-side cochlear nucleus, as compared to the control side, averaged about 20%. Nevertheless, a profound lesion-side reduction of choline acetyltransferase activity was found in a branch connection from the olivocochlear bundle to the cochlear nucleus. The results suggest that branches from the olivocochlear bundle are cholinergic, but contribute a relatively minor proportion of the cholinergic synapses in all regions of the rat cochlear nucleus. In the light of previous results with more extensive lesions, it can be proposed that synapses in all regions of the rat cochlear nucleus. In the light of previous results with more extensive lesions, it can be proposed that most cholinergic input into the rat cochlear nucleus enters by a ventral route along the trapezoid body. It is noted that this represents a quantitatively somewhat different situation from that in the cat.

Acetylcholinesterase↗
PubMed2000

Cytology of large neurons in the guinea pig dorsal cochlear nucleus contacting the inferior colliculus.

Large neurons in the dorsal cochlear nucleus of the guinea pig which project to the inferior colliculus were identified after injections of the neural tracer WGA-HRP. Retrograde labelled cells (pyramidal and giant neurons) in the dorsal cochlear nucleus were glycine and GABA immunonegative and showed a similar ultrastructure. Between 30 and 60% of their perimeter was covered by axo-somatic boutons, most of which (>50%) contained pleomorphic synaptic vesicles. Other boutons (about 40% of total) contained flat vesicles and few (5-6%) contained round vesicles, a characteristic of the excitatory cells innervating the inferior colliculus. Immunogold-cytochemistry, coupled to silver intensification, showed that more than 50% of axo-somatic pleomorphic boutons and over 90% of boutons containing flat and pleomorphic vesicles store glycine. Rare WGA-HRP labelled axo-somatic boutons containing flat-pleomorphic vesicles were seen on pyramidal and giant neurons. This suggests that a few inhibitory collicular terminals contact the excitatory large neurons in the dorsal cochlear nucleus.

Animals↗
PubMed1997

Neurotrophin receptor immunostaining in the rat ventral cochlear nucleus.

By virtue of its known segregated distribution of cell types, their known neurotransmitters and neurophysiologic properties, the cochlear nucleus is an excellent model and provides the opportunity to study the relation between neurotrophins and their receptors along with the functional properties of the adult cochlear nucleus. To investigate the potential role of neurotrophins in the mature cochlear nucleus, we determined the expression of the three major neurotrophin tyrosine kinase receptors (Trk) in the adult rat ventral cochlear nucleus, as revealed by antibodies against the full-Trk proteins. A qualitative and a cartographic analysis showed a widespread distribution of the three Trk receptors throughout the nucleus. Immunostaining was mainly restricted to neurons as shown by the lack of double immunostaining with specific markers for glial cells. However, we observed variability in immunostaining for given receptors. Three classes of cells were distinguished by their specificity for Trk receptors. The first one was a cell population that stained for TrkA or TrkB. This population characterizes the majority of small and small round neurons and fusiform cells. The second group consists of TrkC-immunolabeled cells and comprises the majority of spherical, globular, granule and small multipolar cells. The third group consists of cells that seem to be immunopositive for all three Trk receptors. This group includes in part multipolar, giant and octopus cells. A possible correlation between Trk expression and cell functional properties is suggested: TrkA- and TrkB-immunoreactive cells could include inhibitory neurons while cells stained for TrkC could include excitatory neurons. The abundant and widespread neuronal distribution of signal-transducing forms of TrkA, TrkB and TrkC predicts that their cognate ligands may exert significant effects on a large proportion of neurons within the mature ventral cochlear nucleus.

Age Factors↗
PubMed1993

Morphometric development of the human auditory system: ventral cochlear nucleus.

The development of the human cochlear nucleus was studied in serial sections of the brain of 12 fetuses at 12-40 weeks of gestation, an infant at 2 months of age and an adult of 63 years using an electronic planimeter with a computer. Morphometric analysis of the development of the ventral cochlear nucleus showed that its development accelerates after 18 weeks of gestation in terms of columnar volume, columnar length, neuronal number and neuronal size.

Auditory Pathways↗
PubMed1986

Identification of glycinergic synapses in the cochlear nucleus through immunocytochemical localization of the postsynaptic receptor.

The distribution and morphology of glycinergic synapses in the cochlear nucleus were investigated using monoclonal antibodies against the glycine receptor. Glycine receptor immunoreactivity was seen on somas and proximal processes of most cells in all divisions of the cochlear nucleus; distribution of label in neuropil was denser in the dorsal cochlear nucleus and granule cell cap than in the ventral cochlear nucleus. At the ultrastructural level, glycine receptor immunoreactivity was specifically distributed postsynaptically to terminals that contained flattened vesicles in the guinea pig anteroventral cochlear nucleus. These studies show that the immunocytochemical localization of the glycine receptor can provide a means of identifying and characterizing glycinergic synapses throughout the central nervous system.

Animals↗
PubMed1983

Physiological response properties of cells labeled intracellularly with horseradish peroxidase in cat ventral cochlear nucleus.

To determine the correspondence between anatomical and physiological cell types in the ventral cochlear nucleus of the cat, intracellular injections of horseradish peroxidase were made into cells whose extracellular and intracellular responses to sound had been studied. Three identified cells responded to a short tone burst at their characteristic frequencies with an onset pattern. This pattern is characterized by a strong response to the onset of the stimulus. One was an octopus cell. The second cell, located in the octopus-cell area, was a giant cell with a few somatic spines and thin tapering dendrites; the intracellular record revealed that even in the absence of sound it received continuous synaptic input, while tones at characteristic frequency produced a sustained depolarization. A third cell, which had an onset response at low intensities and a chopper response at high intensities, was a stellate cell located in the intermediate acoustic stria with dendrites oriented parallel to the fiber tract. This cell had an unusually broad dynamic range in response to changes in intensity. Two cells with transient chopper response patterns were stellate cells in the posteroventral cochlear nucleus with many branched and beaded dendrites. Three cells with more sustained chopper response patterns were stellate cells in the anteroventral cochlear nucleus with fewer, less-branched, smooth dendrites. Two cells with primarylike responses to tones were bushy cells with numerous short, thin, highly branched dendrites in the posterior division of the anteroventral cochlear nucleus. Intracellular responses to tones at the characteristic frequency consisted of large brief depolarizations, which were not sustained. Another cell, which responded to tones in a phase-locked fashion, was also located in the anteroventral cochlear nucleus. It was a small, stellate cell with relatively few, smooth dendrites. The labeled cells largely support previous attempts at physiological-morphological correlations: (1) bushy cells exhibit primarylike pattern; (2) stellate cells exhibit chopper patterns; and (3) octopus cells exhibit an onset pattern. It was also demonstrated that more than one cell type can exhibit a particular response pattern.

Animals↗
PubMed1985

Quantitative analyses of axonal endings in the central nucleus of the inferior colliculus and distribution of 3H-labeling after injections in the dorsal cochlear nucleus.

Quantitative analyses of electron microscopic (EM) autoradiographs were used to identify the afferents from the dorsal cochlear nucleus in the central nucleus of the inferior colliculus (IC) in the cat. In order to localize the sources of radioactivity, material from axonal transport experiments was analyzed by means of a hypothetical grain procedure which takes the cross-scatter of beta particles into account. Measurements of the synaptic vesicles in axonal endings and a cluster analysis were used to identify different groups of endings. In order to determine which types of endings arise in the dorsal cochlear nucleus, axonal endings labeled after axonal transport and unlabeled endings were characterized and compared to the groups defined by the cluster analysis. Axonal endings with round synaptic vesicles were labeled with more than 2 grains/micron2 which was about 30% of the radioactivity in the central nucleus of the IC. This was six to seven times greater than if the radioactivity had been randomly distributed. Other tissue compartments usually had less radioactivity. Some myelinated and unmyelinated axons were labeled, but, as a group they had lower amounts of radioactivity than predicted by random labeling. In most cases, only low levels of activity were found in glial and postsynaptic structures. Five groups of axonal endings in the medial part of the central nucleus were identified by an analysis which clustered similar types of endings. The variance of the longest axis, the mean diameter, the variance of area, and the mean area of the synaptic vesicles were the variables most useful in distinguishing these five groups. Axonal endings with round synaptic vesicles were classified as either small, or large, or very large, while endings with pleomorphic vesicles were either large or small. Using measurements of the cross-sectional diameter of dendritic microtubules, samples of digitized axonal endings from normal and experimental cases were normalized and could be compared directly to the groups defined by the cluster analysis. Microtubules were 21.7 nm (+/- 1.6) in average diameter. After injections of 3H-leucine and/or proline in the dorsal cochlear nucleus, most of the labeled endings in the IC contained small, round vesicles (less than 47 nm in diameter) although a very small number of endings with large, round vesicles also were labeled.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed2003

Olivocochlear collaterals evoke excitatory effects in onset neurones of the rat cochlear nucleus.

Axons of medial olivocochlear neurones in the superior olivary complex terminate on the outer hair cells of the cochlea and also give off collaterals that terminate in the cochlear nucleus. Previous work in our laboratory, using extracellular recordings in the cochlear nucleus, has indicated that stimulation of the olivocochlear axons may have an excitatory effect on specific cell populations of the cochlear nucleus, such as onset-choppers, in contrast to the peripheral suppressive action of the same axons. We have investigated whether this excitation is produced by action of the olivocochlear collaterals in the cochlear nucleus or whether it is mediated via the peripheral suppression, by measuring intracellular responses in the rat cochlear nucleus to electrical stimulation of the olivocochlear axons in silence. The results demonstrate that single shocks applied to the olivocochlear axons can evoke excitatory postsynaptic potentials in onset neurones. We observed an inhibitory effect in one chopper only. In the same animals in all other neurones investigated (i.e. three primary-like neurones and eight choppers) the same stimulation was without any effect on cell membrane potential. We conclude that the excitatory effects in onset neurones are not caused by suppression in the auditory peripheral organ, but by activation of olivocochlear collaterals in the cochlear nucleus.

Animals↗
PubMed1993

[The effects of chronic electrical stimulation on cochlear nucleus in normal hearing kittens].

We examined the effects of long-term electrical stimulation of the auditory nerve on the morphology of neurons in the cochlear nucleus in young, normal hearing kittens. These animals received bilateral intra- and extra-cochlear implants and were stimulated unilaterally for periods of 1,000-1,500 hours. After sacrifice, cross-sectional areas of neuron somata of the cochlear nucleus were measured with an image-analysis system. There were no statistically significant difference among anteroventral cochlear nucleus (AVCN), posteroventral cochlear nucleus (PVCN) and dorsal cochlear nucleus (DCN) neurons when stimulated sides were compared with unstimulated sides. These results indicate that there were no significant effects of long-term electrical stimulation of auditory nerve on the cross-sectional areas of neuron somata of AVCN, PVCN and DCN in normal hearing kittens.

Animals↗
PubMed1996

Immunolocalization of muscarinic acetylcholine subtype 2 receptors in rat cochlear nucleus.

It has been suggested that cholinergic effects in the rat cochlear nucleus (CN) are mediated by muscarinic acetylcholine receptors. In this study, immunohistochemistry for muscarinic subtype 2 (m2) receptors using a monoclonal subtype-specific antibody (Levey et al. [1995] J. Comp. Neurol. 351:339-356) revealed an m2-like system in the rat CN. A prominent lamina of m2-immunoreactive fibers and puncta was located in a subgranular layer of the caudal anteroventral cochlear nucleus (AVCN) and the posteroventral cochlear nucleus (PVCN). The superficial granular layer of the rostral AVCN and the medial sheet region also contained notable immunoreactivity for m2. Some labeled somata and their processes were found in magnocellular regions of the ventral CN. A network of neurites and puncta was located in the fusiform soma and deep layers of the dorsal CN. The olivocochlear bundle and its branches to the CN were also m2 immunoreactive and possibly contributed m2-labeled fibers and terminals to the CN. Some similarities and some differences were found between this m2 receptor distribution pattern and previous results for choline acetyltransferase (ChAT), acetylcholinesterase (AChE), and muscarinic acetylcholine receptor immunohistochemistry and binding in the CN. The results suggest that m2 receptors that are located both pre- and postsynaptically mediate many cholinergic effects in the rat CN.

Acetylcholinesterase↗
PubMed1985

Classification of response patterns in cochlear nucleus of barn owl: correlation with functional response properties.

Response patterns of neurons in the cochlear nuclei of the barn owl (Tyto alba) were studied by obtaining poststimulus time histograms (PSTHs) and interspike interval histograms for the response to short tone bursts at the neuron's characteristic frequency. The observed response patterns can be classified according to the scheme developed for neurons of the mammalian cochlear nuclear complex (22). Neurons of the magnocellular cochlear nucleus (n. magnocellularis), which respond in a phase-locked manner to sinusoidal signals and do not show large increases in spike discharge rate with changes in stimulus intensity (26), have "primarylike" (PSTH) discharge patterns and broad interspike interval histograms. This indicates that magnocellular neurons have irregular firing patterns, with the timing of individual spikes being dependent on the phase of the stimulus waveform. Neurons of the angular cochlear nucleus (n. angularis), which show little or no phase-locking and large increases in spike rate with increasing intensity (26), had almost exclusively "transient chopper" discharge patterns. The interspike interval histograms of these angular units are sharp, indicating that their discharge is very regular. At the onset of the response where the chopper pattern is observed, both discharge regularity and rate-intensity sensitivity are at their maximum levels. Several "onset" units were isolated in the angular cochlear nucleus, but no "pauser" or "buildup" units were seen. Also, all of the units in the angular nucleus had monotonic rate-intensity functions. Thus no neural response patterns typical of mammalian dorsal cochlear nucleus units were observed. The relationship of response pattern type to neural function is discussed in relation to the acoustic cues used by the owl for two-dimensional sound localization. The primarylike, phase-locked discharge of magnocellular units is undoubtedly involved in the analysis of interaural differences in stimulus phase, which the owl uses for horizontal localization. There is strong evidence suggesting that the angular nucleus is involved in processing stimulus intensity information, which is important for determining sound elevation (due to asymmetries in vertical directionality of the owl's external ears). The predominant chopper patterns seen in the angular nucleus suggest that in the owl, this response type is correlated with stimulus intensity processing. Similarities in both anatomy and physiology suggest that the magnocellular nucleus is analogous to the spherical cell or bushy cell population of the anterior division of the mammalian anteroventral cochlear nucleus.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1987

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.

Animals↗
PubMed1998

Synaptic inputs to stellate cells in the ventral cochlear nucleus.

Auditory information is carried from the cochlear nuclei to the inferior colliculi through six parallel ascending pathways, one of which is through stellate cells of the ventral cochlear nuclei (VCN) through the trapezoid body. To characterize and identify the synaptic influences on T stellate cells, intracellular recordings were made from anatomically identified stellate cells in parasagittal slices of murine cochlear nuclei. Shocks to the auditory nerve consistently evoked five types of synaptic responses in T stellate cells, which reflect sources intrinsic to the cochlear nuclear complex. 1) Monosynaptic excitatory postsynaptic potentials (EPSPs) that were blocked by 6,7-dinitroquinoxaline-2,3-dione (DNQX), an antagonist of alpha-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptors, probably reflected activation by auditory nerve fibers. Electrophysiological estimates indicate that about five auditory nerve fibers converge on one T stellate cell. 2) Disynaptic, glycinergic inhibitory postsynaptic potentials (IPSPs) arise through inhibitory interneurons in the VCN or in the dorsal cochlear nucleus (DCN). 3) Slow depolarizations, the source of which has not been identified, that lasted between 0.2 and 1 s and were blocked by -2-amino-5-phosphonovaleric acid (APV), the N-methyl-D-aspartate (NMDA) receptor antagonist. 4) Rapid, late glutamatergic EPSPs are polysynaptic and may arise from other T stellate cells. 5) Trains of late glycinergic IPSPs after single or repetitive shocks match the responses of D stellate cells, showing that D stellate cells are one source of glycinergic inhibition to T stellate cells. The source of late, polysynaptic EPSPs and IPSPs was assessed electrophysiologically and pharmacologically. Late synaptic responses in T stellate cells were enhanced by repetitive stimulation, indicating that the interneurons from which they arose should fire trains of action potentials in responses to trains of shocks. Late EPSPs and late IPSPs were blocked by APV and enhanced by the removal of Mg2+, indicating that the interneurons were driven at least in part through NMDA receptors. Bicuculline, a gamma-aminobutyric acid-A (GABAA) receptor antagonist, enhanced the late PSPs, indicating that GABAergic inhibition suppresses both the glycinergic interneurons responsible for the trains of IPSPs in T-stellate cells and the interneuron responsible for late EPSPs in T stellate cells. The glycinergic interneurons that mediate the series of IPSPs are intrinsic to the ventral cochlear nucleus because long series of IPSPs were recorded from T stellate cells in slices in which the DCN was removed. These experiments indicate that T stellate cells are a potential source of late EPSPs and that D stellate cells are a potential source for trains of late IPSPs.

Action Potentials↗