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The insula (Island of Reil) and its role in auditory processing. Literature review.

The insular cortex is a complex structure which contains areas that subserve visceral sensory, motor, vestibular, and somatosensory functions. The role of the insular cortex in auditory processing was poorly understood until recently. However, recent case studies indicate that bilateral damage to the insulae may result in total auditory agnosia. Functional imaging studies demonstrate that the insulae participate in several key auditory processes, such as allocating auditory attention and tuning in to novel auditory stimuli, temporal processing, phonological processing and visual-auditory integration. These studies do not clarify the issue of further specialisation within the insular cortex, e.g. whether the posterior insulae are primarily sensory areas, while the anterior insulae serve mainly as integration/association auditory areas, two hypotheses that would be compatible with the cytoarchitectonic structure and connectivity of the insulae. The functional characterisation of the insulae remains incomplete, underlining the need for further studies.

Animals↗

Infant discrimination of rapid auditory cues predicts later language impairment.

The etiology and mechanisms of specific language impairment (SLI) in children are unknown. Differences in basic auditory processing abilities have been suggested to underlie their language deficits. Studies suggest that the neuropathology, such as atypical patterns of cerebral lateralization and cortical cellular anomalies, implicated in such impairments likely occur early in life. Such anomalies may play a part in the rapid processing deficits seen in this disorder. However, prospective, longitudinal studies in infant populations that are critical to examining these hypotheses have not been done. In the study described, performance on brief, rapidly-presented, successive auditory processing and perceptual-cognitive tasks were assessed in two groups of infants: normal control infants with no family history of language disorders and infants from families with a positive family history for language impairment. Initial assessments were obtained when infants were 6-9 months of age (M=7.5 months) and the sample was then followed through age 36 months. At the first visit, infants' processing of rapid auditory cues as well as global processing speed and memory were assessed. Significant differences in mean thresholds were seen in infants born into families with a history of SLI as compared with controls. Examination of relations between infant processing abilities and emerging language through 24 months-of-age revealed that threshold for rapid auditory processing at 7.5 months was the single best predictor of language outcome. At age 3, rapid auditory processing threshold and being male, together predicted 39-41% of the variance in language outcome. Thus, early deficits in rapid auditory processing abilities both precede and predict subsequent language delays. These findings support an essential role for basic nonlinguistic, central auditory processes, particularly rapid spectrotemporal processing, in early language development. Further, these findings provide a temporal diagnostic window during which future language impairments may be addressed.

Apgar Score↗

Auditory gating processes and binaural inhibition in the inferior colliculus.

Physiological/behavioral/perceptual responses to an auditory stimulus can be inhibited by another leading auditory stimulus at certain stimulus intervals, and have been considered useful models of auditory gating processes. Two typical examples of auditory gating are prepulse inhibition of the startle reflex and the precedence effect (echo suppression). This review summarizes studies of these two auditory gating processes with regard to their biological significance, cognitive modulation, binaural properties, and underlying neural mechanisms. Both prepulse inhibition and the precedence effect have gating functions of reducing the disruptive influence of the lagging sound, but prepulse inhibition has a much longer temporal window than the precedence effect. Attentional processes can modulate prepulse inhibition, and the listener's previous experience can modulate the precedence effect. Compared to monaural hearing, binaural hearing reduces prepulse inhibition but enhances the precedence effect. The inferior colliculus, the major structure of the auditory midbrain, plays an important role in mediating these two auditory gating processes, and inhibitory neural transmissions within the inferior colliculus may account for binaural inhibition observed in prepulse inhibition and lag suppression recorded in the inferior colliculus. The neural mechanisms underlying binaural inhibition in the inferior colliculus are also discussed.

Acoustic Stimulation↗

Brainstem auditory-evoked potentials in iron-deficiency anemia.

Slight-to-moderate impairments may be observed in mental and motor developments of infants with iron- deficiency anemia. Brainstem auditory-evoked potentials provide a noninvasive means of examining the auditory aspect of the central nervous system functions. In this study the effect of iron-deficiency anemia on auditory functions was investigated by using brainstem auditory-evoked potentials. Brainstem auditory-evoked potentials of the 20 iron-deficient infants were not significantly different from those of the control group that included 20 healthy age-matched infants. Furthermore, there was not a statistically significant difference between the brainstem auditory-evoked potentials of the study group performed before and 3 months after oral iron therapy. Although we could not demonstrate a hearing loss in infants with moderate iron-deficiency anemia in this study, the relationship between severe iron-deficiency anemia and hearing loss or auditory dysfunction remains to be determined.

Anemia, Iron-Deficiency↗

Eye position affects activity in primary auditory cortex of primates.

BACKGROUND: Neurons in primary auditory cortex are known to be sensitive to the locations of sounds in space, but the reference frame for this spatial sensitivity has not been investigated. Conventional wisdom holds that the auditory and visual pathways employ different reference frames, with the auditory pathway using a head-centered reference frame and the visual pathway using an eye-centered reference frame. Reconciling these discrepant reference frames is therefore a critical component of multisensory integration. RESULTS: We tested the reference frame of neurons in the auditory cortex of primates trained to fixate visual stimuli at different orbital positions. We found that eye position altered the activity of about one third of the neurons in this region (35 of 113, or 31%). Eye position affected not only the responses to sounds (26 of 113, or 23%), but also the spontaneous activity (14 of 113, or 12%). Such effects were also evident when monkeys moved their eyes freely in the dark. Eye position and sound location interacted to produce a representation for auditory space that was neither head- nor eye-centered in reference frame. CONCLUSIONS: Taken together with emerging results in both visual and other auditory areas, these findings suggest that neurons whose responses reflect complex interactions between stimulus position and eye position set the stage for the eventual convergence of auditory and visual information.

Acoustic Stimulation↗

A unified mechanism for spontaneous-rate and first-spike timing in the auditory nerve.

Recent physiological experiments have provided detailed descriptions of the properties of first-spike latency and variability in auditory cortex and nerve in response to pure tones with different envelopes. The envelope-dependence of first-spike timing and precision in auditory cortical neurons appears to reflect properties established in the nerve. First-spike latency properties in individual auditory nerve fibers are strongly correlated with their spontaneous rate (SR). It is shown here that a minimal, plausible model of auditory transduction with two free parameters accurately reproduces the physiological data from the auditory nerve population. The model consists of a simple gain stage, a bandpass filter, a rectifying saturating non-linearity, and a lowpass filter in series. The output of the lowpass filter drives an inhomogeneous Poisson process. The shape of the non-linearity is determined by SR; in physiological terms, this shape depends upon the resting sensitivity of the synapse between the inner hair cell and the auditory nerve. An alternative model for SR generation, where SR is added to the stimulus-driven output of a fixed nonlinearity, fails to account for the data. The results provide a novel, comprehensive and physiologically-based explanation for the range of experimental results on the envelope-dependence of first-spike latency and precision, and its relationship with SR, in the auditory system.

Acoustic Stimulation↗

A corollary discharge maintains auditory sensitivity during sound production.

Speaking and singing present the auditory system of the caller with two fundamental problems: discriminating between self-generated and external auditory signals and preventing desensitization. In humans and many other vertebrates, auditory neurons in the brain are inhibited during vocalization but little is known about the nature of the inhibition. Here we show, using intracellular recordings of auditory neurons in the singing cricket, that presynaptic inhibition of auditory afferents and postsynaptic inhibition of an identified auditory interneuron occur in phase with the song pattern. Presynaptic and postsynaptic inhibition persist in a fictively singing, isolated cricket central nervous system and are therefore the result of a corollary discharge from the singing motor network. Mimicking inhibition in the interneuron by injecting hyperpolarizing current suppresses its spiking response to a 100-dB sound pressure level (SPL) acoustic stimulus and maintains its response to subsequent, quieter stimuli. Inhibition by the corollary discharge reduces the neural response to self-generated sound and protects the cricket's auditory pathway from self-induced desensitization.

Acoustic Stimulation↗

Classical conditioning modifies cytochrome oxidase activity in the auditory system.

The effects of excitatory classical conditioning on cytochrome oxidase activity in the central auditory system were investigated using quantitative histochemistry. Rats in the conditioned group were trained with consistent pairings of a compound conditional stimulus (a tone and a light) with a mild footshock, to elicit conditioned suppression of drinking. Rats in the pseudorandom group were exposed to pseudorandom presentations of the same tone, light and shock stimuli without consistent pairings. Untrained rats in a naive group did not receive presentations of the experimental stimuli. The findings demonstrated that auditory fear conditioning modifies the metabolic neuronal responses of the auditory system, supporting the hypothesis that sensory neurons are responsive to behavioural stimulus properties acquired by learning. There was a clear distinction between thalamocortical and lower divisions of the auditory system based on the differences in metabolic activity evoked by classical conditioning, which lead to an overt learned behavioural response versus pseudorandom stimulus presentations, which lead to behavioural habituation. Increases in cytochrome oxidase activity indicated that tone processing is enhanced during associative conditioning at upper auditory structures (medial geniculate nucleus and secondary auditory cortices). In contrast, metabolic activation of lower auditory structures (cochlear nuclei and inferior colliculus) in response to the pseudorandom presentation of the experimental stimuli suggest that these areas may be activated during habituation to tone stimuli. Together these findings show that mapping the metabolic activity of cytochrome oxidase with quantitative histochemistry can be successfully used to map regional long-lasting effects of learning on brain systems.

Acoustic Stimulation↗

Visual speech speeds up the neural processing of auditory speech.

Synchronous presentation of stimuli to the auditory and visual systems can modify the formation of a percept in either modality. For example, perception of auditory speech is improved when the speaker's facial articulatory movements are visible. Neural convergence onto multisensory sites exhibiting supra-additivity has been proposed as the principal mechanism for integration. Recent findings, however, have suggested that putative sensory-specific cortices are responsive to inputs presented through a different modality. Consequently, when and where audiovisual representations emerge remain unsettled. In combined psychophysical and electroencephalography experiments we show that visual speech speeds up the cortical processing of auditory signals early (within 100 ms of signal onset). The auditory-visual interaction is reflected as an articulator-specific temporal facilitation (as well as a nonspecific amplitude reduction). The latency facilitation systematically depends on the degree to which the visual signal predicts possible auditory targets. The observed auditory-visual data support the view that there exist abstract internal representations that constrain the analysis of subsequent speech inputs. This is evidence for the existence of an "analysis-by-synthesis" mechanism in auditory-visual speech perception.

Adult↗

Spatial processing in the auditory cortex of the macaque monkey.

The patterns of cortico-cortical and cortico-thalamic connections of auditory cortical areas in the rhesus monkey have led to the hypothesis that acoustic information is processed in series and in parallel in the primate auditory cortex. Recent physiological experiments in the behaving monkey indicate that the response properties of neurons in different cortical areas are both functionally distinct from each other, which is indicative of parallel processing, and functionally similar to each other, which is indicative of serial processing. Thus, auditory cortical processing may be similar to the serial and parallel "what" and "where" processing by the primate visual cortex. If "where" information is serially processed in the primate auditory cortex, neurons in cortical areas along this pathway should have progressively better spatial tuning properties. This prediction is supported by recent experiments that have shown that neurons in the caudomedial field have better spatial tuning properties than neurons in the primary auditory cortex. Neurons in the caudomedial field are also better than primary auditory cortex neurons at predicting the sound localization ability across different stimulus frequencies and bandwidths in both azimuth and elevation. These data support the hypothesis that the primate auditory cortex processes acoustic information in a serial and parallel manner and suggest that this may be a general cortical mechanism for sensory perception.

Acoustic Stimulation↗

Auditory neuropathy.

Ten patients presented as children or young adults with hearing impairments that, by behavioural and physiological testing, were compatible with a disorder of the auditory portion of the VIII cranial nerve. Evidence of normal cochlear outer hair cell function was provided by preservation of otoacoustic emissions and cochlear microphonics in all of the patients. Auditory brainstem potentials showed evidence of abnormal auditory pathway function beginning with the VIII nerve: the potentials were absent in nine patients and severely distorted in one patient. Auditory brainstem reflexes (middle ear muscles; crossed suppression of otoacoustic emissions) were absent in all of the tested patients. Behavioural audiometric testing showed a mild to moderate elevation of pure tone threshold in nine patients. The extent of the hearing loss, if due to cochlear receptor damage, should not have resulted in the loss of auditory brainstem potentials. The shape of the pure tone loss varied, being predominantly low frequency in five patients, flat across all frequencies in three patients and predominantly high frequency in two patients. Speech intelligibility was tested in eight patients, and in six was affected out of proportion to what would have been expected if the pure tone loss were of cochlear origin. The patients were otherwise neurologically normal when the hearing impairment was first manifest. Subsequently, eight of these patients developed evidence for a peripheral neuropathy. The neuropathy was hereditary in three and sporadic in five. We suggest that this type of hearing impairment is due to a disorder of auditory nerve function and may have, as one of its causes, a neuropathy of the auditory nerve, occurring either in isolation or as part of a generalized neuropathic process.

Adolescent↗

Sequential effects of increasing propofol sedation on frontal and temporal cortices as indexed by auditory event-related potentials.

BACKGROUND: It is an open question whether cognitive processes of auditory perception that are mediated by functionally different cortices exhibit the same sensitivity to sedation. The auditory event-related potentials P1, mismatch negativity (MMN), and early right anterior negativity (ERAN) originate from different cortical areas and reflect different stages of auditory processing. The P1 originates mainly from the primary auditory cortex. The MMN is generated in or in the close vicinity of the primary auditory cortex but is also dependent on frontal sources. The ERAN mainly originates from frontal generators. The purpose of the study was to investigate the effects of increasing propofol sedation on different stages of auditory processing as reflected in P1, MMN, and ERAN. METHODS: The P1, the MMN, and the ERAN were recorded preoperatively in 18 patients during four levels of anesthesia adjusted with target-controlled infusion: awake state (target concentration of propofol 0.0 microg/ml), light sedation (0.5 microg/ml), deep sedation (1.5 microg/ml), and unconsciousness (2.5-3.0 microg/ml). Simultaneously, propofol anesthesia was assessed using the Bispectral Index. RESULTS: Propofol sedation resulted in a progressive decrease in amplitudes and an increase of latencies with a similar pattern for MMN and ERAN. MMN and ERAN were elicited during sedation but were abolished during unconsciousness. In contrast, the amplitude of the P1 was unchanged by sedation but markedly decreased during unconsciousness. CONCLUSION: The results indicate differential effects of propofol sedation on cognitive functions that involve mainly the auditory cortices and cognitive functions that involve the frontal cortices.

Acoustic Stimulation↗

A cortical region sensitive to auditory spectral motion.

The functional architecture of human auditory cortex is still poorly understood compared with that of visual cortex, yet anatomical and electrophysiological studies in non-human primates suggest that the auditory cortex also might be functionally specialized, in a model of parallel and hierarchical organization. In particular, spectral changes such as the formant transitions of speech, or spectral motion (SM) by analogy with visual motion, could be processed in specialized cortical regions. In this study, positron emission tomography (PET) was used to identify which auditory cortical region are involved in SM analysis. We found that a bilateral secondary auditory cortical region, located in the caudal-lateral belt of auditory cortex, was more sensitive to auditory stimuli containing spectral changes than to matched stimuli with a stationary spectral profile. This result suggests that analogies between sensory systems could prove useful in the research into the functional organization of the auditory cortex.

Acoustic Stimulation↗

Requirement of the auditory association cortex for discrimination of vowel-like sounds in rats.

We investigated the roles of the auditory cortex in discrimination learning of vowel-like sounds consisting of multiple formants. Rats were trained to discriminate between synthetic sounds with four formants. Bilateral electrolytic lesions including the primary auditory cortex and the dorsal auditory association cortex impaired multiformant discrimination, whereas they did not significantly affect discrimination between sounds with a single formant or between pure tones. Local lesions restricted to the dorsal/rostral auditory association cortex were sufficient to attenuate multiformant discrimination learning, and lesions restricted to the primary auditory cortex had no significant effects. These findings indicate that the dorsal/rostral auditory association cortex but not the primary auditory cortex is required for discrimination learning of vowel-like sounds with multiple formants in rats.

Acoustic Stimulation↗

Primitive auditory stream segregation: a neurophysiological study in the songbird forebrain.

Auditory stream segregation refers to the perceptual grouping of sounds, to form coherent representations of objects in the acoustic scene, and is a fundamental aspect of hearing and speech perception. The perceptual segregation of simple interleaved tone sequences has been studied in humans and European starlings (Sturnus vulgaris) using sequences of 2 alternating tones differing in frequency (ABA-ABA-ABA-...). The segregation of A and B tones into separate auditory streams is believed to be promoted by preattentive auditory processes that increase the separation of excitation patterns along a tonotopic gradient. We tested the hypothesis that frequency selectivity and forward masking operate as 2 preattentive processes in sequential stream segregation by recording neural responses in the auditory forebrain of awake starlings to repeated ABA- sequences in which we varied the frequency separation (DeltaF) between the A and B tones and the tone repetition time (TRT). The A tones were presented at the neurons' characteristic frequency (CF), and B tones differed from the CF over a one-octave range. Larger DeltaF values and shorter TRTs promote the perceptual segregation of alternating tone sequences in humans and also resulted in larger differences in neural responses to alternating CF (A) and non-CF (B) tones. Our results are consistent with the hypothesis that preattentive auditory processes, such as frequency selectivity and forward masking, contribute to the perceptual segregation of sequential acoustic events having different frequencies into separate auditory streams, but also suggest that additional processes may be required to account for all known perceptual effects related to sequential auditory stream segregation.

Acoustic Stimulation↗

A corollary discharge mechanism modulates central auditory processing in singing crickets.

Crickets communicate using loud (100 dB SPL) sound signals that could adversely affect their own auditory system. To examine how they cope with this self-generated acoustic stimulation, intracellular recordings were made from auditory afferent neurons and an identified auditory interneuron-the Omega 1 neuron (ON1)-during pharmacologically elicited singing (stridulation). During sonorous stridulation, the auditory afferents and ON1 responded with bursts of spikes to the crickets' own song. When the crickets were stridulating silently, after one wing had been removed, only a few spikes were recorded in the afferents and ON1. Primary afferent depolarizations (PADs) occurred in the terminals of the auditory afferents, and inhibitory postsynaptic potentials (IPSPs) were apparent in ON1. The PADs and IPSPs were composed of many summed, small-amplitude potentials that occurred at a rate of about 230 Hz. The PADs and the IPSPs started during the closing wing movement and peaked in amplitude during the subsequent opening wing movement. As a consequence, during silent stridulation, ON1's response to acoustic stimuli was maximally inhibited during wing opening. Inhibition coincides with the time when ON1 would otherwise be most strongly excited by self-generated sounds in a sonorously stridulating cricket. The PADs and the IPSPs persisted in fictively stridulating crickets whose ventral nerve cord had been isolated from muscles and sense organs. This strongly suggests that the inhibition of the auditory pathway is the result of a corollary discharge from the stridulation motor network. The central inhibition was mimicked by hyperpolarizing current injection into ON1 while it was responding to a 100 dB SPL sound pulse. This suppressed its spiking response to the acoustic stimulus and maintained its response to subsequent, quieter stimuli. The corollary discharge therefore prevents auditory desensitization in stridulating crickets and allows the animals to respond to external acoustic signals during the production of calling song.

Acoustic Stimulation↗

Consonance and dissonance of musical chords: neural correlates in auditory cortex of monkeys and humans.

Some musical chords sound pleasant, or consonant, while others sound unpleasant, or dissonant. Helmholtz's psychoacoustic theory of consonance and dissonance attributes the perception of dissonance to the sensation of "beats" and "roughness" caused by interactions in the auditory periphery between adjacent partials of complex tones comprising a musical chord. Conversely, consonance is characterized by the relative absence of beats and roughness. Physiological studies in monkeys suggest that roughness may be represented in primary auditory cortex (A1) by oscillatory neuronal ensemble responses phase-locked to the amplitude-modulated temporal envelope of complex sounds. However, it remains unknown whether phase-locked responses also underlie the representation of dissonance in auditory cortex. In the present study, responses evoked by musical chords with varying degrees of consonance and dissonance were recorded in A1 of awake macaques and evaluated using auditory-evoked potential (AEP), multiunit activity (MUA), and current-source density (CSD) techniques. In parallel studies, intracranial AEPs evoked by the same musical chords were recorded directly from the auditory cortex of two human subjects undergoing surgical evaluation for medically intractable epilepsy. Chords were composed of two simultaneous harmonic complex tones. The magnitude of oscillatory phase-locked activity in A1 of the monkey correlates with the perceived dissonance of the musical chords. Responses evoked by dissonant chords, such as minor and major seconds, display oscillations phase-locked to the predicted difference frequencies, whereas responses evoked by consonant chords, such as octaves and perfect fifths, display little or no phase-locked activity. AEPs recorded in Heschl's gyrus display strikingly similar oscillatory patterns to those observed in monkey A1, with dissonant chords eliciting greater phase-locked activity than consonant chords. In contrast to recordings in Heschl's gyrus, AEPs recorded in the planum temporale do not display significant phase-locked activity, suggesting functional differentiation of auditory cortical regions in humans. These findings support the relevance of synchronous phase-locked neural ensemble activity in A1 for the physiological representation of sensory dissonance in humans and highlight the merits of complementary monkey/human studies in the investigation of neural substrates underlying auditory perception.

Acoustic Stimulation↗

Functional imaging of the auditory system: the use of positron emission tomography.

Modern brain imaging methods now afford unprecedented opportunities for the in vivo study of central auditory system function. Positron emission tomography (PET) has been used as a functional imaging technique for more than 15 years to study the distribution of cerebral haemodynamic changes associated with auditory stimulation, in subjects with normal and abnormal auditory function. Many of these studies concern processes related to, but not identical with, audition, such as speech perception, melodic processing and directed attention. Additionally, PET has been used to explore auditory perception in clinical populations such as cochlear implantees, neurosurgical candidates and people with tinnitus and auditory hallucinations. The spatial resolution of PET does not appear sufficient to address questions of a fine anatomical grain, e.g. exploring functional specialization within the primary and secondary auditory cortex. Nevertheless, PET has considerable potential as a tool in basic research on, and clinical assessment of, many auditory phenomena. Although functional magnetic resonance imaging can now be used for many studies for which formerly only PET was suitable, PET still possesses unique advantages. For example, it permits acquisition of data from inferior frontal and anterior temporal areas, can be used with subjects with cochlear and other implants (such as pacemakers) and can be used to map neurochemical pathways and receptors.

Auditory Cortex↗