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Activity-dependent developmental plasticity of the auditory brain stem in children who use cochlear implants.

OBJECTIVES: 1) To determine if a period of early auditory deprivation influences neural activity patterns as revealed by human auditory brain stem potentials evoked by electrical stimulation from a cochlear implant. 2) To examine the potential for plasticity in the human auditory brain stem. Specifically, we asked if electrically evoked auditory potentials from the auditory nerve and brain stem in children show evidence of development as a result of implant use. 3) To assess whether a sensitive or critical period exists in auditory brain stem development. Specifically, is there an age of implantation after which there are no longer developmental changes in auditory brain stem activity as revealed by electrically evoked potentials? DESIGN: The electrically evoked compound potential of the auditory nerve (ECAP) and the electrically evoked auditory brain stem response (EABR) were recorded repeatedly during the first year of implant use in each of 50 children. The children all had pre- or peri-lingual onset of severe to profound sensorineural hearing loss and received their implants at ages ranging from 12 mo to 17 yr. All children received Nucleus cochlear implant devices. All children were in therapy and in school programs that emphasized listening and required the children to wear their implants consistently. RESULTS: Initial stimulation from the cochlear implant evoked clear responses from the auditory nerve and auditory brain stem in most children. There was no correlation between minimum latency, maximum amplitude, or slope of amplitude growth of initial responses with age at implantation for ECAP eN1, EABR eIII and eV components (p > 0.05). During the first year of implant use, minimum latency of these waves significantly decreased (p < 0.01, p < 0.0001, p < 0.0001, respectively). Neural conduction time, measured using the interwave latency of ECAP eN1-EABR eIII for lower brain stem and EABR eIII-eV for upper brain stem, decreased during the period of 6 to 12 mo of cochlear implant use (p < 0.01 (lower), p < 0.0001(upper)). The ECAP wave eN1 and the EABR wave eV showed significant increases in amplitude during time of implant use (p < 0.05 and p < 0.01, respectively). There were no correlations between the rate of interwave latency decrease and the rate of amplitude increases and the age at which children underwent implantation (p < 0.05). CONCLUSIONS: Activity in the auditory pathways to the level of the midbrain can be evoked by acute stimulation from a cochlear implant. EABR measures are not influenced by any period of auditory deprivation. Auditory development proceeds once the implant is activated and involves improvements in neural conduction velocity and neural synchrony. Underlying mechanisms likely include improvements in synaptic efficacy and possibly increased myelination. The developmental plasticity that we have shown in the human auditory brain stem does not appear from EABR data to be limited by a critical period during childhood.

Adolescent↗

Tone-evoked oscillations in the rat auditory cortex result from interactions between the thalamus and reticular nucleus.

This study investigates the origins of tone-evoked oscillations (5-13 Hz) in the thalamo-cortical auditory system of anaesthetized rats. In three separate experiments, the auditory sector of the reticular nucleus (RE), the auditory cortex and the auditory thalamus were inactivated by local applications of muscimol (1 mg/mL). To assess the efficacy of this procedure, recordings were performed in the inactivated structure in each experiment; and to determine the extent of the drug diffusion autoradiographic experiments were carried out. The evolution of the strength of the oscillations was followed using power spectra during the whole recording session. In the first experiment, muscimol injection in the auditory RE totally suppressed the tone-evoked oscillations in the auditory thalamus and cortex. In the second experiment, inactivation of the auditory cortex did not interfere with the presence of tone-evoked oscillations in the auditory RE. In the third experiment, inactivation of the auditory thalamus impaired the oscillations produced by cortical stimulation in the auditory RE. From these results, it appears that both the auditory thalamus and the auditory sector of the RE, but not the auditory cortex, are involved in the generation of stimulus-evoked oscillations in the thalamo-cortical auditory system.

Acoustic Stimulation↗

Eye-centered, head-centered, and complex coding of visual and auditory targets in the intraparietal sulcus.

The integration of visual and auditory events is thought to require a joint representation of visual and auditory space in a common reference frame. We investigated the coding of visual and auditory space in the lateral and medial intraparietal areas (LIP, MIP) as a candidate for such a representation. We recorded the activity of 275 neurons in LIP and MIP of two monkeys while they performed saccades to a row of visual and auditory targets from three different eye positions. We found 45% of these neurons to be modulated by the locations of visual targets, 19% by auditory targets, and 9% by both visual and auditory targets. The reference frame for both visual and auditory receptive fields ranged along a continuum between eye- and head-centered reference frames with approximately 10% of auditory and 33% of visual neurons having receptive fields that were more consistent with an eye- than a head-centered frame of reference and 23 and 18% having receptive fields that were more consistent with a head- than an eye-centered frame of reference, leaving a large fraction of both visual and auditory response patterns inconsistent with both head- and eye-centered reference frames. The results were similar to the reference frame we have previously found for auditory stimuli in the inferior colliculus and core auditory cortex. The correspondence between the visual and auditory receptive fields of individual neurons was weak. Nevertheless, the visual and auditory responses were sufficiently well correlated that a simple one-layer network constructed to calculate target location from the activity of the neurons in our sample performed successfully for auditory targets even though the weights were fit based only on the visual responses. We interpret these results as suggesting that although the representations of space in areas LIP and MIP are not easily described within the conventional conceptual framework of reference frames, they nevertheless process visual and auditory spatial information in a similar fashion.

Acoustic Stimulation↗

The organization of central auditory pathways in a reptile, Iguana iguana.

The present experiments were designed to trace the central auditory pathways in an extant reptile, the New Worlkd lizard--Iguana iguana, utilizing anterograde axonal degeneration stained by the Fink-Heimer ('67) method and the retrograde axonal transport of horseradish peroxidase (LaVail and LaVail, '74). Beginning with the projections of the auditory portion of the VIIIth nerve, the ascending pathways were traced through successive relay nuclei to the telencephalon. The auditory portion of the VIIIth nerve projects to two nuclei in the dorsomedial medulla-nucleus angularis and nucleus magnocellularis medialis. These two nuclei together with a third cll group, nucleus magnocellularis lateralis (intercalated between nucleus angularis and nucleus magnocellularis medialis), have been referred to as the auditory tubercle in previous studies (cf. Miller, '75). The axonal degeneration following large lesions of the auditory tubercle and small lesions of nucleus angularis demonstrated the second order auditory pathways. Fibers leave nucleus angularis ventrally and travel to the ventral surface of the medulla where they cross the midline and ascend to the midbrain in pathways resembling the trapezoid body and the lateral lemniscus of mammals. Along these pathways, terminal arborizations of some fibers were seen in three lower brainstem nuclei while other fibers ascent to the midbrain and terminate in the central nucleus of the torus semicircularis. Experiments in which horseradish peroxidase injections were made in the torus semicircularis demonstrated that nucleus angularis is a primary source of second order auditory fibers to the midbrain and, in addition, that two of the lower brainstem targets of the auditory tubercle project to the torus semicircularis. These lower brainstem pathways were shown to be associated with the auditory system by electrophysiologically recording sound-evoked responses from clusters of cells in the torus semicircularis. Ascending fibers arising from the central nucleus of the torus semicircularis were followed rostrally where they entered the dorsal thalamus and terminated throughout nucleus medialis. Finally, a thalamotelencephalic auditory pathway was traced from nucleus medialis into the lateral forebrain bundle. Terminations of this pathway from nucleus medialis were seen in the medial dorsal ventricular ridge and in the striatum. It was concluded that the ascending auditory pathways of the iguana bear a remarkable resemblance to both the mammalian and avian auditory pathways from the level of the first order neurons in the VIIIth nerve to the level of the telencephalon. At the same time, there are important specializations of the auditory system in birds and mammals such as the development of particular lower brainstem nuclei. Nevertheless, a basic plan for the organization of the auditory system in terrestrial vertebrates can be recognized which invites comparisons with the vertebrate classes that remained in aquatic habitats...

Animals↗

Bilirubin and the auditory system.

The auditory system is highly sensitive to bilirubin toxicity. Damage to the auditory nervous system includes auditory neuropathy or auditory dyssynchrony and auditory processing problems which may occur with or without deafness, hearing loss. Auditory dysfunction may occur in children with or without other signs of classical kernicterus. Bilirubin selectively damages the brainstem auditory nuclei, and may also damage the auditory nerve and spiral ganglion containing cell bodies of primary auditory neurons. The inner ear, thalamic and cortical auditory pathways appear to be spared. Noninvasive auditory neurophysiological tests such as the auditory brainstem response (ABR) or brainstem auditory response (BAER) play an important role in the early detection of bilirubin-induced auditory and central nervous system dysfunction in the neonate.

Animals↗

Auditory brainstem implant as a salvage treatment after unsuccessful cochlear implantation.

OBJECTIVE: The present article investigates on an individual basis the performance achieved with the auditory brainstem implant in patients who had been treated unsuccessfully with a cochlear implant. STUDY DESIGN: An intrasubject comparison between results achieved with the cochlear implant and the auditory brainstem implant is reported. SETTING: Tertiary referral care. PATIENTS: Five subjects were fitted with an auditory brainstem implant in our department because of the poor results achieved with cochlear implants. Two were children, one with bilateral cochlear nerve aplasia and one suffering from auditory neuropathy. Three were adults with complete cochlear ossification. INTERVENTION: A retrosigmoid approach was used in all subjects. Electrically evoked auditory brainstem responses and neural response telemetry were used to monitor electrode positioning. RESULTS: No complications were observed due to implantation surgery or related to activation or long-term use of the auditory brainstem implant. Auditory sensations were induced in all patients with varying numbers of electrodes (from 9-16). In all three adults, the cochlear implant did not allow either word/sentence discrimination or speech tracking, whereas the auditory brainstem implant permitted discrimination of two- or three-syllable words with scores from 85 to 100%. In the two adults with a follow-up of 5 and 6 months after auditory brainstem implant activation, the open-set sentence recognition scores (auditory-only mode) were 70% and 100%, respectively, and the speech-tracking scores were 27 and 40 words/min, respectively. One patient with a follow-up of only 3 months scored 0% in both sentence recognition and speech tracking. The two children who had achieved no hearing ability with the cochlear implant were already able to detect sounds and words as early as 2 months after activation of the auditory brainstem implant and are showing progressive improvement in their performance. CONCLUSION: Auditory brainstem implantation may be a very powerful rehabilitative treatment after cochlear implant failure. The possibility of using the auditory brainstem implant as first-choice therapy in some categories of deaf patients (e.g., subjects with auditory neuropathy or cochlear ossification) who are currently treated with cochlear implantation is discussed.

Adult↗

Involvement of subcortical and cortical afferents to the lateral nucleus of the amygdala in fear conditioning measured with fear-potentiated startle in rats trained concurrently with auditory and visual conditioned stimuli.

The goal of this work was to test the involvement, in fear conditioning, of afferents to the lateral nucleus of the amygdala originating from the auditory thalamus, auditory cortex, and perirhinal area. The acoustic startle reflex was used as the behavioral index of conditioning because it is reliably enhanced in the presence of a conditioned stimulus (CS) previously paired with a footshock. Auditory and visual CSs were used to assess the modality specificity of lesions to the above brain areas. Pre- or posttraining lesions of the entire auditory thalamus including the ventral, dorsal, and medial divisions of the medial geniculate body, the posterior intralaminar nucleus, and the suprageniculate nucleus, completely blocked fear-potentiated startle to the auditory CS, but had no effect on fear-potentiated startle to the visual CS. Posttraining lesions mostly restricted to the ventral and dorsal divisions of the medial geniculate body specifically disrupted fear-potentiated startle to the auditory CS. However, retraining in rats sustaining ventral and dorsal medial geniculate body lesions led to reliable fear-potentiated startle to the auditory CS. Posttraining lesions mostly restricted to the medial division of the medial geniculate body, posterior intralaminar, and suprageniculate nuclei did not disrupt fear-potentiated startle. These results indicate that the auditory thalamus is specifically involved, through either its direct or indirect amygdaloid afferents, in fear conditioning to auditory CSs. Pre- or posttraining lesions mostly restricted to the primary auditory cortex did not reliably attenuate fear-potentiated startle to the auditory or visual CSs. Extensive posttraining lesions of the perirhinal area (including secondary auditory cortices), but not its rostral aspect alone, blocked fear-potentiated startle to both CSs. However, reliable potentiated startle was observed to both CSs following similarly extensive pretraining lesions of the perirhinal area. Because post- but not pretraining lesions of the perirhinal area blocked fear-potentiated startle nonspecifically, at least with regard to auditory and visual CSs, the results are consistent with the involvement of the perirhinal area in general memory functions such as information storage or retrieval. Alternatively, the secondary auditory and/or perirhinal cortices might function as multimodal sensory relays to the amygdala.

Acoustic Stimulation↗

NMDA and non-NMDA glutamate receptors in auditory transmission in the barn owl inferior colliculus.

The pharmacology of auditory responses in the inferior colliculus (IC) of the barn owl was investigated by iontophoresis of excitatory amino acid receptor antagonists into two different functional subdivisions of the IC, the external nucleus (ICx) and the lateral shell of the central nucleus (lateral shell), both of which carry out important computations in the processing of auditory spatial information. Combined application of the NMDA receptor antagonist 2-amino-5-phosphonovaleric acid (AP5) and the non-NMDA receptor antagonist 6-cyano-5-nitroquinoxaline-2,3-dione (CNQX) significantly reduced auditory-evoked spikes at all sites in these two subdivisions, and completely eliminated responses at many locations. This suggests that excitatory amino acid receptors mediate the bulk, if not all, of auditory responses in the ICx and lateral shell. NMDA and non-NMDA receptors contributed differently to auditory responses in the two subdivisions. In the ICx, AP5 significantly reduced the number of auditory-evoked spikes at every site tested. On average, AP5 eliminated 55% of auditory-evoked spikes at multiunit sites and 64% at single-unit sites in this structure. In contrast, in the lateral shell, AP5 significantly reduced responses at less than half the sites tested, and, on average, AP5 eliminated only 19% of spikes at multiunit sites and 25% at single-unit sites. When the magnitude of response blockade produced by AP5 at individual multiunit sites was normalized to adjust for site-to-site differences in the efficacy of iontophoresed AP5 and CNQX, AP5 blockade was still significantly greater in the ICx than the lateral shell. CNQX application strongly reduced responses in both subdivisions. These data suggest that NMDA receptor currents make a major contribution to auditory responses in the ICx, while they make only a small contribution to auditory responses in the lateral shell. Non-NMDA receptor currents, on the other hand, contribute to auditory responses in both subdivisions, and mediate the bulk of auditory transmission in the lateral shell. The time course of the NMDA receptor contribution to ICx auditory responses and the dependence of this contribution on stimulus level were both examined in detail. AP5 preferentially blocked spikes late in ICx auditory responses, while CNQX blocked spikes equally throughout the responses. This pattern is consistent with a simple model in which slow NMDA receptor currents and faster non-NMDA receptor currents are both activated by auditory inputs to ICx neurons.

2-Amino-5-phosphonovalerate↗

Mismatch negativity in the neurophysiologic/behavioral evaluation of auditory processing deficits: a case study.

The subject of this case report is an 18-year-old woman with grossly abnormal auditory brain stem response (ABR), normal peripheral hearing, and specific behavioral auditory processing deficits. Auditory middle latency responses (MLRs) and cortical potentials N1, P2, and P300 were intact. The mismatch negativity (MMN) was normal in response to certain synthesized speech stimuli and impaired to others--consistent with her behavioral discrimination of these stimuli. Behavioral tests of auditory processing were consistent with auditory brain stem dysfunction. A neuropsychological evaluation revealed normal intellectual and academic performance. The subject was in her first year of college at the time of the evaluation. This case study is important because: (1) Although there have been several reports of absent/abnormal ABR with preserved peripheral hearing and deficits in auditory processing, little is known about the specific nature of the auditory deficits experienced by these individuals. Such information may be valuable to the clinical management of patients with this constellation of findings. (2) Of interest is the information that the mismatch negativity (MMN) cortical event-related potential can bring to the evaluation of patients with auditory processing deficits. The MMN reflects central auditory processing of small acoustic differences and may provide an objective measure of auditory discrimination. (3) From a theorectical standpoint, a patient with neural deficits affecting specific components of the auditory pathway provides insight into the relationship between evoked potentials and physiological mechanisms of auditory processing. How do various components of the auditory pathway contribute to speech discrimination? How might evoked potentials reflect the processes underlying the neural coding of specific features of speech stimuli such as timing and spectral cues?

Achievement↗

[Auditory training in workshops: group therapy option].

BACKGROUND: auditory training in groups. AIM: to verify in a group of individuals with mental retardation the efficacy of auditory training in a workshop environment. METHOD a longitudinal prospective study with 13 mentally retarded individuals from the Associação de Pais e Amigos do Excepcional (APAE) of Congonhas divided in two groups: case (n=5) and control (n=8) and who were submitted to ten auditory training sessions after verifying the integrity of the peripheral auditory system through evoked otoacoustic emissions. Participants were evaluated using a specific protocol concerning the auditory abilities (sound localization, auditory identification, memory, sequencing, auditory discrimination and auditory comprehension) at the beginning and at the end of the project. Data (entering, processing and analyses) were analyzed by the Epi Info 6.04 software. RESULTS: the groups did not differ regarding aspects of age (mean = 23.6 years) and gender (40% male). In the first evaluation both groups presented similar performances. In the final evaluation an improvement in the auditory abilities was observed for the individuals in the case group. When comparing the mean number of correct answers obtained by both groups in the first and final evaluations, a statistically significant result was obtained for sound localization (p=0.02), auditory sequencing (p=0.006) and auditory discrimination (p=0.03). CONCLUSION: group auditory training demonstrated to be effective in individuals with mental retardation, observing an improvement in the auditory abilities. More studies, with a larger number of participants, are necessary in order to confirm the findings of the present research. These results will help public health professionals to reanalyze the theory models used for therapy, so that they can use specific methods according to individual needs, such as auditory training workshops.

Acoustic Stimulation↗

Parvalbumin and calbindin are differentially distributed within primary and secondary subregions of the mouse auditory forebrain.

The calcium binding proteins parvalbumin and calbindin are thought to differentially regulate physiological functions and often show complementary distributions in the CNS. Our goal was to determine parvalbumin and calbindin distributions in the different subdivisions of mouse auditory thalamus and auditory cortex. Following fixation, FVB mouse brains (postnatal days 38-80) were sectioned along coronal and horizontal planes, then processed for parvalbumin and calbindin immunohistochemistry (antibodies: parvalbumin pa-235, calbindin-d-28k cl-300). Strong complementary differences in calcium binding protein distributions were found in mouse auditory thalamus. The ventral division of the medial geniculate, which is the principal relay to primary auditory cortex, exhibited dense parvalbumin but weak calbindin immunoreactivity. In contrast, most of the 'secondary' auditory thalamic regions surrounding the ventral division showed strong calbindin and lighter parvalbumin levels. Thus, the mouse auditory thalamus is composed of a parvalbumin positive 'core' surrounded by a calbindin positive 'shell'. Parvalbumin immunoreactivity was also more prominent in the primary auditory cortex than in the secondary belt auditory cortex. Calbindin immunoreactivity in auditory cortex was less clearly divided along primary/secondary lines, especially in supragranular layers. However, within infragranular layers, there was heavier staining in belt areas than in primary auditory cortex. In auditory thalamus, parvalbumin labeling was largely confined to the neuropil, whereas calbindin labeling involved somata and neuropil. In auditory cortex, somata and neuropil were positive for both proteins.In summary, the calcium binding proteins parvalbumin and calbindin were found to be differentially distributed within the primary and non-primary regions of mouse auditory forebrain. These differences in protein distribution may contribute to the distinct types of physiological responses that occur in the primary vs. non-primary areas.

Animals↗

Tonotopic organization and parcellation of auditory cortex in the FM-bat Carollia perspicillata.

In the short-tailed fruit bat (Carollia perspicillata), the auditory cortex was localized autoradiographically and studied electrophysiologically in detail by using metal microelectrodes and 10-ms tone stimuli. Because, in the weakly-anaesthetized preparation, neuronal responses to pure-tones were even found throughout the non-primary auditory cortex, characteristic frequencies and minimum thresholds of neuron clusters (multiunits) could be mapped consistently and used to define auditory cortical fields conventionally (i.e. as in studies of auditory cortex of non-echolocating mammals). Thus, within the electrophysiologically demarcated auditory cortex, six auditory fields were defined by criteria, as for example a gradient of characteristic frequencies (primary auditory cortex, AI; anterior auditory field, AAF; secondary auditory cortex, AII), reversal of the gradient across the field border (AI, AAF), uniform representation of a restricted band of frequencies (i.e. > 60 kHz; high-frequency fields I and II, HFI and HFII), and transition from low to high minimum thresholds or vice versa [dorsoposterior field (DP), AII, HFI, HFII]. As supportive evidence for the distinction of these auditory cortical fields, differences in neuronal response properties were also used. In comparison with other mammals (e.g. cat and mouse), both the relative position of the auditory fields (mainly AI, AAF, DP and AII) and the representational principles for sound parameters within these forebrain areas seem to reflect a 'fundamental plan' (discussion below) of mammalian auditory cortical organization. Two coherent dorsally displaced high-frequency representations (HFI, HFII) covering approximately 40% of the total auditory cortical surface seem particularly suited for the processing of the dominant biosonar second and third harmonic of this species, and hence can be regarded as an adaptation for echolocation.

Acoustic Stimulation↗

Auditory responses in multiple sensorimotor song system nuclei are co-modulated by behavioral state.

Auditory responsiveness in nucleus HVC, a high-order sensorimotor area of the avian song system, is modulated by changes in behavioral state. Modulation is not observed in the primary thalamo-recipient auditory area Field L, the indirect source of auditory input to HVC. In this study, we show that auditory responsiveness in nucleus interfacialis (NIf), the immediate auditory afferent to HVC, is modulated by behavioral state. While auditory responsiveness is generally greater in NIf during wakefulness and in HVC during sedation, simultaneous recordings reveal a co-variation of auditory response magnitude. This co-variation is observed both in awake birds, where responses are spontaneously variable, and in sedated birds during manipulations of arousal levels. Auditory responses in NIf and HVC, which are selective for the bird's own song (BOS) during sedation, become predominantly unselective during wakefulness. This loss of selectivity is accompanied by a decrease in the similarity of NIf and HVC response patterns. To explore the role of NIf in shaping HVC auditory responses, we pharmacologically manipulated NIf while recording in HVC. Injection of the GABA(A) agonist muscimol into NIf eliminated most spontaneous activity and all auditory responses in the ipsilateral HVC, while injections of the GABA(A) antagonist bicuculline increased HVC auditory responsiveness and selectivity. These findings indicate that HVC is not the initial site of behavioral state-dependent modulation in the song system. Together with the suppression of HVC auditory responses by muscimol in NIf, these results suggest that NIf plays an important role in the flow of auditory information to HVC.

Acoustic Stimulation↗

Auditory dysfunction in stroke.

The auditory and vestibular systems share the same end organ and cranial nerve, yet vestibular signs and symptoms are common with stroke, whereas hearing disturbances are much less frequent. Several reasons would appear to account for this striking dissimilarity. One is that the auditory pathway is less ubiquitous than the vestibular pathways. The likelihood that a stroke involves the auditory pathway is, therefore, less on this basis alone. A second difference, to our knowledge not previously reported, is that the auditory pathway is often spared by the most common strokes. This is because major parts of the auditory pathway, such as the cochlear nucleus, inferior colliculus and medial geniculate body, have multiple sources of blood supply. A third well-recognized factor is the redundancy of the central auditory system and its strong bilateral representation above the level of the cochlear nuclei. Consequently, rostral to the cochlear nuclei gross deficits in hearing, such as those measured by standard pure-tone audiometry and speech discrimination, only occur if lesions are bilateral. Furthermore, widespread bilateral lesions of the auditory system typically render the patient unable to respond or are incompatible with life. In contrast, language disorders are more frequent because language is usually unilaterally represented in the cortex. Certainly, cerebral stroke often includes the auditory system, resulting in various types of auditory disorders, but most hemispherical lesions produce subtle hearing dysfunctions that can only be detected with sophisticated psychoacoustic and electrophysiological testing. The purpose of this review is to provide an overview of the auditory system and its blood supply and to review how auditory processing can be affected by stroke. Psychoacoustic and electrophysiological test procedures for identifying lesions in the central auditory system are described. The literature of hearing disorders due to stroke is reviewed and illustrative cases are presented.

Audiometry, Pure-Tone↗

Evidence of functional connectivity between auditory cortical areas revealed by amplitude modulation sound processing.

The human auditory cortex includes several interconnected areas. A better understanding of the mechanisms involved in auditory cortical functions requires a detailed knowledge of neuronal connectivity between functional cortical regions. In human, it is difficult to track in vivo neuronal connectivity. We investigated the interarea connection in vivo in the auditory cortex using a method of directed coherence (DCOH) applied to depth auditory evoked potentials (AEPs). This paper presents simultaneous AEPs recordings from insular gyrus (IG), primary and secondary cortices (Heschl's gyrus and planum temporale), and associative areas (Brodmann area [BA] 22) with multilead intracerebral electrodes in response to sinusoidal modulated white noises in 4 epileptic patients who underwent invasive monitoring with depth electrodes for epilepsy surgery. DCOH allowed estimation of the causality between 2 signals recorded from different cortical sites. The results showed 1) a predominant auditory stream within the primary auditory cortex from the most medial region to the most lateral one whatever the modulation frequency, 2) unidirectional functional connection from the primary to secondary auditory cortex, 3) a major auditory propagation from the posterior areas to the anterior ones, particularly at 8, 16, and 32 Hz, and 4) a particular role of Heschl's sulcus dispatching information to the different auditory areas. These findings suggest that cortical processing of auditory information is performed in serial and parallel streams. Our data showed that the auditory propagation could not be associated to a unidirectional traveling wave but to a constant interaction between these areas that could reflect the large adaptive and plastic capacities of auditory cortex. The role of the IG is discussed.

Acoustic Stimulation↗

Song system auditory responses are stable and highly tuned during sedation, rapidly modulated and unselective during wakefulness, and suppressed by arousal.

We used auditory responsiveness in the avian song system to investigate the complex relationship between behavioral state and sensory processing in a high-order sensorimotor brain area. We present evidence from recordings in awake, anesthetized, and sleeping male zebra finches (Taeniopygia guttata) that auditory responsiveness in nucleus HVc is profoundly affected by changes in behavioral state. In anesthetized and sleeping birds, auditory responses were characterized by an increase in firing rate that was selective for the bird's own song (BOS) and highly stable over time. In contrast, HVc responses during wakefulness were extremely variable and transitioned between undetectable and robust levels over short intervals. Surprisingly, auditory responses in awake birds were not selective for the BOS stimulus. The variability of HVc auditory responses in awake birds suggests that, as in mammals, wakefulness is not a uniform behavioral state. Rather, auditory responsiveness likely is continually influenced by variables such as arousal state. We therefore developed several experimental paradigms in which we could manipulate arousal levels during auditory stimulus presentation. In all cases, arousal suppressed HVc auditory responses. This effect was specific to the song system, as auditory responses in Field L, a primary auditory area that is a source of auditory input to HVc, were unaffected. While arousal acts as a negative regulator of HVc auditory responsiveness, the presence and variability of the responses observed in awake, alert birds suggests that other mechanisms, such as attention, may enhance auditory responsiveness. The interplay between behavioral state and sensory processing may regulate song system responsiveness according to the bird's behavioral and social context.

Acoustic Stimulation↗

Frequency and space representation in the primary auditory cortex of the frequency modulating bat Eptesicus fuscus.

1. Frequency and space representation in the auditory cortex of the big brown bat, Eptesicus fuscus, were studied by recording responses of 223 neurons to acoustic stimuli presented in the bat's frontal auditory space. 2. The majority of the auditory cortical neurons were recorded at a depth of less than 500 microns with a response latency between 8 and 20 ms. They generally discharged phasically and had nonmonotonic intensity-rate functions. The minimum threshold, (MT) of these neurons was between 8 and 82 dB sound pressure level (SPL). Half of the cortical neurons showed spontaneous activity. All 55 threshold curves are V-shaped and can be described as broad, intermediate, or narrow. 3. Auditory cortical neurons are tonotopically organized along the anteroposterior axis of the auditory cortex. High-frequency-sensitive neurons are located anteriorly and low-frequency-sensitive neurons posteriorly. An overwhelming majority of neurons were sensitive to a frequency range between 30 and 75 kHz. 4. When a sound was delivered from the response center of a neuron on the bat's frontal auditory space, the neuron had its lowest MT. When the stimulus amplitude was increased above the MT, the neuron responded to sound delivered within a defined spatial area. The response center was not always at the geometric center of the spatial response area. The latter also expanded with stimulus amplitude. High-frequency-sensitive neurons tended to have smaller spatial response areas than low-frequency-sensitive neurons. 5. Response centers of all 223 neurons were located between 0 degrees and 50 degrees in azimuth, 2 degrees up and 25 degrees down in elevation of the contralateral frontal auditory space. Response centers of auditory cortical neurons tended to move toward the midline and slightly downward with increasing best frequency. 6. Auditory space representation appears to be systematically arranged according to the tonotopic axis of the auditory cortex. Thus, the lateral space is represented posteriorly and the middle space anteriorly. Space representation, however, is less systematic in the vertical direction. 7. Auditory cortical neurons are columnarly organized. Thus, the BFs, MTs, threshold curves, azimuthal location of response centers, and auditory spatial response areas of neurons sequentially isolated from an orthogonal electrode penetration are similar.

Acoustic Stimulation↗

The role of predictive models in the formation of auditory streams.

Sounds provide us with useful information about our environment which complements that provided by other senses, but also poses specific processing problems. How does the auditory system distentangle sounds from different sound sources? And what is it that allows intermittent sound events from the same source to be associated with each other? Here we review findings from a wide range of studies using the auditory streaming paradigm in order to formulate a unified account of the processes underlying auditory perceptual organization. We present new computational modelling results which replicate responses in primary auditory cortex [Fishman, Y.I., Arezzo, J.C., Steinschneider, M., 2004. Auditory stream segregation in monkey auditory cortex: effects of frequency separation, presentation rate, and tone duration. J. Acoust. Soc. Am. 116, 1656-1670; Fishman, Y. I., Reser, D. H., Arezzo, J.C., Steinschneider, M., 2001. Neural correlates of auditory stream segregation in primary auditory cortex of the awake monkey. Hear. Res. 151, 167-187] to tone sequences. We also present the results of a perceptual experiment which confirm the bi-stable nature of auditory streaming, and the proposal that the gradual build-up of streaming may be an artefact of averaging across many subjects [Pressnitzer, D., Hupé, J. M., 2006. Temporal dynamics of auditory and visual bi-stability reveal common principles of perceptual organization. Curr. Biol. 16(13), 1351-1357.]. Finally we argue that in order to account for all of the experimental findings, computational models of auditory stream segregation require four basic processing elements; segregation, predictive modelling, competition and adaptation, and that it is the formation of effective predictive models which allows the system to keep track of different sound sources in a complex auditory environment.

Acoustic Stimulation↗