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Binaural tuning of auditory units in the forebrain archistriatal gaze fields of the barn owl: local organization but no space map.

We identified a region in the archistriatum of the barn owl forebrain that contains neurons sensitive to auditory stimuli. Nearly all of these neurons are tuned for binaural localization cues. The archistriatum is known to be the primary source of motor-related output from the avian forebrain and, in barn owls, contributes to the control of gaze, much like the frontal eye fields in monkeys. The auditory region is located in the medial portion of the archistriatum, at the level of the anterior commissure, and is within the region of the archistriatum from which head saccades can be elicited by electrical microstimulation (see preceding companion article, Knudsen et al., 1995). Free-field measurements revealed that auditory sites have large, spatial receptive fields. However, within these large receptive fields, responses are tuned sharply for sound source location. Dichotic measurements showed that auditory sites are tuned broadly for frequency and that the majority are tuned to particular values of interaural time differences and interaural level differences, the principal cues used by barn owls for sound localization. The tuning of sites to these binaural cues is essentially independent of sound level. The auditory properties of units in the medial archistriatum are similar to those of units in the optic tectum, a structure that also contributes to gaze control. Unlike the optic tectum, however, the auditory region of the archistriatum does not contain a single, continuous auditory map of space. Instead, it is organized into dorsoventral clusters of sites with similar binaural (spatial) tuning. The different representations of auditory space in closely related structures in the forebrain (archistriatum) and midbrain (optic tectum) probably reflect the fact that the forebrain contributes to a wide variety of sensorimotor tasks more complicated than gaze control.

Acoustic Stimulation↗

Registration of neural maps through value-dependent learning: modeling the alignment of auditory and visual maps in the barn owl's optic tectum.

In the optic tectum (OT) of the barn owl, visual and auditory maps of space are found in close alignment with each other. Experiments in which such alignment has been disrupted have shown a considerable degree of plasticity in the auditory map. The external nucleus of the inferior colliculus (ICx), an auditory center that projects massively to the tectum, is the main site of plasticity; however, it is unclear by what mechanisms the alignment between the auditory map in the ICx and the visual map in the tectum is established and maintained. In this paper, we propose that such map alignment occurs through a process of value-dependent learning. According to this paradigm, value systems, identifiable with neuromodulatory systems having diffuse projections, respond to innate or acquired salient cues and modulate changes in synaptic efficacy in many brain regions. To test the self-consistency of this proposal, we have developed a computer model of the principal neural structures involved in the process of auditory localization in the barn owl. This is complemented by simulations of aspects of the barn owl phenotype and of the experimental environment. In the model, a value system is activated whenever the owl carries out a foveation toward an auditory stimulus. A term representing the diffuse release of a neuromodulator interacts with local pre- and postsynaptic events to determine synaptic changes in the ICx. Through large-scale simulations, we have replicated a number of experimental observations on the development of spatial alignment between the auditory and visual maps during normal visual experience, after the retinal image is shifted through prismatic goggles, and after the reestablishment of normal visual input. The results suggest that value-dependent learning is sufficient to account for the registration of auditory and visual maps of space in the OT of the barn owl, and they lead to a number of experimental predictions.

Animals↗

Professional education and assessment practices in central auditory processing.

A 17-item questionnaire probing professional preparation and current practices in central auditory assessment was mailed to 500 audiologists selected randomly from the membership directory of the American Academy of Audiology. Data from 183 respondents, representing a 37 percent response rate, were analyzed. The majority of respondents reported minimal academic and clinical preparation in assessment of the central auditory nervous system. Eighty percent of respondents had not taken any graduate course explicitly dedicated to central auditory processing. However, 80 percent had taken at least one basic science course in central audition and 83 percent reported having taken at least one graduate course that included some coverage of central auditory processing and/or the central auditory nervous system. A mean of 3 clinical clock hours accrued in this area was reported. Not surprisingly, 78 percent reported a satisfaction rating of < or =50 percent relative to the graduate education they received in this area and only 41 percent reported providing central auditory assessment. Comparisons with prior surveys show substantial change in the preferred test battery. Most notable is the pivotal role of physiologic measures, with the acoustic reflex and auditory brainstem response listed along with the SCAN as the three most frequently used assessment tests/procedures. Overall, the results suggest a need for improvement in professional preparation in evaluation of central auditory function.

Audiology↗

Tonotopic organization in auditory cortex of the cat.

Microelectrode mapping techniques were employed in the cat's auditory cortex to relate the best frequencies of a large population of neurons with their spatial loci. Based upon the best-frequency distribution, the auditory region was divided into four complete and orderly tonotopic representations and a surrounding belt of cortex in which the tonotopic organization was more complex. The four auditory fields occupy a crescent-shaped band of tissue which comprises portions of both the exposed gyral surfaces and sulcal banks of the ectosylvian cortex. The anterior auditory field (A) is situated most rostrally upon the anterior ectosylvian gyrus. It extends upon the ventral bank of the suprasylvian sulcus and upon the banks of the anterior ectosylvian sulcus. Adjoining field A caudally is the primary auditory field (AI), which extends across the middle ectosylvian gyrus and portions of both banks of the posterior ectosylvian sulcus. The representations of the highest best frequencies in fields A and AI are contiguous. Caudal and ventral to AI are located the posterior (P) and ventroposterior (VP) auditory fields. They lie mainly upon the caudal bank of the posterior ectosylvian sulcus but also extend upon the rostral bank and upon the posterior ectosylvian gyrus. The low best-frequency representations of fields AI and P are contiguous, whereas the low best-frequency representation of field VP lies near the ventral end of the posterior ectosylvian sulcus. Fields P and VP are joined along their middle and high best-frequency representations. Within each auditory field isofrequency lines defined by the spatial loci of neurons with similar best frequencies are oriented orthogonal to the low-to-high best-frequency gradients.

Animals↗

Corticocortical connections of cat primary auditory cortex (AI): laminar organization and identification of supragranular neurons projecting to area AII.

The laminar distribution and structure of the supragranular cells projecting from primary auditory cortex (AI) to the second auditory cortex (AII) in the cat were studied with horseradish peroxidase. Injections in AII retrogradely labeled somata in ipsilateral cortical layers I-VI of AI. A bimodal laminar disposition was found, with approximately 40% of the labeled cells in layer III, 25% in layer V, and 10-15% each in layers II, IV, and VI; only a few cells were found in layer I. The labeled cells were scattered in small aggregates between which unlabeled neurons were interspersed. There was some, though not a strict, topographical distribution of the labeled cells according to the locus of the injection in AII. Injections in the caudal part of AII labeled cells in more rostral AI, while rostral AII injections labeled cells in more caudal AI. Ventral AII injections labeled more ventrally located AI cells, while more dorsal AII injections labeled more dorsally situated AI cells. AII injections also labeled cells in other auditory cortex subdivisions, including the posterior ectosylvian, ventroposterior, temporal, and dorsal auditory zone/suprasylvian fringe cortical areas, and in some non-auditory cortical areas. In layers II and III, both pyramidal and non-pyramidal cells were labeled. More pyramidal cells were labeled in layer III than layer II (80% vs. 62%), and the proportion of non-pyramidal cells in layer II was more than twice that in layer IV (27% vs. 12%). The types of labeled cells were distinguished from one another on the basis of size, somatic and dendritic shape, and laminar distribution. The profiles of labeled cells in these experiments were compared to, and correlated with, those in Golgi-impregnated material. In layer II, the classes of corticocortical projecting cells consisted of small and medium-sized pyramidal, bipolar, and multipolar cells. Those in layer III included small, medium-sized, and large pyramidal neurons, and bipolar and multipolar cells. The average somatic area of the labeled cells did not differ significantly from that of the unlabeled cells, and both were about equal in somatic size to neurons accumulating tritiated gamma-aminobutyric acid in layers II and III. These findings suggest that there is convergent, ipsilateral input onto AII from every layer in AI, and from other cortical auditory and non-auditory areas. A morphologically heterogeneous population of cells in AI contributes to these projections. Diversity in the cytological origins of corticocortical projections implies functional differences between layers II and III since the latter also projects commissural, while layer II in the cat, does not.

Animals↗

A comparative study of avian auditory brainstem responses: correlations with phylogeny and vocal complexity, and seasonal effects.

We conducted a comparative study of the peripheral auditory system in six avian species (downy woodpeckers, Carolina chickadees, tufted titmice, white-breasted nuthatches, house sparrows, and European starlings). These species differ in the complexity and frequency characteristics of their vocal repertoires. Physiological measures of hearing were collected on anesthetized birds using the auditory brainstem response to broadband click stimuli. If auditory brainstem response patterns are phylogenetically conserved, we predicted woodpeckers, sparrows, and starlings to be outliers relative to the other species, because woodpeckers are in a different Order (Piciformes) and, within the Order Passeriformes, sparrows and starlings are in different Superfamilies than the nuthatches, chickadees, and titmice. However, nuthatches and woodpeckers have the simplest vocal repertoires at the lowest frequencies of these six species. If auditory brainstem responses correlate with vocal complexity, therefore, we would predict nuthatches and woodpeckers to be outliers relative to the other four species. Our results indicate that auditory brainstem responses measures in the spring broadly correlated with both vocal complexity and, in some cases, phylogeny. However, these auditory brainstem response patterns shift from spring to winter due to species-specific seasonal changes. These seasonal changes suggest plasticity at the auditory periphery in adult birds.

Acoustic Stimulation↗

The response of single units in the auditory cortex of rhesus monkeys to predicted and to unpredicted sound stimuli.

Rhesus monkeys were trained to predict the nature of short auditory signals of two different types, to which they responded differentially. Prediction was based on visual signals that preceded the auditory ones. The monkey's use of the visual signals as predictors was assessed through two behavioural criteria. (1) performance in trials in which the visual signals was followed by the correct auditory signal (true conditioning) vs performance in trials in which the visual signal was followed by the wrong auditory signal (false conditioning). (2) Reaction time in trials with different types of conditioning. The response of single auditory cortex units to correctly and to incorrectly predicted auditory signals was recorded. The unconditioned response of every unit to each type of auditory stimulus was also obtained. Of 92 units that were analysed in detail, the response of about half was affected by the predictability of the stimulus. Units were affected in two different ways. One involved facilitation of the response to correctly predicted signals and inhibition of the response to incorrectly predicted signals. The other involved facilitation of the response to incorrectly predicted signals and base line response to correctly predicted signals. These findings were discussed in terms of neural mechanisms that relate to prediction.

Animals↗

Bitemporal lesions dissociate auditory evoked potentials and perception.

We studied auditory evoked potentials (AEPs) in an 82-year-old female patient who became suddenly deaf following the second of two strokes. The patient showed markedly elevated pure tone thresholds, was unable to discriminate sounds and could not understand speech. Brain-stem auditory evoked potentials (BAEPs) were normal. CT scans revealed bilateral lesions of the superior temporal plane which included auditory cortex. Two experiments were performed. In the first, tones, complex sounds and speech stimuli were presented at intensities above and below the patient's perceptual threshold. P1, N1 and P2 components were elicited by each of the stimuli--whether or not they were perceived. In particular, stimuli presented below threshold evoked large amplitude, short latency responses comparable to those produced in a control subject. In a second experiment, the refractory properties of the N1-P2 were examined using trains of tones. They were also found to be similar to those of normal subjects. Shifts in the pitch of the tones near the end of the train (when refractory effects were maximal) evoked N1-P2s with enhanced amplitudes, although the change in pitch was not perceived by the patient. In both experiments AEP scalp topographies were normal. The results suggest that bitemporal lesions of auditory cortex can dissociate auditory perception and long-latency auditory evoked potentials. A review of evoked potential studies of cortical deafness suggests that the neural circuits responsible for N1-P2 generation lie in close proximity to those necessary for auditory perception.

Aged↗

Noise impairment in the guinea pig. I. Changes in electrical evoked activity along the auditory pathway.

Changes in the cochlear microphonics (CM), auditory nerve action potential (AP), and evoked responses from the inferior colliculus (IC-ER) and auditory cortex (AC-ER) of the guinea pig were assessed after exposure to white noise of 115 dB for 30 min. Both continuous and intermittent (200 ms noise and 200 ms pause) exposures were used. In comparison with the pre-exposure level, CM isopotential curves were shifted by 1.1 +/- 0.5 dB (means +/- S.E.) on the average in the range of 0.5-8 kHz (recorded at the round window). The amplitude-intensity function of the click-evoked auditory nerve action potential decreased by 8.4 +/- 1.2 dB, that of the inferior colliculus evoked response by 20.9 +/- 3.7 dB, and the amplitude-intensity function of the auditory cortex evoked potential decreased by 6.2 +/- 4.7 dB. A similar reduction in the amplitude was found after both continuous and intermittent noise exposure. In contrast to the decrease in amplitudes of evoked potentials, the latency-intensity functions of the individual waves of potentials evoked along the auditory pathway did not change when compared at the same click intensity before and after the exposure. The results suggest that individual auditory nuclei are impaired by the noise to different extents and that the impairment does not increase linearly up to the auditory cortex.

Action Potentials↗

Reduced auditory efferent activity in childhood selective mutism.

BACKGROUND: Selective mutism is a psychiatric disorder of childhood characterized by consistent inability to speak in specific situations despite the ability to speak normally in others. The objective of this study was to test whether reduced auditory efferent activity, which may have direct bearings on speaking behavior, is compromised in selectively mute children. METHODS: Participants were 16 children with selective mutism and 16 normally developing control children matched for age and gender. All children were tested for pure-tone audiometry, speech reception thresholds, speech discrimination, middle-ear acoustic reflex thresholds and decay function, transient evoked otoacoustic emission, suppression of transient evoked otoacoustic emission, and auditory brainstem response. RESULTS: Compared with control children, selectively mute children displayed specific deficiencies in auditory efferent activity. These aberrations in efferent activity appear along with normal pure-tone and speech audiometry and normal brainstem transmission as indicated by auditory brainstem response latencies. CONCLUSIONS: The diminished auditory efferent activity detected in some children with SM may result in desensitization of their auditory pathways by self-vocalization and in reduced control of masking and distortion of incoming speech sounds. These children may gradually learn to restrict vocalization to the minimal amount possible in contexts that require complex auditory processing.

Acoustic Impedance Tests↗

Experimental-neuromodeling framework for understanding auditory object processing: integrating data across multiple scales.

In this article, we review a combined experimental-neuromodeling framework for understanding brain function with a specific application to auditory object processing. Within this framework, a model is constructed using the best available experimental data and is used to make predictions. The predictions are verified by conducting specific or directed experiments and the resulting data are matched with the simulated data. The model is refined or tested on new data and generates new predictions. The predictions in turn lead to better-focused experiments. The auditory object processing model was constructed using available neurophysiological and neuroanatomical data from mammalian studies of auditory object processing in the cortex. Auditory objects are brief sounds such as syllables, words, melodic fragments, etc. The model can simultaneously simulate neuronal activity at a columnar level and neuroimaging activity at a systems level while processing frequency-modulated tones in a delayed-match-to-sample task. The simulated neuroimaging activity was quantitatively matched with neuroimaging data obtained from experiments; both the simulations and the experiments used similar tasks, sounds, and other experimental parameters. We then used the model to investigate the neural bases of the auditory continuity illusion, a type of perceptual grouping phenomenon, without changing any of its parameters. Perceptual grouping enables the auditory system to integrate brief, disparate sounds into cohesive perceptual units. The neural mechanisms underlying auditory continuity illusion have not been studied extensively with conventional neuroimaging or electrophysiological techniques. Our modeling results agree with behavioral studies in humans and an electrophysiological study in cats. The results predict a particular set of bottom-up cortical processing mechanisms that implement perceptual grouping, and also attest to the robustness of our model.

Acoustic Stimulation↗

Sounds do-able: auditory-motor transformations and the posterior temporal plane.

Accumulating evidence in humans and non-human primates implicates the posterior superior temporal plane (STP) in the processing of both auditory spatial information and vocal sounds. Such evidence is difficult to reconcile with existing accounts of the primate auditory brain. We propose that the posteromedial STP generates sequenced auditory representations by matching incoming auditory information with stored templates. These sequenced auditory representations are subsequently used to constrain motor responses. We argue for a re-assessment of the much-debated dorsal auditory pathway in terms of its generic behavioral role as an auditory "do" pathway.

Animal Communication↗

Mitosis and apoptosis in postnatal auditory system of the C3H/He strain.

The mouse auditory neurons, hair cells and their supporting cells in the cochlea are considered to be generated mainly in the embryonic days and to be sustained throughout the whole life. In the present study, however, we observed that auditory ganglion cells in the spiral ganglia undergo apoptosis and mitosis in the suckling mouse (1- to 2-week-old C3H/HeJ mice) with a normal auditory system. In spiral ganglia at postnatal days 7 (P7) and 10 (P10), TUNEL (TdT-mediated dUTP nick-end labeling)-positive and morphologically apoptotic ganglion cells were found. Furthermore, by bromodeoxyuridine labeling, mitosis of auditory ganglion cells was found at P10 to P14. In a functional study of auditory brainstem response, we demonstrated that the C3H/HeJ mouse acquires the ability to hear airborne sound at P12 and this is the same time as the opening of their external acoustic meatus (EAM). These results indicate that C3H/HeJ auditory ganglion cells have the ability to proliferate even after opening of the EAM and the initial input of airborne sound. We found that postnatal apoptosis and mitosis after P7 also occurred in the greater epithelial ridge (GER) which is an important organ for maturation of the organ of Corti and is located around the inner hair cells. This indicates that GER cells are not only degenerated but also regenerated until their disappearance around P12. This is the first report on mammals to demonstrate that neuronal mitosis of spiral ganglion cells and that of GER cells occur not only in embryonic and neonatal development but also in postnatal development of the normal auditory system.

Age Factors↗

Latency as a function of intensity in auditory neurons: influences of central processing.

The response latencies of sensory neurons typically shorten with increases in stimulus intensity. In the central auditory system this phenomenon should have a significant impact on a number of auditory functions that depend critically on an integration of precisely timed neural inputs. Evidence from previous studies suggests that the auditory system not only copes with the potential problems associated with intensity-dependent latency change, but that it also modifies latency change to shape the response properties of many cells for specific functions. This observation suggests that intensity-dependent latency change may undergo functional transformations along the auditory neuraxis. The goal of our study was to explore these transformations by making a direct, quantitative comparison of intensity-dependent latency change among a number of auditory centers from the lower brainstem to the thalamus. We found two main ways in which intensity-dependent latency change transformed along the neuraxis: (1) the range of latency change increased substantially and (2) one particular type of latency change, which has been suggested to be associated with sensitivity to temporally segregated stimulus components, occurred only at the highest centers tested, the midbrain and thalamus. Additional testing in the midbrain (inferior colliculus) indicated that inhibitory inputs are involved in shaping latency change. Our findings demonstrate that the central auditory system modifies intensity-dependent latency changes. We suggest that these changes may be functionally incorporated, actively enhanced, or modified to suit specific functions of the auditory system.

Animals↗

Neural plasticity following auditory training in children with learning problems.

OBJECTIVE: This study examined the plasticity of the central auditory pathway and accompanying cognitive changes in children with learning problems. METHODS: Children diagnosed with a learning disability and/or attention deficit disorder worked with commercial auditory processing training software for 8 weeks; control groups consisted of normal-learning and learning-impaired children who did not participate in any remedial programs. Auditory brainstem function was evaluated in response to click and speech stimuli in quiet; cortical responses to speech stimuli were obtained in quiet and noise. Academic achievement and cognitive abilities were assessed with standardized measures. RESULTS: Compared to controls, the trained group improved on measures of auditory processing and exhibited changes in cortical responses in quiet and in noise. In quiet, cortical responses reflected an accelerated maturational pattern; in background noise, cortical responses became more resistant to degradation. Brainstem responses did not change with training. CONCLUSIONS: Children with learning problems who practiced with auditory training software exhibited plasticity of neural encoding of speech sounds at the cortical, but not subcortical, level of the auditory pathway. This plasticity was accompanied by improvement in behavioral performance. SIGNIFICANCE: This study demonstrates that in learning-impaired children working with commercial auditory processing training programs affects both the perception and the cortical representation of sound.

Achievement↗

A comparison of higher order auditory processes in younger and older adults.

The authors conducted 3 experiments investigating the effects of aging on higher order auditory processes. They compared younger and older adults with respect to (a) their auditory channel capacity, (b) the extent of their top-down control over auditory gain, and (c) their ability to focus attention on a narrow band of frequencies. To ensure that subclinical cochlear processing deficits in older adults (e.g., higher thresholds, poorer discrimination of frequency and intensity differences) did not limit performance, the authors used only stimuli that were perfectly discriminable by all participants. No age differences were found in any of these experiments, suggesting that some higher order auditory processes (e.g., top-down control over auditory gain, auditory attention) are preserved in normal aging, despite numerous age-related declines in peripheral auditory functionality.

Adult↗

Functional brain imaging of tinnitus-like perception induced by aversive auditory stimuli.

Tinnitus is an aversive auditory percept of unknown origin. We tested the speculation that tinnitus may share neuronal processing mechanisms with aversive auditory percepts of known origin. This study revealed the functional neuroanatomy of the perception of aversive auditory stimuli. The stimuli were presented to 12 healthy volunteers so as to mimic the psychoacoustical features of tinnitus and its affective response in tinnitus sufferers. The regional cerebral blood flow distribution was measured by PET during four auditory processing conditions and one control condition. The aversive auditory stimuli activated primary and secondary auditory areas bilaterally, dorsolateral prefrontal attention areas, and structures in the limbic system which subserve emotional processing. Based on these results and findings from other functional neuroimages of tinnitus, we hypothesize that the perception of tinnitus may involve the functional linkage of these brain areas: secondary auditory cortex, dorsolateral prefrontal cortex, and limbic system.

Acoustic Stimulation↗

A review of objective methods of evaluating auditory neural pathways.

OBJECTIVES: Review physiological methods of evaluating function of the auditory neural pathways in infants, children, and adults. Present two case studies to demonstrate the usefulness of physiological measures in assessing abnormalities of the auditory neural pathways. STUDY DESIGN: Review of applications of physiological measures of auditory neural function. METHODS: Review otoacoustic emissions, auditory evoked potentials, and efferent reflexes, and discuss their use in identifying disorders of the auditory neural pathways from the cochlea to the cortex. RESULTS: Auditory disorders occur from peripheral to central areas of the neural system. Patients with disorders of the peripheral nerve and/or the input from the cochlear inner hair cells, such as the patients presently described as having auditory neuropathy, demonstrate abnormal efferent reflexes, including middle ear muscle reflexes and efferent suppression of otoacoustic emissions, as well as grossly abnormal brainstem evoked potentials. In contrast, patients with more central disorders can be differentiated by normal results on tests of the neural periphery and abnormal findings on cortical evoked potentials and other measures of central function. CONCLUSIONS: Physiological measures are sensitive, objective, and less variable in assessing neural disorders than traditional behavioral measures such as pure-tone or speech audiometry.

Adolescent↗