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PubMed1995

Fiber outgrowth and pathfinding in the developing auditory brainstem.

In order to study the initial outgrowth of cochlear nucleus fibers, the lipophilic carbocyanine dye DiI was used to label these neurites in rats at E13 and E15. At E13, prior to leaving the ventricular zone, cochlear neurons have already extended axons along the marginal edge of the hindbrain and a few fibers have reached the midline. By E15, many more fibers have entered the midline region and some appear to contact glial cells at the midline. Based on axon trajectory and growth cone morphology it does not appear likely that there are significant intermediate decision points for trapezoid body fibers at these early stages of development. Contact between growth cones and glial cells at the midline is consistent with these cells playing a role in providing guidance cues for developing auditory fibers.

Animals↗
PubMed1994

Loudness-coding mechanisms inferred from electric stimulation of the human auditory system.

Two distinct physiological mechanisms underlying loudness sensation were inferred from electric stimulation of the human auditory nerve and brainstem. In contrast to a power function relating loudness and stimulus intensity in acoustic hearing, loudness in electric stimulation of the auditory nerve depends on stimulus frequency. Loudness is an exponential function of electric amplitude for high frequencies and is a power function for low frequencies. A frequency-dependent, two-stage model is suggested to explain the loudness function, in which the first stage of processing is performed by a mechanical mechanism in the cochlea for high-frequency stimuli and by a neural mechanism in the cochlear nucleus for low-frequency stimuli.

Acoustic Stimulation↗
PubMed2000

AMPA receptor binding in adult guinea pig brain stem auditory nuclei after unilateral cochlear ablation.

This study determined if an asymmetric hearing loss, due to unilateral cochlear ablation, could induce the regulation of intracellular AMPA receptors in brain stem auditory nuclei. In young adult guinea pigs, the high-affinity specific binding of [(3)H]AMPA was measured in the cochlear nucleus (CN), the superior olivary complex (SOC), and the auditory midbrain at 2-147 postlesion days. After correction for tissue shrinkage, changes in specific binding relative to that in age-matched unlesioned controls were interpreted as altered numbers and/or activity of intracellular AMPA receptors. In the CN, transient elevations and/or deficits in binding were evident in most regions, which usually recovered by 147 days. However, persistently deficient binding was evident ipsilaterally in the anterior part of the anteroventral CN (AVCNa). In the SOC, transient elevations in binding were evident at 2 days in the medial limb of the lateral superior olive (LSOmed) and the medial superior olive. Between 7 and 147 days, most SOC nuclei exhibited transient, temporally synchronized postlesion deficits in binding. However, late in the survival period, deficits persisted ipsilaterally in the LSOmed and the lateral (LSOlat) limb of the lateral superior olive. In the midbrain, transient elevations and/or deficits in binding were evident in the dorsal nucleus of the lateral lemniscus as well as in the central and dorsal nucleus of the inferior colliculus. A persistent deficit was evident in the intermediate nucleus of the lateral lemniscus. The findings implied that auditory neurons contain regulatory mechanisms that control the numbers and/or activity of intracellular AMPA receptors. Regulation was induced by cochlear nerve destruction and probably by changes in the excitation of glutamatergic neurons. Many of the regulatory changes were transient, except in the ipsilateral AVCNa and LSO, where postlesion downregulations were persistent. The downregulation in the ipsilateral AVCNa was probably induced directly by the loss of cochlear nerve endings. However, other regulatory changes may have been induced by signals carried on pathways emerging from the ipsilateral CN and on centrifugal auditory pathways.

Animals↗
PubMed1987

Sources of projections to subdivisions of the inferior colliculus in the rat.

Brainstem and forebrain projections to major subdivisions of the rat inferior colliculus were studied by using retrograde and anterograde transport of horseradish peroxidase. Retrograde label from injection into the external cortex of the inferior colliculus appears bilaterally in cells of the inferior colliculus, as well as in other brainstem auditory groups including the ipsilateral dorsal nucleus of the lateral lemniscus and contralateral dorsal cochlear nucleus. The external cortex is the only collicular subdivision where an injection labels cells in the contralateral cuneate nucleus, gracile nucleus, and spinal trigeminal nucleus. Other projecting cells to the external cortex are found in the lateral nucleus of substantia nigra, the parabrachial region, the deep superior colliculus, the midbrain central gray, the periventricular nucleus, and area 39 of auditory cortex. Injection of the dorsal cortex of inferior colliculus heavily labels pyramidal cells of areas 41, 20, and 36 of the ipsilateral neocortex. Anterograde label from a large injection of auditory cortex is densely distributed in the dorsal cortex, lesser so in the external cortex, and only slightly in the central nucleus. Labelled cells appear in the central nucleus, dorsal cortex, and external cortex, primarily ipsilaterally, following dorsal cortex injection. Relatively few cells from other brainstem auditory groups show projections to the dorsal cortex. Injection of the central nucleus of the inferior colliculus results in robust labelling of nuclei of the ascending auditory pathway including the anteroventral, posteroventral, and dorsal cochlear nuclei (mainly contralaterally), and bilaterally the lateral superior olive, lateral nucleus of the trapezoid body, dorsal nucleus of the lateral lemniscus, and the central nucleus, dorsal cortex, and external cortex of the colliculus. The medial superior olive, superior paraolivary nucleus, and ventral nucleus of the trapezoid body essentially show ipsilateral projections to the central nucleus. The differential distribution of afferents to the inferior colliculus provides a substrate for functional parcellation of collicular subdivisions.

Animals↗
PubMed1997

Speech recognition at simulated soft, conversational, and raised-to-loud vocal efforts by adults with cochlear implants.

Ten postlinguistically deaf adults who used the Nucleus Cochlear Implant System and SPEAK speech coding strategy responded to vowels, consonants, words, and sentences presented sound-only at 70, 60, and 50 dB sound-pressure level. Highest group mean scores were at a raised-to-loud level of 70 dB for consonants (73%), words (44%), and sentences (87%); the highest score for vowels (70%) was at a conversational level of 60 dB. Lowest group mean scores were at a soft level of 50 dB for vowels (56%), consonants (47%), words (10%), and sentences (29%); all except subject 7 had some open-set speech recognition at this level. For the conversational level (60 dB), group mean scores for sentences and words were 72% and 29%, respectively. With this performance and sound-pressure level, it was observed that these subjects communicated successfully in a variety of listening situations. Given these subjects' speech recognition scores at 60 dB and the fact that 70 dB does not simulate the vocal effort used in everyday speaking situations, it is suggested that cochlear implant candidates and implantees be evaluated with speech tests presented at 60 dB instead of the customary 70 dB sound-pressure level to simulate benefit provided by implants in everyday life. Analysis of individuals' scores at the three levels for the four speech materials revealed different patterns of speech recognition among subjects (e.g., subjects 1 and 5). Future research on the relation between stimuli, sound processing, and subjects' responses associated with these different patterns may provide guidelines to select parameter values with which to map incoming sound onto an individual's electrical dynamic range between threshold and maximum acceptable loudness level to improve speech recognition.

Adult↗
PubMed2002

Conductive hearing loss results in changes in cytochrome oxidase activity in gerbil central auditory system.

Conductive hearing loss (CHL) restricts auditory input to an intact peripheral auditory system. Effects of deprivation on the central auditory system (CAS) have been debated, although a number of studies support the hypothesis that CHL can cause modification of CAS structure and function. The present study was designed to test the hypothesis that unilateral CHL results in a decrease in cytochrome oxidase (CO) activity in CAS nuclei that receive major afferent input from the affected ear. Gerbils at postnatal day 12 (P21) or 6-8 weeks underwent left unilateral CHL (malleus removal), cochlear ablation, or a sham surgical procedure. After a survival time of 48 hours or 3 weeks, animals were sacrificed and tissue was processed for cytochrome oxidase histochemistry. Optical density (OD) measurements were made from individual neurons in the anteroventral cochlear nucleus (AVCN) and from medial and lateral dendritic fields in the medial superior olivary nucleus (MSO), the lateral superior olivary nucleus, and the inferior colliculus. The width of the CO-stained neuropil in MSO was also measured as an estimate of dendritic length. OD measures were corrected to neutral areas of the brain. Cochlear ablation caused significant decreases in CO activity in left lower brainstem nuclei, particularly in adult animals. Following CHL, a significant decrease in CO activity was observed in the ipsilateral AVCN and a significant increase was observed in the contralateral AVCN. Cochlear ablation resulted in decreased width of MSO neuropil containing dendrites that receive primary input from the ablated ear. CHL resulted in a significant increase in the width of MSO neuropil on both sides of the brain in the P21 animals that survived 3 weeks but not in P21 animals that survived only 48 hours or in the adult animals. Unilateral CHL is associated with changes in CO activity in the AVCN and may affect MSO dendritic length in younger animals.

Animals↗
PubMed2005

Morphologic and neurochemical abnormalities in the auditory brainstem of the genetically epilepsy-prone hamster (GPG/Vall).

PURPOSE: This study was performed to evaluate whether audiogenic seizures, in a strain of genetically epilepsy-prone hamsters (GPG/Vall), might be associated with morphologic alterations in the cochlea and auditory brainstem. In addition, we used parvalbumin as a marker of neurons with high levels of activity to examine changes within neurons. METHODS: Cochlear histology as well as parvalbumin immunohistochemistry were performed to assess possible abnormalities in the GPG/Vall hamster. Densitometry also was used to quantify levels of parvalbumin immunostaining within neurons and fibers in auditory nuclei. RESULTS: In the present study, missing outer hair cells and spiral ganglion cells were observed in the GPG/Vall hamster. In addition, an increase was noted in the size of spiral ganglion cells as well as a decrease in the volume and cell size of the cochlear nucleus (CN), the superior olivary complex nuclei (SOC), and the nuclei of the lateral lemniscus (LL) and the inferior colliculus (IC). These alterations were accompanied by an increase in levels of parvalbumin immunostaining within CN, SOC, and LL neurons, as well as within parvalbumin-immunostained fibers in the CN and IC. CONCLUSIONS: These data are consistent with a cascade of atrophic changes starting in the cochlea and extending along the auditory brainstem in an animal model of inherited epilepsy. Our data also show an upregulation in parvalbumin immunostaining in the neuropil of the IC that may reflect a protective mechanism to prevent cell death in the afferent sources to this nucleus.

Animals↗
PubMed1996

Monitoring auditory function during operations to remove acoustic tumors.

The use of auditory evoked potentials in the preservation of hearing is reviewed, and the feasibility of using brainstem auditory evoked potentials, electrocochleography, and compound action potentials directly recorded from the eighth nerve is discussed. The method of recording directly from the surface of the cochlear nucleus using an electrode placed in the lateral recess of the fourth ventricle is described, and the advantages of its use are demonstrated.

Audiometry, Evoked Response↗
PubMed1997

Input from the medial nucleus of trapezoid body to an interaural level detector.

The medial nucleus of the trapezoid body (MNTB) contains components of a neural network that functions as an interaural level difference (ILD) detector. In the cat, lateral superior olivary (LSO) neurons compare the contralateral inhibitory input from the MNTB with an excitatory input form the ipsilateral anteroventral cochlear nucleus to extract information about binaural stimuli. To better specify the inhibitory inputs to the LSO and gain a better understanding of the inhibitory component of the LSO network, the response characteristics of MNTB neurons were examined in cats under stimulus conditions similar to those used to study LSO inhibitory responses. The inhibitory tuning curves of LSO units were wider than the tuning curves of MNTB units. Hence, MNTB neurons with similar, but not identical, characteristic frequencies converge to provide inhibitory input to single LSO neurons. Variations in the number of converging MNTB inputs produced a range of LSO excitatory-inhibitory threshold differences, thus creating a coding mechanism for representing the ILD. Convergence of MNTB inputs also increased the dynamic range over which contralateral stimulus level effects LSO binaural responses beyond the dynamic ranges of individual MNTB units, thus expanding the ILD range encoded by the LSO network. The differences between the first-spike latencies of MNTB and LSO tone burst responses were small and the precision of the LSO first-spike discharges was significantly greater than that of MNTB units. As tone bursts delivered simultaneously to the two ears can consistently inhibit LSO first-spike discharges, the inhibitory input must match the LSO precision by converging a number of the more variably timed MNTB discharges. Because of their precision LSO first-spike discharges may be used to encode interaural time-of-arrival differences of mid- to high-frequency transients. These findings add to the foundation for a comprehensive network model that describes the inputs to the LSO as point processes, delimits the biophysical mechanisms underlying excitatory and inhibitory interactions at the single neuron level, and reveals how these inputs determine the response to different binaural stimulus conditions.

Acoustic Stimulation↗
PubMed2005

Reconnecting neuronal networks in the auditory brainstem following unilateral deafening.

When we disturbed the auditory input of the adult rat by cochleotomy or noise trauma on one side, several substantial anatomical, cellular, and molecular changes took place in the auditory brainstem. We found that: (1) cochleotomy or severe noise trauma both lead to a considerable increase of immunoreactivity of the growth-associated protein GAP-43 in the ventral cochlear nucleus (VCN) of the affected side; (2) the expression of GAP-43 in VCN is restricted to presynaptic endings and short fiber segments; (3) axon collaterals of the cholinergic medial olivocochlear (MOC) neurons are the path along which GAP-43 reaches VCN; (4) partial cochlear lesions induce the emergence of GAP-43 positive presynaptic endings only in regions tonotopically corresponding to the extent of the lesion; (5) judging from the presence of immature fibers and growth cones in VCN on the deafened side, at least part of the GAP-43 positive presynaptic endings appear to be newly formed neuronal contacts following axonal sprouting while others may be modified pre-existing contacts; and (6) GAP-43 positive synapses are formed only on specific postsynaptic profiles, i.e., glutamatergic, glycinergic and calretinin containing cell bodies, but not GABAergic cell bodies. We conclude that unilateral deafening, be it partial or total, induces complex patterns of reconnecting neurons in the adult auditory brainstem, and we evaluate the possibility that the deafness-induced chain of events is optimized to remedy the loss of a bilaterally balanced activity in the auditory brainstem.

Animals↗
PubMed1997

Axogenesis in the human fetal auditory system, demonstrated by neurofilament immunohistochemistry.

Morphological, electrophysiological and behavioral evidence indicates that the onset of rapid, synchronized conduction of auditory impulses occurs in the human brainstem during the 28th-29th fetal weeks. This implies that axonal connections in the brainstem auditory pathway are generated prior to this time. In order to investigate the sequence of axogenic events in the human brainstem pathway, we employed immunohistochemical techniques and an antibody to neurofilament protein. Immunostaining for axonal neurofilaments in an age-graded series of fetal brains demonstrates that a small number of cochlear nerve axons have invaded the ventral cochlear nucleus by the 16th fetal week. By this same time point, a limited number of trapezoid body-lateral lemniscus axons have reached the superior olivary complex and inferior colliculus. Between gestational weeks 16 and 26, there is marked expansion and collateralization of the ascending pathway from cochlear nerve to inferior colliculus. By week 26, ascending axons have begun to form plexuses of terminal neuropil within all of the brainstem auditory nuclei. Beginning in week 22, there is development of commissural axons (dorsal commissure of the lateral lemniscus and commissure of the inferior colliculus) and descending projections (descending collicular axons and olivocochlear bundle). This early establishment of a mature pattern of axonal connections presumably forms the basis for the appearance of myelin, acousticomotor reflexes and recordable brainstem responses by fetal week 29.

Auditory Pathways↗
PubMed1997

Neuronal and transneuronal degeneration of auditory axons in the brainstem after cochlear lesions in the chinchilla: cochleotopic and non-cochleotopic patterns.

Terminal axonal degeneration in the brain following cochlear lesions was studied with the Nauta-Rasmussen method. Losses of hair cells and myelinated cochlear fibers were assessed. The cochleotopic map projected, from apex to base, on the ventral-to-dorsal axes of the cochlear nuclei. The cochleotopic correspondence was better for loss of cochlear nerve fibers and inner hair cells, than for outer hair cells. Cochlear fibers were traced to all parts of the cochlear nucleus, including the small-cell shell, also to cell-group Y and the flocculus. Terminal axonal degeneration in nuclei of the superior olivary complex, lateral lemniscus, and inferior colliculus was interpreted as transynaptic, since degenerated axons could not be traced to these locations from the cochlear nerve or trapezoid body. Moreover, biotinylated dextran amine injection in the basal turn of scala media of a normal cochlea labeled cochlear nerve fibers projecting to the high-frequency regions of the cochlear nuclei and to the flocculus, but not to more central auditory nuclei. This is the first detailed account of transynaptic degeneration in the ascending auditory pathway resulting from cochlear damage in an adult mammal. These findings are consistent with a dystrophic process depending on hair-cell loss and/or direct damage to cochlear nerve fibers.

Animals↗
PubMed2002

The medial nucleus of the trapezoid body in the gerbil is more than a relay: comparison of pre- and postsynaptic activity.

The medial nucleus of the trapezoid body (MNTB) plays an important role in the processing of interaural intensity differences, a feature that is critical for the localization of sound sources. It is generally believed that the MNTB functions primarily as a passive relay in converting excitatory input originating from the contralateral cochlear nucleus (CN) into an inhibitory input to the ipsilateral lateral superior olive. However, studies showing that the MNTB itself is also the target of inhibitory input suggest that the MNTB may serve more than a sign-converting function. To examine the fidelity of signal transmission at the CN-MNTB synapse, presynaptic calyceal potentials ("prepotentials"), reflecting the excitatory input to the MNTB neuron, and postsynaptic action potentials were simultaneously monitored with the same electrode during in vivo extracellular recordings from the gerbil's MNTB. Presynaptic activity differed from postsynaptic activity in several respects: (1) Spontaneous and sound-evoked discharge rates were greater presynaptically than postsynaptically. (2) Frequency tuning was sharper postsynaptically than presynaptically. (3) Calyceal terminals and MNTB neurons both showed phasic-tonic response patterns to tonal stimulation, but the duration of the onset response and the level of the tonic component were reduced postsynaptically. (4) Phase-locking to sound frequencies up to 1 kHz was greater postsynaptically than presynaptically. (5) The rate-intensity characteristics of pre- and postsynaptic activities differed significantly from each other in half of the MNTB neurons. To test the hypothesis that acoustically evoked inhibition of MNTB neurons contributed to the relatively lower levels of postsynaptic discharge, two-tone stimulation was applied, wherein the response to one tone-burst, set at the neuron's characteristic frequency, can be reduced by addition of a second "inhibitory" tone. The inhibitory tone caused a much larger reduction in post- than in presynaptic activity, indicating an acoustically evoked inhibitory influence directly on MNTB units. These findings show that transmission at the CN-MNTB synapse does not occur in a fixed one-to-one manner and that the response of MNTB neurons reflects the integration of their excitatory and inhibitory inputs.

Action Potentials↗
PubMed1991

Central nervous system changes associated with noise-induced hearing loss: an electron microscopic and freeze-fracture study of the chick nucleus magnocellularis.

This investigation assessed the effects of noise-induced hearing loss on the avian nucleus magnocellularis, the homologue of the mammalian cochlear nucleus. Fifteen-day-old chicks were exposed to high-intensity broad-band noise (132 dBA) for 24 to 72 hours. Hearing loss was documented with auditory brain-stem response recordings and scanning electron microscopy of the basilar papilla. Nucleus magnocellularis was assessed with thin-section electron microscopy and freeze-fracture techniques. Freeze-fracture preparations of noise-exposed animals showed a decrease in synaptic vesicle fusion sites at the synaptic active zone, a decrease in re-uptake of presynaptic membrane by coated vesicles, and a shift of the preferential fracture plane at the synapse from the presynaptic membrane to the post-synaptic membrane. These findings support the hypothesis that the effects of noise are not limited to the auditory periphery, but that transneuronal changes extend into the auditory central nervous system.

Animals↗
PubMed2002

Altered tone-induced Fos expression in the mouse inferior colliculus after early exposure to intense noise.

Mice become highly susceptible to audiogenic seizures (AGS) after being exposed to intense, high-frequency noise during a critical period of early life (priming). To determine the critical site for AGS priming in the auditory brainstem, animals in the experimental group were primed at 21 days, and the tone-induced Fos immunoreactivity was examined 1, 7, and 14 days after priming as an index of excitability of neurons. Enhanced Fos immunoreactivity was observed in the inferior colliculus (IC) of the primed mice 7 and 14 days after priming as compared to that of non-primed mice and attenuated Fos expression was observed 1 day after priming. No significant elevation of Fos expression was observed in the cochlear nucleus and the deep layer of the superior colliculus of either type of mice. These results strongly suggest that the IC is the target site of AGS priming.

Acoustic Stimulation↗
PubMed2000

Differential expression of voltage-gated potassium channel genes in auditory nuclei of the mouse brainstem.

Voltage-gated potassium (Kv) channels may play an important role in the encoding of auditory information. Towards understanding the roles of Shaker and Shaw-like channels in this process, we examine here the expression of Kv1.1, Kv1.2, Kv3.1, and Kv3.3 in the central auditory nuclei of the mouse using quantitative in situ hybridization techniques. We establish rank order for each channel's expression in each region, finding that the medial nucleus of the trapezoid body shows the highest signal for each of the four channel genes. In other auditory nuclei differential expression is found among and between members of both Shaker and Shaw subfamilies. Of particular interest is the stark contrast between high level expression of Kv1.1 and very low level expression of Kv3.1 in the octopus cell area of the cochlear nucleus and in the lateral superior olivary nucleus. These unique expression patterns suggest that Kv channel gene expression is regulated to allow brainstem auditory neurons to transmit temporally patterned signals with high fidelity. In instances where specific cell types can be tentatively identified, we discuss the possible contribution made by these channel genes to the physiological properties of those neurons.

Animals↗
PubMed1996

Immunohistochemical mapping of perineuronal nets containing chondroitin unsulfated proteoglycan in the rat central nervous system.

Subsets of neurons ensheathed by perineuronal nets containing chondroitin unsulfated proteoglycan have been immunohistochemically mapped throughout the rat central nervous system from the olfactory bulb to the spinal cord. A variable proportion of neurons were outlined by immunoreactivity for the monoclonal antibody (Mab 1B5), but only after chondroitinase ABC digestion. In forebrain cortical structures the only immunoreactive nets were around interneurons; in contrast, throughout the brainstem and spinal cord a large proportion of projection neurons were surrounded by intense immunoreactivity. Immunoreactivity was ordinarily found in the neuropil between neurons surrounded by an immunopositive net. By contrast, within the pyriform cortex the neuropil of the plexiform layer was intensely immunoreactive even though no perineuronal net could be found. The presence of perineuronal nets could not be correlated with any single class of neurons; however a few functionally related groups (e.g., motor and motor-related structures: motor neurons both in the spinal cord and in the efferent somatic nuclei of the brainstem, deep cerebellar nuclei, vestibular nuclei; red nucleus, reticular formation; central auditory pathway: ventral cochlear nucleus, trapezoid body, superior olive, nucleus of the lateral lemniscus, inferior colliculus, medial geniculate body) were the main components of the neuronal subpopulation displaying chondroitin unsulfated proteoglycans in the surrounding extracellular matrix. The immunodecorated neurons found in the present study and those shown by different monoclonal antibodies or by lectin cytochemistry, revealed consistent overlapping of their distribution patterns.

Animals↗
PubMed1985

The localization and origin of somatostatin-containing fibers in an auditory brainstem nucleus.

Immunoreactive perikarya and terminals have been visualized in the rat brainstem using antibodies raised against somatostatin 14 and somatostatin 28 (4-14) in a peroxidase antiperoxidase procedure. Labeling was observed in several areas of the brainstem, including the solitary nucleus, the dorsal lateral lemniscus, the ventral cochlear nucleus, several areas of the reticular formation, the nucleus of the spinal trigeminal tract, and the locus ceruleus. There was no obvious difference obtained with antisera against SS 14 and SS 28 (4-14). Several auditory areas were found to contain SS in discrete sub-groups of perikarya and terminals, suggesting the existence of one or more specific SS-containing auditory pathways. The origin of the SS-positive terminals in the medial nucleus of the trapezoid body was investigated using thermal and chemical lesions. It was concluded that these terminals originate from a specific SS-positive group of perikarya located either in or adjacent to the medial portion of the nucleus of the dorsal lateral lemniscus.

Animals↗