The role of the right parietal cortex in sound localization: a chronometric single pulse transcranial magnetic stimulation study.

Auditory spatial functions, including the ability to discriminate between the positions of nearby sound sources, are subserved by a large temporo-parieto-frontal network. With the aim of determining whether and when the parietal contribution is critical for auditory spatial discrimination, we applied single pulse transcranial magnetic stimulation on the right parietal cortex 20, 80, 90 and 150 ms post-stimulus onset while participants completed a two-alternative forced choice auditory spatial discrimination task in the left or right hemispace. Our results reveal that transient TMS disruption of right parietal activity impairs spatial discrimination when applied at 20 ms post-stimulus onset for sounds presented in the left (controlateral) hemispace and at 80 ms for sounds presented in the right hemispace. We interpret our finding in terms of a critical role for controlateral temporo-parietal cortices over initial stages of the building-up of auditory spatial representation and for a right hemispheric specialization in integrating the whole auditory space over subsequent, higher-order processing stages.

The spatio-temporal brain dynamics of processing and integrating sound localization cues in humans.

Interaural intensity and time differences (IID and ITD) are two binaural auditory cues for localizing sounds in space. This study investigated the spatio-temporal brain mechanisms for processing and integrating IID and ITD cues in humans. Auditory-evoked potentials were recorded, while subjects passively listened to noise bursts lateralized with IID, ITD or both cues simultaneously, as well as a more frequent centrally presented noise. In a separate psychophysical experiment, subjects actively discriminated lateralized from centrally presented stimuli. IID and ITD cues elicited different electric field topographies starting at approximately 75 ms post-stimulus onset, indicative of the engagement of distinct cortical networks. By contrast, no performance differences were observed between IID and ITD cues during the psychophysical experiment. Subjects did, however, respond significantly faster and more accurately when both cues were presented simultaneously. This performance facilitation exceeded predictions from probability summation, suggestive of interactions in neural processing of IID and ITD cues. Supra-additive neural response interactions as well as topographic modulations were indeed observed approximately 200 ms post-stimulus for the comparison of responses to the simultaneous presentation of both cues with the mean of those to separate IID and ITD cues. Source estimations revealed differential processing of IID and ITD cues initially within superior temporal cortices and also at later stages within temporo-parietal and inferior frontal cortices. Differences were principally in terms of hemispheric lateralization. The collective psychophysical and electrophysiological results support the hypothesis that IID and ITD cues are processed by distinct, but interacting, cortical networks that can in turn facilitate auditory localization.

Representation of sound localization cues in the auditory thalamus of the barn owl

Barn owls can localize a sound source using either the map of auditory space contained in the optic tectum or the auditory forebrain. The auditory thalamus, nucleus ovoidalis (N.Ov), is situated between these two auditory areas, and its inactivation precludes the use of the auditory forebrain for sound localization. We examined the sources of inputs to the N.Ov as well as their patterns of termination within the nucleus. We also examined the response of single neurons within the N.Ov to tonal stimuli and sound localization cues. Afferents to the N.Ov originated with a diffuse population of neurons located bilaterally within the lateral shell, core, and medial shell subdivisions of the central nucleus of the inferior colliculus. Additional afferent input originated from the ipsilateral ventral nucleus of the lateral lemniscus. No afferent input was provided to the N.Ov from the external nucleus of the inferior colliculus or the optic tectum. The N.Ov was tonotopically organized with high frequencies represented dorsally and low frequencies ventrally. Although neurons in the N.Ov responded to localization cues, there was no apparent topographic mapping of these cues within the nucleus, in contrast to the tectal pathway. However, nearly all possible types of binaural response to sound localization cues were represented. These findings suggest that in the thalamo-telencephalic auditory pathway, sound localization is subserved by a nontopographic representation of auditory space.

Roaring lions and chirruping lemurs: How the brain encodes sound objects in space.

The dual-stream model of auditory processing postulates separate processing streams for sound meaning and for sound location. The present review draws on evidence from human behavioral and activation studies as well as from lesion studies to argue for a position-linked representation of sound objects that is distinct both from the position-independent representation within the ventral/What stream and from the explicit sound localization processing within the dorsal/Where stream.

A spiking neural network implementation of sound localisation

The focus of this paper is the implementation of a spiking neural network to achieve sound localization; the model is based on the influential short paper by Jeffress in 1948. The SNN has a two-layer topology which can accommodate a limited number of angles in the azimuthal plane. The model accommodates multiple inter-neuron connections with associated delays, and a supervised STDP algorithm is applied to select the optimal pathway for sound localization. Also an analysis of previous relevant work in the area of auditory modelling supports this research.

Modularity of auditory spatial representations

Abstract : Auditory spatial functions are of crucial importance in everyday life. Determining the origin of sound sources in space plays a key role in a variety of tasks including orientation of attention, disentangling of complex acoustic patterns reaching our ears in noisy environments. Following brain damage, auditory spatial processing can be disrupted, resulting in severe handicaps. Complaints of patients with sound localization deficits include the inability to locate their crying child or being over-loaded by sounds in crowded public places. Yet, the brain bears a large capacity for reorganization following damage and/or learning. This phenomenon is referred as plasticity and is believed to underlie post-lesional functional recovery as well as learning-induced improvement. The aim of this thesis was to investigate the organization and plasticity of different aspects of auditory spatial functions. Overall, we report the outcomes of three studies: In the study entitled "Learning-induced plasticity in auditory spatial representations" (Spierer et al., 2007b), we focused on the neurophysiological and behavioral changes induced by auditory spatial training in healthy subjects. We found that relatively brief auditory spatial discrimination training improves performance and modifies the cortical representation of the trained sound locations, suggesting that cortical auditory representations of space are dynamic and subject to rapid reorganization. In the same study, we tested the generalization and persistence of training effects over time, as these are two determining factors in the development of neurorehabilitative intervention. In "The path to success in auditory spatial discrimination" (Spierer et al., 2007c), we investigated the neurophysiological correlates of successful spatial discrimination and contribute to the modeling of the anatomo-functional organization of auditory spatial processing in healthy subjects. We showed that discrimination accuracy depends on superior temporal plane (STP) activity in response to the first sound of a pair of stimuli. Our data support a model wherein refinement of spatial representations occurs within the STP and that interactions with parietal structures allow for transformations into coordinate frames that are required for higher-order computations including absolute localization of sound sources. In "Extinction of auditory stimuli in hemineglect: space versus ear" (Spierer et al., 2007a), we investigated auditory attentional deficits in brain-damaged patients. This work provides insight into the auditory neglect syndrome and its relation with neglect symptoms within the visual modality. Apart from contributing to a basic understanding of the cortical mechanisms underlying auditory spatial functions, the outcomes of the studies also contribute to develop neurorehabilitation strategies, which are currently being tested in clinical populations.

Hemispheric competence for auditory spatial representation.

Sound localization relies on the analysis of interaural time and intensity differences, as well as attenuation patterns by the outer ear. We investigated the relative contributions of interaural time and intensity difference cues to sound localization by testing 60 healthy subjects: 25 with focal left and 25 with focal right hemispheric brain damage. Group and single-case behavioural analyses, as well as anatomo-clinical correlations, confirmed that deficits were more frequent and much more severe after right than left hemispheric lesions and for the processing of interaural time than intensity difference cues. For spatial processing based on interaural time difference cues, different error types were evident in the individual data. Deficits in discriminating between neighbouring positions occurred in both hemispaces after focal right hemispheric brain damage, but were restricted to the contralesional hemispace after focal left hemispheric brain damage. Alloacusis (perceptual shifts across the midline) occurred only after focal right hemispheric brain damage and was associated with minor or severe deficits in position discrimination. During spatial processing based on interaural intensity cues, deficits were less severe in the right hemispheric brain damage than left hemispheric brain damage group and no alloacusis occurred. These results, matched to anatomical data, suggest the existence of a binaural sound localization system predominantly based on interaural time difference cues and primarily supported by the right hemisphere. More generally, our data suggest that two distinct mechanisms contribute to: (i) the precise computation of spatial coordinates allowing spatial comparison within the contralateral hemispace for the left hemisphere and the whole space for the right hemisphere; and (ii) the building up of global auditory spatial representations in right temporo-parietal cortices.

Auditory spatio-temporal brain dynamics and their consequences for multisensory interactions in humans.

Recent multisensory research has emphasized the occurrence of early, low-level interactions in humans. As such, it is proving increasingly necessary to also consider the kinds of information likely extracted from the unisensory signals that are available at the time and location of these interaction effects. This review addresses current evidence regarding how the spatio-temporal brain dynamics of auditory information processing likely curtails the information content of multisensory interactions observable in humans at a given latency and within a given brain region. First, we consider the time course of signal propagation as a limitation on when auditory information (of any kind) can impact the responsiveness of a given brain region. Next, we overview the dual pathway model for the treatment of auditory spatial and object information ranging from rudimentary to complex environmental stimuli. These dual pathways are considered an intrinsic feature of auditory information processing, which are not only partially distinct in their associated brain networks, but also (and perhaps more importantly) manifest only after several tens of milliseconds of cortical signal processing. This architecture of auditory functioning would thus pose a constraint on when and in which brain regions specific spatial and object information are available for multisensory interactions. We then separately consider evidence regarding mechanisms and dynamics of spatial and object processing with a particular emphasis on when discriminations along either dimension are likely performed by specific brain regions. We conclude by discussing open issues and directions for future research.

Organization and plasticity of auditory spatial representations

Spatial hearing refers to a set of abilities enabling us to determine the location of sound sources, redirect our attention toward relevant acoustic events, and recognize separate sound sources in noisy environments. Determining the location of sound sources plays a key role in the way in which humans perceive and interact with their environment. Deficits in sound localization abilities are observed after lesions to the neural tissues supporting these functions and can result in serious handicaps in everyday life. These deficits can, however, be remediated (at least to a certain degree) by the surprising capacity of reorganization that the human brain possesses following damage and/or learning, namely, the brain plasticity. In this thesis, our aim was to investigate the functional organization of auditory spatial functions and the learning-induced plasticity of these functions. Overall, we describe the results of three studies. The first study entitled "The role of the right parietal cortex in sound localization: A chronometric single pulse transcranial magnetic stimulation study" (At et al., 2011), study A, investigated the role of the right parietal cortex in spatial functions and its chronometry (i.e. the critical time window of its contribution to sound localizations). We concentrated on the behavioral changes produced by the temporarily inactivation of the parietal cortex with transcranial magnetic stimulation (TMS). We found that the integrity of the right parietal cortex is crucial for localizing sounds in the space and determined a critical time window of its involvement, suggesting a right parietal dominance for auditory spatial discrimination in both hemispaces. In "Distributed coding of the auditory space in man: evidence from training-induced plasticity" (At et al., 2013a), study B, we investigated the neurophysiological correlates and changes of the different sub-parties of the right auditory hemispace induced by a multi-day auditory spatial training in healthy subjects with electroencephalography (EEG). We report a distributed coding for sound locations over numerous auditory regions, particular auditory areas code specifically for precise parts of the auditory space, and this specificity for a distinct region is enhanced with training. In the third study "Training-induced changes in auditory spatial mismatch negativity" (At et al., 2013b), study C, we investigated the pre-attentive neurophysiological changes induced with a training over 4 days in healthy subjects with a passive mismatch negativity (MMN) paradigm. We showed that training changed the mechanisms for the relative representation of sound positions and not the specific lateralization themselves and that it changed the coding in right parahippocampal regions. - L'audition spatiale désigne notre capacité à localiser des sources sonores dans l'espace, de diriger notre attention vers les événements acoustiques pertinents et de reconnaître des sources sonores appartenant à des objets distincts dans un environnement bruyant. La localisation des sources sonores joue un rôle important dans la façon dont les humains perçoivent et interagissent avec leur environnement. Des déficits dans la localisation de sons sont souvent observés quand les réseaux neuronaux impliqués dans cette fonction sont endommagés. Ces déficits peuvent handicaper sévèrement les patients dans leur vie de tous les jours. Cependant, ces déficits peuvent (au moins à un certain degré) être réhabilités grâce à la plasticité cérébrale, la capacité du cerveau humain à se réorganiser après des lésions ou un apprentissage. L'objectif de cette thèse était d'étudier l'organisation fonctionnelle de l'audition spatiale et la plasticité induite par l'apprentissage de ces fonctions. Dans la première étude intitulé « The role of the right parietal cortex in sound localization : A chronometric single pulse study » (At et al., 2011), étude A, nous avons examiné le rôle du cortex pariétal droit dans l'audition spatiale et sa chronométrie, c'est-à- dire le moment critique de son intervention dans la localisation de sons. Nous nous sommes concentrés sur les changements comportementaux induits par l'inactivation temporaire du cortex pariétal droit par le biais de la Stimulation Transcrânienne Magnétique (TMS). Nous avons démontré que l'intégrité du cortex pariétal droit est cruciale pour localiser des sons dans l'espace. Nous avons aussi défini le moment critique de l'intervention de cette structure. Dans « Distributed coding of the auditory space : evidence from training-induced plasticity » (At et al., 2013a), étude B, nous avons examiné la plasticité cérébrale induite par un entraînement des capacités de discrimination auditive spatiale de plusieurs jours. Nous avons montré que le codage des positions spatiales est distribué dans de nombreuses régions auditives, que des aires auditives spécifiques codent pour des parties données de l'espace et que cette spécificité pour des régions distinctes est augmentée par l'entraînement. Dans « Training-induced changes in auditory spatial mismatch negativity » (At et al., 2013b), étude C, nous avons examiné les changements neurophysiologiques pré- attentionnels induits par un entraînement de quatre jours. Nous avons montré que l'entraînement modifie la représentation des positions spatiales entraînées et non-entrainées, et que le codage de ces positions est modifié dans des régions parahippocampales.

Temporal dynamics of auditory what and where processing in humans

Recent evidence suggests the human auditory system is organized,like the visual system, into a ventral 'what' pathway, devoted toidentifying objects and a dorsal 'where' pathway devoted to thelocalization of objects in space w1x. Several brain regions have beenidentified in these two different pathways, but until now little isknown about the temporal dynamics of these regions. We investigatedthis issue using 128-channel auditory evoked potentials(AEPs).Stimuli were stationary sounds created by varying interaural timedifferences and environmental real recorded sounds. Stimuli ofeach condition (localization, recognition) were presented throughearphones in a blocked design, while subjects determined theirposition or meaning, respectively.AEPs were analyzed in terms of their topographical scalp potentialdistributions (segmentation maps) and underlying neuronalgenerators (source estimation) w2x.Fourteen scalp potential distributions (maps) best explained theentire data set.Ten maps were nonspecific (associated with auditory stimulationin general), two were specific for sound localization and two werespecific for sound recognition (P-values ranging from 0.02 to0.045).Condition-specific maps appeared at two distinct time periods:;200 ms and ;375-550 ms post-stimulus.The brain sources associated with the maps specific for soundlocalization were mainly situated in the inferior frontal cortices,confirming previous findings w3x. The sources associated withsound recognition were predominantly located in the temporal cortices,with a weaker activation in the frontal cortex.The data show that sound localization and sound recognitionengage different brain networks that are apparent at two distincttime periods.References1. Maeder et al. Neuroimage 2001.2. Michel et al. Brain Research Review 2001.3. Ducommun et al. Neuroimage 2002.

Localisation sonore chez les aveugles : l'influence de l'âge de survenue de la cécité

Thèse numérisée par la Division de la gestion de documents et des archives de l'Université de Montréal

Adaptation aux changements induits aux indices spectraux de l’audition spatiale chez l’humain.

Dans le cas de perte auditive, la localisation spatiale est amoindrie et vient entraver la compréhension de la parole et ce, malgré le port de prothèses auditives. La présente étude modifie la forme de l’oreille externe d’individus à l’aide de silicone afin d’induire des changements aux indices spectraux (HRTFs), similaires à ceux causés par des prothèses auditives, et d’explorer les mécanismes perceptifs (visuel, spectral, ou tactile) permettant d’alterner d’un nouvel ensemble à l’ensemble originel de HRTFs une fois les prothèses enlevées. Les résultats démontrent que les participants s’adaptent aux nouveaux HRTFs à l’intérieur de quatre séances d’entraînement. Dès le retrait des prothèses, les participants reviennent à leur performance originale. Il n’est pas possible de conclure avec les données présentes si le changement d’un ensemble de HRTFs à un autre est influencé par un des mécanismes de rétroaction perceptuelle étudié. L’adaptation aux prothèses perdure jusqu’à quatre semaines après leur retrait.

Desempenho de escolares com distúrbio de aprendizagem e dislexia em testes de processamento auditivo

OBJETIVO: caracterizar e comparar, por meio de testes comportamentais, o processamento auditivo de escolares com diagnóstico interdisciplinar de (I) distúrbio da aprendizagem, (II) dislexia e (III) escolares com bom desempenho acadêmico. MÉTODOS: participaram deste estudo 30 escolares na faixa etária de 8 a 16 anos de idade, de ambos os gêneros, de 2ª a 4ª séries do ensino fundamental, divididos em três grupos: GI composto por 10 escolares com diagnóstico interdisciplinar de distúrbio de aprendizagem, GII: composto por 10 escolares com diagnóstico interdisciplinar de dislexia e GIII composto por 10 escolares sem dificuldades de aprendizagem, pareados segundo gênero e faixa etária com os grupos GI e GII. Foram realizadas avaliação audiológica e de processamento auditivo. RESULTADOS: os escolares de GIII apresentaram desempenho superior nos testes de processamento auditivo em relação aos escolares de GI e GII. GI apresentou desempenho inferior nas habilidades auditivas avaliadas para testes dicóticos de dígitos e dissílabos alternados, logoaudiometria pediátrica, localização sonora, memória verbal e não-verbal, ao passo que GII apresentou as mesmas alterações de GI, com exceção do teste de logoaudiometria pediátrica. CONCLUSÃO: os escolares com transtornos de aprendizagem apresentaram desempenho inferior nos testes de processamento auditivo, sendo que os escolares com distúrbio de aprendizagem apresentaram maior número de habilidades auditivas alteradas, em comparação com os escolares com dislexia, por terem apresentado atenção sustentada reduzida. O grupo de escolares com dislexia apresentou alterações decorrentes da dificuldade relacionada à codificação e decodificação de estímulos sonoros.

Processamento auditivo na perda auditiva unilateral: relato de caso

A perda auditiva unilateral representa grande risco para o atraso acadêmico, a comunicação, o desenvolvimento social e também para o processamento auditivo. Assim, o objetivo deste estudo foi avaliar as habilidades auditivas de localização, fechamento, figura-fundo, resolução temporal e ordenação temporal simples de um sujeito do gênero masculino, 17 anos de idade, com diagnóstico de perda auditiva sensorioneural unilateral de grau profundo, de causa idiopática, sem outros comprometimentos. O processo de avaliação constituiu da aplicação de um questionário, da realização da avaliação audiológica clínica convencional (audiometria tonal, logoaudiometria e imitanciometria) e de testes de processamento auditivo monóticos (SSI ipsilateral, Teste de fala filtrada) e dióticos (Localização sonora, Memória auditiva para sons verbais, Memória auditiva para sons não verbais, AFT-R). Apenas o teste de Localização sonora apresentou resultados alterados. Não foram relatadas queixas relacionadas à habilidade de localização sonora, atenção, discriminação e compreensão auditiva. No caso em estudo, a perda auditiva sensorioneural unilateral de grau profundo não pareceu restringir o desenvolvimento das habilidades do processamento auditivo avaliadas, exceto no que se refere à localização da fonte sonora.

Relação entre potenciais evocados auditivos de média latência e distúrbio de processamento auditivo: estudo de casos

The Auditory Evoked Middle Latency Response is one of the most promising objective tests in audiology and in revealing brain dysfunction and neuro-audiologic findings. The main advantages of its clinical use are precision and objectivity in evaluating children. This study aimed to analyze the auditory evoked middle latency response in two patients with auditory processing disorder and relate objective and behavioral measures. This case study was conducted in 2 patients (P1 = 12 years, female, P2 = 17 years old, male), both with the absence of sensory abnormalities, neurological and neuropsychiatric disorders. Both were submitted to anamnesis, inspection of the external ear canal, hearing test and evaluation of Auditory Evoked Middle latency Response. There was a significant association between behavioral test and objectives results. In the interview, there were complaints about the difficulty in listening in a noisy environment, sound localization, inattention, and phonological changes in writing and speaking, as confirmed by evaluation of auditory processing and Auditory Evoked Middle Latency Response. Changes were observed in the right decoding process hearing in both cases on the behavioral assessment of auditory processing; auditory evoked potential test middle latency shows that the right contralateral via response was deficient, confirming the difficulties of the patients in the assignment of meaning in acoustic information in a competitive sound condition at right, in both cases. In these cases it was shown the association between the results, but there is a need for further studies with larger sample population to confirm the data.