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Auditory training improves neural timing in the human brainstem.

The auditory brainstem response reflects neural encoding of the acoustic characteristic of a speech syllable with remarkable precision. Some children with learning impairments demonstrate abnormalities in this preconscious measure of neural encoding especially in background noise. This study investigated whether auditory training targeted to remediate perceptually-based learning problems would alter the neural brainstem encoding of the acoustic sound structure of speech in such children. Nine subjects, clinically diagnosed with a language-based learning problem (e.g., dyslexia), worked with auditory perceptual training software. Prior to beginning and within three months after completing the training program, brainstem responses to the syllable /da/ were recorded in quiet and background noise. Subjects underwent additional auditory neurophysiological, perceptual, and cognitive testing. Ten control subjects, who did not participate in any remediation program, underwent the same battery of tests at time intervals equivalent to the trained subjects. Transient and sustained (frequency-following response) components of the brainstem response were evaluated. The primary pathway afferent volley -- neural events occurring earlier than 11 ms after stimulus onset -- did not demonstrate plasticity. However, quiet-to-noise inter-response correlations of the sustained response ( approximately 11-50 ms) increased significantly in the trained children, reflecting improved stimulus encoding precision, whereas control subjects did not exhibit this change. Thus, auditory training can alter the preconscious neural encoding of complex sounds by improving neural synchrony in the auditory brainstem. Additionally, several measures of brainstem response timing were related to changes in cortical physiology, as well as perceptual, academic, and cognitive measures from pre- to post-training.

Attention↗

Electrophysiologic studies of the auditory cortex in the awake monkey.

An overview of some recent developments in the study of central auditory processes is presented. We describe findings from single cell physiologic studies that contribute to our understanding of central auditory function, the development of behavioral techniques permitting precise evaluation of hearing in animals, and the power and potential of integrative neurophysiologic-behavioiral investigations in defining and analyzing central neural mechanisms that underlie normal perception and imperception. The influence of changes in intermodality attention on evoked activity of 25 cells of the auditory cortex is described and compared to effects of other attentional changes. A small but consistent increase in excitatory cell evoked discharge rate and a reduction in initial latency of response are shown to be correlated with a shift from a visual to an auditory task. In general, behavioral states ranging from sleep-waking to attention to a specific acoustic cue are shown to influence responsiveness of the auditory system and variability of response. Procedures are described for effective control of attentional factors and reduction of electrophysiologic variability. The contributions of these data and procedures to studies of the encoding of complex auditory signals and studies of neural mechanisms underlying such disorders as noise induced hearing loss are discussed.

Animals↗

40-Hz steady state response in awake cats after bilateral chronic lesions in auditory cortices or inferior colliculi.

To reveal the generators of 40-Hz steady state response, the following investigations in awake cat were performed: (i) analyses of 40-Hz SSR without any lesions, and study of stimulus rate; (ii) effect of chronic lesions in the bilateral auditory cortex; (iii) effect of chronic lesions in the bilateral inferior colliculus. The maximum amplitudes were obtained at stimulus rates of 40-50 Hz. In cat with lesions in the bilateral auditory cortex, the wave configurations and the phase were retained but the amplitude was reduced to 90%. In cat with lesions in the inferior colliculus, the wave configurations and the phase were retained but the amplitude was reduced to 60%. These data suggest the following conclusions: (i) cat was a suitable animal model for human's 40-Hz SSR; (ii) the auditory cortex could be one of the generators of 40-Hz SSR; (iii) the inferior colliculus could be an important generator of 40-Hz SSR; (iv) 40-Hz SSR seems to be generated through not only primary auditory pathways, but also nonspecific auditory pathways, which receive auditory information in parallel from the brainstem, thalamus, and cerebrum.

Animals↗

Peripheral auditory asymmetry in infantile autism.

Difficulty in filtering relevant auditory information in background noise is one of the features of autism. Auditory filtering processes can be investigated at the peripheral level as they are hypothesized to involve active cochlear mechanisms which are regulated by the efferent activity of the medial olivocochlear (MOC) system. The aim of the present work was therefore to assess these peripheral auditory processes in 22 children and adolescents with autism compared with age- and gender-matched normal controls. Active cochlear mechanisms were evaluated with transiently evoked otoacoustic emissions (TEOAEs) and MOC system efficiency was assessed via TEOAEs which are decreased when stimulating the contralateral ear with noise. The MOC system evaluation was performed on 18 of the 22 children. In both studies, results were analysed according to age (from 4 to 10 years and from 11 to 20 years). The main result concerns the asymmetry of the efferent system which differs in individuals with autism. Several neural processes might be hypothesized as involved in the results obtained as the MOC system which originates in the brainstem received regulating controls from upper brain structures including auditory cortex. Lateralization abnormalities at the auditory periphery may reflect indirectly a problem at a higher level of auditory processing. A second important result shows a decrease in TEOAE amplitude with age, in patients, that may correspond to a decrease in hearing sensitivity.

Adolescent↗

A case study of pure word deafness: modularity in auditory processing?

AL, a woman with an acquired disturbance of auditory processing beginning in the second decade, was originally diagnosed as having pure word deafness. Recent analysis with a wide range of stimuli suggests that her comprehension deficit also extends to a subset of musical and non-verbal environmental sounds. The perceptual demands of the different auditory stimuli appear to account for part of the apparent material specificity. Additionally, over the years, the presumed temporal lobe cortical pathology has been supplemented by a mild to moderate, peripheral low-frequency hearing loss and evidence of dysfunction in lower level auditory processing pathways. The current peripheral dysfunction closely resembles cases recently labeled as auditory neuropathy. The diagnosis of pure word deafness should not be based on a limited set of auditory stimuli; additionally, a careful assessment using modern audiological techniques should be performed to evaluate peripheral auditory functions.

Achievement↗

Evaluation of TEOAE and DPOAE measurements for the assessment of auditory thresholds in sensorineural hearing loss.

Transiently evoked otoacoustic emissions (TEOAE) and distortion product otoacoustic emissions (DPOAE) can be quantified concerning their amplitude and frequency. They are known to be diminished or absent in sensorineural hearing loss. It is therefore of interest how TEOAE and DPOAE correlate with the auditory threshold and whether the auditory threshold can be predicted by these variables. In a cross sectional study of 61 patients (102 ears) with sensorineural hearing loss, auditory threshold, tympanometry, stapedius reflexes, TEOAE and DPOAE were measured. Correlation coefficients of the hearing loss (0.5-6 kHz) and the amplitude of the distortion product 2f1-f2 (0.46 4 kHz) respectively TEOAE amplitude (1-4 kHz) were computed. TEOAE showed lower correlation coefficients and less frequency specificity than DPOAE. In order to increase the correlations with the auditory threshold we fitted a multivariate linear regression model with TEOAE and DPOAE simultaneously as predictors for the auditory threshold gaining 95% prediction intervals of 19-39 dB depending on the frequency investigated. By restricting the hearing loss to a maximum of 70 dB HL the 95% prediction interval of the auditory threshold could be decreased to 18-26 dB. Further improvements can be expected if the high inter- and intraindividual variability of TEOAE and particularly DPOAE measurements can be reduced. The results allow us to use TEOAE and DPOAE in addition to click-evoked brainstem audiometry in order to provide more frequency specific information about the hearing loss in newborns, which is of the utmost importance for an ideal fitting of hearing aids.

Acoustic Impedance Tests↗

Mapping auditory hallucinations in schizophrenia using functional magnetic resonance imaging.

BACKGROUND: Perceptions of speech in the absence of an auditory stimulus (auditory verbal hallucinations) are a cardinal feature of schizophrenia. Functional neuroimaging provides a powerful means of measuring neural activity during auditory hallucinations, but the results from previous studies have been inconsistent. This may reflect the acquisition of small numbers of images in each subject and the confounding effects of patients actively signaling when hallucinations occur. METHODS: We examined 6 patients with schizophrenia who were experiencing frequent auditory hallucinations, using a novel functional magnetic resonance imaging method that permitted the measurement of spontaneous neural activity without requiring subjects to signal when hallucinations occurred. Approximately 50 individual scans were acquired at unpredictable intervals in each subject while they were intermittently hallucinating. Immediately after each scan, subjects reported whether they had been hallucinating at that instant. Neural activity when patients were and were not experiencing hallucinations was compared in each subject and the group as a whole. RESULTS: Auditory hallucinations were associated with activation in the inferior frontal/insular, anterior cingulate, and temporal cortex bilaterally (with greater responses on the right), the right thalamus and inferior colliculus, and the left hippocampus and parahippocampal cortex (P<.0001). CONCLUSIONS: Auditory hallucinations may be mediated by a distributed network of cortical and subcortical areas. Previous neuroimaging studies of auditory hallucinations may have identified different components of this network.

Auditory Perception↗

Ultrastructure of synaptic contacts between identified neurons of the auditory pathway in Gryllus bimaculatus DeGeer.

The synaptic contacts between the auditory sensory cells and identified auditory interneurons ON1 and AN2 have been examined at the ultrastructural level by selective electron-dense labeling of two interneurons or of one interneuron and the sensory fibers in the same preparation. The experiments have provided the following information. The auditory afferent fibers have a monosynaptic connection with the lateral inhibitors ON1 and the ascending interneuron AN2, allowing direct activation of these interneurons. Furthermore, our work proves that the paired, lateral, inhibitor ON1 neurons have direct output synapses onto each other. The results also show that the auditory afferent axons, themselves, receive synaptic inputs just before entering the central auditory neuropil. The effects of current injection into the ON1 neuron during auditory processing indicate that these synaptic inputs onto the afferents originate, in part, from the lateral branches of the ipsilateral ON1 neuron and that they have inhibitory function. The significance of these results for auditory processing and a future perspective for electron microscopic analysis of neuropil are discussed.

Animals↗

Auditory thalamocortical projections in the cat: laminar and areal patterns of input.

Thalamocortical projections were studied in adult cats using biotinylated dextran amines, wheat germ agglutinin conjugated to horseradish peroxidase, and autoradiography with tritiated leucine and/or proline. The input from 7 architectonically defined nuclei to 14 auditory cortical fields was characterized qualitatively and quantitatively. The principal results were that 1) every thalamic nucleus projected to more than 1 field (range, 4-14 fields; mean, 7 fields); 2) only the projection from the ventral division to some primary fields (primary auditory cortex and posterior auditory cortex) had a periodic, clustered distribution, whereas the input from other divisions to nonprimary areas was continuous; 3) layers III-V received >85% of the total axonal profiles; 4) in most experiments, five or more layers were labeled; 5) the projections to nonprimary auditory areas had many laterally oriented axons; 6) the heaviest input to layer I in all experiments was usually in its upper half, suggesting a sublaminar arrangement; 7) the largest axonal trunks (up to 6 microm in diameter) arose from the medial division and ended in layer Ia, where they ran laterally for long distances; 8) there were three projection patterns: type 1 had its peak in layers III-IV with little input to layer I, and it arose from the ventral division and the dorsal superficial, dorsal, and suprageniculate nuclei of the dorsal division; type 2 had heavy labeling in layer I and less in layers III-IV, arising from the dorsal division nuclei primarily, especially the caudal dorsal and deep dorsal nuclei; and type 3 was a trimodal concentration in layers I, III-IV, and VI that originated chiefly in the medial division and had the lowest density of labeling; and 9) the quantitative profiles with the three methods were very similar. The results suggest that the subdivisions of the auditory thalamus have consistent patterns of laminar distribution to different cortical areas, that an average of five or more layers receive significant input in a specific area, that a given thalamic nucleus can influence areas as far as 20 mm apart, that the first information to arrive at the cortex may reach layer I by virtue of the giant axons, and that several laminar patterns of auditory thalamocortical projection exist. The view that the auditory thalamus (and perhaps other thalamic nuclei) serves mainly a relay function underestimates its many modes for influencing the cortex on a laminar basis.

Animals↗

Projections from auditory cortex to cochlear nucleus: A comparative analysis of rat and mouse.

Mammalian hearing is a complex special sense that involves detection, localization, and identification of the auditory stimulus. The cerebral cortex may subserve higher auditory processes by providing direct modulatory cortical projections to the auditory brainstem. To support the hypothesis that corticofugal projections are a conserved feature in the mammalian brain, this article reviews features of the rat corticofugal pathway and presents new data supporting the presence of similar projections in the mouse. The mouse auditory cortex was localized with electrophysiological recording and neuronal tracers were injected into AI. The cochlear nucleus was dissected and examined for terminal fibers by light and electron microscopy. Bouton endings were found bilaterally forming synapses with dendrites of granule cells of the cochlear nucleus. This report provides evidence for direct auditory cortex projections to the cochlear nucleus in the mouse. The distribution of projections to the granule cell domain and the synapses onto granule cell dendrites are consistent with what has been reported for rats and guinea pigs. These findings suggest a general plan for corticofugal modulation of ascending auditory information in mammals. Corticobulbar inputs to the auditory brainstem likely provided a survival advantage by improving sound detection and identification, thus allowing the development of complex social behaviors and the navigation of varied environments.

Animals↗

Visual and auditory association areas of the cat's posterior ectosylvian gyrus: cortical afferents.

In a preceding report, we described patterns of thalamic retrograde labeling following 17 tracer deposits on the cat's posterior ectosylvian gyrus and concluded, on the basis of patterns of thalamic connectivity, that the posterior ectosylvian gyrus is composed of three major divisions: a tonotopic auditory zone located anteriorly, a belt of auditory association cortex occupying the gyral crown, and a visual belt located posteriorly. We describe here patterns of transcortical retrograde labeling obtained from tracer deposits in the three zones so defined. Our results indicate that the tonotopic auditory strip is innervated primarily by axons from low-order auditory areas (AAF, AI, P, VP, and V), that the auditory belt receives its strongest input from nontonotopic auditory fields (AII, temporal cortex, and other parts of the auditory belt), and that projections to the visual belt derive primarily from extrastriate visual areas (ALLS, PLLS, DLS, 19, 20, and 21) and from association areas affiliated with the visual system (insular cortex, posterior cingulate gyrus, area 7p, and frontal cortex). We discuss the results in relation to previous systems for parcellating the posterior ectosylvian gyrus of the cat and consider the possibility that divisions of the feline posterior ectosylvian gyrus correspond directly to areas making up the superior temporal gyrus in primates.

Animals↗

Differential glycosylation of auditory and vestibular hair bundle proteins revealed by peanut agglutinin.

Up to four morphologically distinct types of cross-link are found between the stereocilia in the hair bundles of avian hair cells. These links are involved in mechanotransduction, force transmission across the bundle, and maintenance of hair bundle structure. They appear to be specialisations of the cell coat, but very little is known about their molecular composition. Chick inner ear tissues were therefore screened with a number of different lectins to find markers for specialisations of the hair bundle surface. One lectin, peanut agglutinin (PNA), which recognises the dissacharide Gal beta 1-3GalNAc, was found to be a fairly selective marker for vestibular hair bundles, but it does not stain the stereocilia of auditory hair cells. The staining patterns observed with PNA in the vestibular system closely resemble those seen with a monoclonal antibody (mab) directed against a 275 kD component of the hair cell's apical surface known as the hair-cell antigen (HCA). However, unlike PNA, the mab recognises both vestibular and auditory hair cells. A detailed comparison of the fluorescence staining patterns observed with PNA and the anti-HCA mab indicates that binding sites for both ligands spatially codistribute on the surface of vestibular hair cells. The lectin and the anti-HCA mab binding sites are both sensitive to trypsin treatment, and, with sections of the vestibular system, PNA pretreatment blocks subsequent anti-HCA mab staining. Immunoelectron microscopy of vestibular hair bundles shows that PNA and the anti-HCA mab both label a type of cross-link known as the shaft connector. This link type is present on both auditory and vestibular hair bundles but reacts with PNA only in the vestibular system. The lectin jacalin, which has greater specificity for Gal beta 1-3GalNAc than does PNA, also only labels vestibular and not auditory hair bundles. Although terminal sialic acid residues can block both PNA and jacalin binding, neuraminidase treatment does not unmask cryptic binding sites for these lectins on auditory hair cells but does reveal PNA and jacalin staining at a number of other locations in the inner ear. The results obtained with the lectins PNA and jacalin indicate that either the HCA or other components of the shaft links are differentially glycosylated in the vestibular and auditory epithelia of the bird. The functional significance for such a difference in glycosylation remains to be determined, but auditory and vestibular hair cells operate over different frequency ranges, and variations in glycosylation might confer different micromechanical properties on the hair bundles in these two systems.

Animals↗

On the development of low-level auditory discrimination and deficits in dyslexia.

Absolute auditory thresholds, frequency resolution and temporal resolution develop with age. It is still discussed whether low-level auditory performance is of clinical significance--specifically, for delayed maturation of central auditory processing. Recently, five new auditory tasks were used to study the development of low-level auditory discrimination. It was found that the development lasts up to the age of 16-18 years (on an average). Very similar tasks were now used with 432 controls and 250 dyslexic subjects in the age range of 7-22 years. For both groups the performance in one of the tasks was not related to the performance in another task indicating that the five tasks challenge independent subfunctions of auditory processing. Surprisingly high numbers of subjects were classified as low performers (LP), because they could not perform one or the other task at its easiest level and no threshold value could be assigned. For the dyslexics the incidence of LP was considerably increased in all tasks and age groups as compared with the age matched controls. The development of dynamic visual and optomotor functions and the corresponding deficits in dyslexia are discussed in relation to the auditory data presented here.

Adolescent↗

Embryonic development and evolutionary origin of the Orthopteran auditory organs.

Two different types of ears characterize the order of Orthopteran insects. The auditory organs of grasshoppers and locusts (Caelifera) are located in the first abdominal segment, those of bushcrickets and crickets (Ensifera) are found in the tibiae of the prothoracic legs. Using neuron-specific antibody labelling, we describe the ontogenetic origin of these two types of auditory organs, use comparative developmental studies to identify their segmental homologs, and on the basis of homology postulate their evolutionary origin. In grasshoppers the auditory receptors develop by epithelial invagination of the body wall ectoderm in the first abdominal segment. Subsequently, at least a part of the receptor cells undergo active migration and project their out-growing axons onto the next anterior intersegmental nerve. During this time the receptor cells and their axons express the cell-cell adhesion molecule, Fasciclin I. Similar cellular and molecular differentiation processes in neighboring segments give rise to serially homologous sensory organs, the pleural chordotonal organs in the pregenital abdominal segments, and the wing-hinge chordotonal organs in the thoracic segments. In more primitive earless grasshoppers pleural chordotonal organs are found in place of auditory organs in the first abdominal segment. In bushcrickets the auditory receptors develop in association with the prothoracic subgenual organ from a common developmental precursor. The auditory receptor neurons in these insects are homologous to identified mechanoreceptors in the meso- and metathoracic legs. The established intra- and interspecies homologies provide insight into the evolution of the auditory organs of Orthopterans.

Animals↗

Song-selective auditory input to a forebrain vocal control nucleus in the zebra finch.

Neurons in nuclei on the motor pathway for vocalizations in songbirds are known to respond to sound stimuli. The auditory responses in one such nucleus, robustus archistriatalis (RA), were characterized by making multi-unit recordings in awake and anesthetized adult male zebra finches and in birds that had received lesions of the input to RA from the lateral part of the magnocellular nucleus of the anterior neostriatum (LMAN) or the Higher Vocal Center (HVC). In awake birds, RA neurons have a high level of spontaneous activity and vigorous auditory responses to song stimuli. Significantly greater responses are seen to the bird's own song (BOS) than to BOS played in reverse (REV) or to the songs of conspecifics (CON). Under ketamine-xylazine anesthesia, spontaneous activity is reduced, response latency increases and responses to BOS, REV and CON are indistinguishable. Responses obtained under urethane anesthesia are similar to those seen in awake birds. Thus, the pattern and selectivity of auditory responses in RA depend on the animal's state. Auditory responses in RA are qualitatively unchanged following lesion of the input to RA from LMAN, indicating that this pathway is not required for the sensory processing that underlies the preference for BOS on the vocal production pathway. Our results show that an input other than that from LMAN must be primarily responsible for auditory responses in RA. The direct projection from HVC is the most likely pathway by which song selective auditory information arrives in RA, since lesioning HVC abolished auditory responses in RA.

Animals↗

Functional differentiation in the human auditory and language areas revealed by a dichotic listening task.

The human auditory cortex plays a special role in speech recognition. It is therefore necessary to clarify the functional roles of individual auditory areas. We applied functional magnetic resonance imaging (fMRI) to examine cortical responses to speech sounds, which were presented under the dichotic and diotic (binaural) listening conditions. We found two different response patterns in multiple auditory areas and language-related areas. In the auditory cortex, the medial portion of the secondary auditory area (A2), as well as a part of the planum temporale (PT) and the superior temporal gyrus and sulcus (ST), showed greater responses under the dichotic condition than under the diotic condition. This dichotic selectivity may reflect acoustic differences and attention-related factors such as spatial attention and selective attention to targets. In contrast, other parts of the auditory cortex showed comparable responses to the dichotic and diotic conditions. We found similar functional differentiation in the inferior frontal (IF) cortex. These results suggest that multiple auditory and language areas may play a pivotal role in integrating the functional differentiation for speech recognition.

Acoustic Stimulation↗

Human primary auditory cortex: cytoarchitectonic subdivisions and mapping into a spatial reference system.

The transverse temporal gyrus of Heschl contains the human auditory cortex. Several schematic maps of the cytoarchitectonic correlate of this functional entity are available, but they present partly conflicting data (number and position of borders of the primary auditory areas) and they do not enable reliable comparisons with functional imaging data in a common spatial reference system. In order to provide a 3-D data set of the precise position and extent of the human primary auditory cortex, its putative subdivisions, and its topographical intersubject variability, we performed a quantitative cytoarchitectonic analysis of 10 brains using a recently established technique for observer-independent definition of areal borders. Three areas, Te1.1, Te1.0, and Te1.2, with a well-developed layer IV, which represent the primary auditory cortex (Brodmann area 41), can be identified along the mediolateral axis of the Heschl gyrus. The cell density was significantly higher in Te1.1 compared to Te1.2 in the left but not in the right hemisphere. The cytoarchitectonically defined areal borders of the primary auditory cortex do not consistently match macroanatomic landmarks like gyral and sulcal borders. The three primary auditory areas of each postmortem brain were mapped to a spatial reference system which is based on a brain registered by in vivo magnetic resonance imaging. The integration of a sample of postmortem brains in a spatial reference system allows one to estimate the spatial variability of each cytoarchitectonically defined region with respect to this reference system. In future, the transfer of in vivo structural and functional data into the same spatial reference system will enable accurate comparisons of cytoarchitectonic maps of the primary auditory cortex with activation centers as established with functional imaging procedures.

Adult↗

Auditory and somatosensory event-related brain potentials in early blind humans.

Previous event-related potential (ERP) studies have suggested a possible participation of the visual cortex of the blind in auditory processing. In the present study, somatosensory and auditory ERPs of blind and sighted subjects were recorded when subjects were instructed to attend to stimuli of one modality and to ignore those of the other. Both modalities were stimulated with frequent ("standard") and infrequent ("deviant") stimuli, which differed from one another in their spatial locus of origin. In the sighted, deviant stimuli of the attended modality elicited N2 type of deflections (auditory N2b and somatosensory N250) over the lateral scalp areas. In contrast, in the blind, these ERP components were centroposteriorly distributed, suggesting an involvement of posterior brain areas in auditory and somatosensory stimulus discrimination. In addition, the mismatch negativity, elicited by deviant auditory stimuli even when the somatosensory stimuli were attended, was larger in the blind than in the sighted. This appears to indicate enhanced automatic processing of auditory stimulus changes in the blind. Thus, the present data suggest several compensatory changes in both auditory and somatosensory modalities after the onset of early visual deprivation.

Adult↗