Rapid change of tonotopic maps in the human auditory cortex during pitch discrimination.

OBJECTIVE: To study early cognitive processes and hemispheric differences in the primary auditory cortex during selective attention. METHODS: We measured auditory evoked magnetic fields (AEFs) to 400 and 4000 Hz tone pips that were randomly presented at the right or left ear. Subjects paid attention to target stimuli during pitch (high or low) or laterality (left or right) discrimination tasks. In the control session, 400 or 4000 Hz tone alone was presented at the left or right ear. We calculated the location and strength of N100m dipole for 400 and 4000 Hz tones, based on the AEFs obtained from the hemisphere contralateral to the stimulated ear. RESULTS: N100m amplitude increased in both hemispheres in pitch or laterality discriminating conditions. N100m latency also shortened during selective attention. The N100m dipole distance between 400 and 4000 Hz tones was enlarged, especially in the right auditory cortex during pitch discrimination task, but was unchanged during the laterality discrimination task. CONCLUSIONS: We conclude that these dynamic changes in the N100m dipole reflect short-term plastic changes in the primary auditory cortex, supporting early selection models. SIGNIFICANCE: This work is the first to disclose short-term plastic changes during pitch discrimination in the human auditory cortex based on the analysis of magnetoencephalography.     

Dynamic movement of N100m dipoles in evoked magnetic field reflects sequential activation of isofrequency bands in human auditory cortex.

OBJECTIVE: To investigate spatiotemporal features of the isofrequency bands for 400 and 4000 Hz tones in human auditory cortex and on the hemispheric differences in the arrangement of isofrequency bands. METHODS: We recorded auditory evoked magnetic fields (AEFs) to 400 or 4000 Hz tone pips presented at right or left ear from 31 normal subjects. The dipole location for the N100m sources was successively calculated from the AEFs obtained from the hemisphere contralateral to the stimulated ear. RESULTS: In the right hemisphere, the current sources for 400 and 4000 Hz moved toward the anterolateral direction before the N100m peak, showing parallel arrangement of the isofrequency bands (4000 Hz in medial location). In the left hemisphere, the movement direction of 400 Hz dipoles was anterolateral, while that of 4000 Hz dipoles was lateral. CONCLUSIONS: This difference in the organization of isofrequency bands between right and left auditory cortices reflects distinct functional roles in auditory information processing such as pitch vs. language discrimination. SIGNIFICANCE: This work is the first to disclose isofrequency bands in human auditory cortex based on the analysis of magnetoencephalography.     

Brain-speech alignment enhances auditory cortical responses and speech perception

Asymmetry in auditory cortical oscillations could play a role in speech perception by fostering hemispheric triage of information across the two hemispheres. Due to this asymmetry, fast speech temporal modulations relevant for phonemic analysis could be best perceived by the left auditory cortex, while slower modulations conveying vocal and paralinguistic information would be better captured by the right one. It is unclear, however, whether and how early oscillation-based selection influences speech perception. Using a dichotic listening paradigm in human participants, where we provided different parts of the speech envelope to each ear, we show that word recognition is facilitated when the temporal properties of speech match the rhythmic properties of auditory cortices. We further show that the interaction between speech envelope and auditory cortices rhythms translates in their level of neural activity (as measured with fMRI). In the left auditory cortex, the neural activity level related to stimulus-brain rhythm interaction predicts speech perception facilitation. These data demonstrate that speech interacts with auditory cortical rhythms differently in right and left auditory cortex, and that in the latter, the interaction directly impacts speech perception performance.     

Hemispheric lateralization of cerebral blood-flow changes during selective listening to dichotically presented continuous speech

Regional cerebral blood flow (rCBF) was measured with positron emission tomography (PET) while subjects were selectively listening to continuous speech delivered to one ear and ignoring concurrent speech delivered to the opposite ear, as well as concurrent text or letter strings running on a screen. rCBF patterns associated with selective listening either to the left-ear or right-ear speech message were compared with each other and with rCBF patterns in two visual-attention conditions in which the subjects ignored both speech messages and either read the text or discriminated the meaningless letter strings moving on the screen. Attention to either speech message was associated with enhanced activity in the superior temporal cortex of the language-dominant left hemisphere, as well as in the superior and middle temporal cortex of the right hemisphere suggesting enhanced processing of prosodic features in the attended speech. Moreover, enhanced activity during attention to either speech message was observed in the right parietal areas known to have an important role in directing spatial attention. Evidence was also found for attentional tuning of the left and right auditory cortices to select information from the contralateral auditory hemispace.     

Inhibition of auditory cortical responses to ipsilateral stimuli during dichotic listening: evidence from magnetoencephalography

The present magnetoencephalography (MEG) study on auditory evoked magnetic fields (AEFs) was aimed at verifying whether during dichotic listening the contralateral auditory pathway inhibits the ipsilateral one, as suggested by behavioural and patient studies. Ten healthy subjects were given a randomized series of three complex tones (261, 293 and 391 Hz, 500 ms duration), which were delivered monotically and dichotically with different intensities [60, 70 or 80 dBA (audio decibels)]. MEG data were recorded from the right auditory cortex. Results showed that the M100 amplitude over the right auditory cortex increased progressively when tones of increasing intensity were provided at the ipsilateral (right) ear. This effect on M100 was abolished when a concurrent tone of constant intensity was delivered dichotically at the contralateral (left) ear, suggesting that the contralateral pathway inhibited the ipsilateral one. The ipsilateral inhibition was present only when the contralateral tone fundamental frequency was similar to the ipsilateral tone. It was proposed that the occlusion mechanism would be exerted in cortical auditory areas as the dichotic effects were observed at M100 but not M50 component. This is the first evidence showing a neurophysiological inhibition driven by the contralateral auditory pathway over the ipsilateral one during dichotic listening.     

Early unilateral cochlear implantation promotes mature cortical asymmetries in adolescents who are deaf

Unilateral cochlear implant (CI) stimulation establishes hearing to children who are deaf but compromises bilateral auditory development if a second implant is not provided within ～1.5 years. In this study we asked: 1) What are the cortical consequences of missing this early sensitive period once children reach adolescence? 2) What are the effects of unilateral deprivation on the pathways from the opposite ear? Cortical responses were recorded from 64‐cephalic electrodes within the first week of bilateral CI activation in 34 adolescents who had over 10 years of unilateral right CI experience and in 16 normal hearing peers. Cortical activation underlying the evoked peaks was localized to areas of the brain using beamformer imaging. The first CI evoked activity which was more strongly lateralized to the contralateral left hemisphere than normal, with abnormal recruitment of the left prefrontal cortex (involved in cognition/attention), left temporo‐parietal‐occipital junction (multi‐modal integration), and right precuneus (visual processing) region. CI stimulation in the opposite deprived ear evoked atypical cortical responses with abnormally large and widespread dipole activity across the cortex. Thus, using a unilateral CI to hear beyond the period of cortical maturation causes lasting asymmetries in the auditory system, requires recruitment of additional cortical areas to support hearing, and does little to protect the unstimulated pathways from effects of auditory deprivation. The persistence of this reorganization into maturity could signal a closing of a sensitive period for promoting auditory development on the deprived side. Hum Brain Mapp 37:135–152, 2016. © 2015 Wiley Periodicals, Inc.     

Contribution of the anterior insula to temporal auditory processing deficits in developmental dyslexia

Developmental dyslexia has been assumed to arise from general auditory deficits, compromising rapid temporal integration both of linguistic and nonlinguistic acoustic stimuli. Because the effort of auditory temporal processing of speech and nonspeech test materials may depend on presentation rate, fMRI measurements were performed in dyslexics and controls during passive listening to series of syllable and click sounds, using a parametric approach. Controls showed a decrease of hemodynamic brain activation within the right and an increase within the left anterior insula as a function of the presentation rate both of click as well as syllable trains. By contrast, dyslexics exhibited this profile of hemodynamic responses under the nonspeech condition only. As concerns syllables, activation in dyslexics did not depend on presentation rate. Moreover, a subtraction analysis of hemodynamic main effects across conditions and groups revealed decreased activation both of the left and right anterior insula in dyslexics compared to controls during application both of click and syllables. These results indicate, in line with preceding studies, that the insula of both hemispheres is involved in auditory temporal processing of nonlinguistic auditory stimuli and demonstrate, furthermore, that these operations of intrasylvian cortex also extend to the linguistic domain. In addition, our data suggest that the anterior insula represents an important neural correlate of deficient temporal processing of speech and nonspeech sounds in dyslexia. Hum Brain Mapp 2009. © 2008 Wiley‐Liss, Inc.     

Late auditory evoked potentials asymmetry revisited.

OBJECTIVE: To investigate brain asymmetries of the auditory evoked potential (AEP) N100, T-complex, and P200 in response to monaural stimulation. METHODS: Electroencephalographic (EEG) recordings from 68 channels were used to record auditory cortex responses to monaural stimulation from normal hearing participants (N=16). White-noise stimuli and 1000Hz tones were repeatedly presented to either the left or right ear. Source localization of the AEP N100 response was carried out with two symmetric regional sources placed into left and right auditory cortex. Regional source waveform amplitude and latency asymmetries were analyzed for tangential and radial activity explaining the N100, T-complex and P200 AEP components. RESULTS: Regional source waveform analysis showed that early tangential activity in the N100 latency range exhibited larger contralateral amplitudes and shorter latencies for both tone and noise monaural stimuli. Lateralized activity was significantly greater when tones or noise was presented to the left compared to the right ear (p<.001). The ear difference in the degree of lateralization arose due to hemispheric asymmetry. Significantly more tangential activity in the N100 latency range was recorded in the right compared to the left hemisphere in response to stimulation by either tones or noise (p<.001). Neither the radial activity modelling the T-complex, nor activity modelling the P200, showed robust ear or hemisphere differences. CONCLUSIONS: Regional source waveform analysis revealed that the extent of auditory evoked potential asymmetries depends on the ear and hemisphere examined. These findings have implications for future studies utilizing AEP asymmetries to examine normal auditory function or experience-related changes in the auditory cortex. SIGNIFICANCE: The right compared to the left auditory cortex may be more involved in processing monaurally presented tone and noise stimuli.     

Deconvolution of 40 Hz steady-state fields reveals two overlapping source activities of the human auditory cortex.

Steady-state auditory evoked fields were recorded from 15 subjects using a whole head MEG system. Stimuli were 800 ms trains of binaural clicks with constant stimulus onset asynchrony (SOA). Seven different SOA settings (19, 21, 23, 25, 27, 29 and 31 ms) were used to give click rates near 40 Hz. Transient responses to each click were reconstructed using a new algorithm that deconvoluted the averaged responses to the different trains. Spatio-temporal multiple dipole modelling in relation to 3D MRI scans revealed two overlapping source components in both the left and right auditory cortex. The primary sources in the medial part of Heschl's gyrus exhibited a N19-P30-N40 m pattern. The secondary, weaker sources at more lateral sites on Heschl's gyrus showed a N24-P36-N46 m pattern. When applied to transient middle latency auditory evoked fields (MAEFs) recorded at SOAs of 95-135 ms, the primary sources imaged activities similar to the deconvoluted steady-state responses, but the secondary source activities were inconsistent. Linear summation of the deconvoluted source waveforms accounted for more than 96% of the steady-state variance. This indicates that the primary activity of the auditory cortex remains constant up to high stimulation rates and is not specifically enhanced around 40 Hz.     

Specialized neural dynamics for verbal and tonal memory: fMRI evidence in congenital amusia

Behavioral and neuropsychological studies have suggested that tonal and verbal short‐term memory are supported by specialized neural networks. To date however, neuroimaging investigations have failed to confirm this hypothesis. In this study, we investigated the hypothesis of distinct neural resources for tonal and verbal memory by comparing typical nonmusician listeners to individuals with congenital amusia, who exhibit pitch memory impairments with preserved verbal memory. During fMRI, amusics and matched controls performed delayed‐match‐to‐sample tasks with tones and words and perceptual control tasks with the same stimuli. For tonal maintenance, amusics showed decreased activity in the right auditory cortex, inferior frontal gyrus (IFG) and dorso‐lateral‐prefrontal cortex (DLPFC). Moreover, they exhibited reduced right‐lateralized functional connectivity between the auditory cortex and the IFG during tonal encoding and between the IFG and the DLPFC during tonal maintenance. In contrasts, amusics showed no difference compared with the controls for verbal memory, with activation in the left IFG and left fronto‐temporal connectivity. Critically, we observed a group‐by‐material interaction in right fronto‐temporal regions: while amusics recruited these regions less strongly for tonal memory than verbal memory, control participants showed the reversed pattern (tonal > verbal). By benefitting from the rare condition of amusia, our findings suggest specialized cortical systems for tonal and verbal short‐term memory in the human brain.     

N100m in adults possessing absolute pitch

We recorded the auditory evoked magnetic fields from adults with and without absolute pitch under the following conditions: hearing 1000 Hz pure tones inattentively (single tone session) and listening to eight random tones and identifying each tone (labeling session). In the adults with absolute pitch, the bilateral N100m dipole moments increased significantly in the labeling session. While, in the adults without absolute pitch, the left N100m dipole moment alone increased in the labeling session. These results suggest that the adults with absolute pitch execute the labeling task in the bilateral auditory cortices with interhemispheric cooperation, which does not operate in the adults without absolute pitch.     

Human auditory cortex is activated by omissions of auditory stimuli

Cortical signals associated with infrequent tone omissions were recorded from 9 healthy adults with a whole-head 122-channel neuromagnetometer. The stimulus sequence consisted of monaural (left or right) 50-ms 1-kHz tones repeated every 0.2 or 0.5 s, with 7% of the tones randomly omitted. Tones elicited typical responses in the supratemporal auditory cortices. Omissions evoked strong responses over temporal and frontal areas, independently of the side of stimulation, with peak amplitudes at 145–195 ms. Response amplitudes were 60% weaker when the subject was not attending to the stimuli. Omission responses originated in supratemporal auditory cortices bilaterally, indicating that auditory cortex plays an important role in the brain's modelling of temporal characteristics of the auditory environment. Additional activity was observed in the posterolateral frontal cortex and in the superior temporal sulcus, more often in the right than in the left hemisphere.     

Neuromagnetic functional coupling during dichotic listening of speech sounds

The present magnetoencephalography (MEG) study tested the hypothesis of a phase synchronization (functional coupling) of cortical alpha rhythms (about 6-12 Hz) within a "speech" cortical neural network comprising bilateral primary auditory cortex and Wernicke's areas, during dichotic listening (DL) of consonant-vowel (CV) syllables. Dichotic stimulation was done with the CV-syllable pairs /da/-/ba/ (true DL, yielded by stimuli having high spectral overlap) and /da/-/ka/ (sham DL, obtained with stimuli having poor spectral overlap). Whole-head MEG activity (165 sensors) was recorded from 10 healthy right-handed non-musicians showing right ear advantage in a speech DL task. Functional coupling of alpha rhythms was defined as the spectral coherence at the following bands: alpha 1 (about 6-8 Hz), alpha 2 (about 8-10 Hz), and alpha 3 (about 10-12) with respect to the peak of individual alpha frequency. Results showed an inverse pattern of functional coupling: during DL of speech sounds, spectral coherence of the high-band alpha rhythms increased between left auditory and Wernicke's areas with respect to sham DL, whereas it decreased between left and right auditory areas. The increase of functional coupling within the left hemisphere would underlie the processing of the syllable presented to the right ear, which arrives to the left auditory cortex without the interference of the other syllable presented to the left ear. Conversely, the decrease of inter-hemispherical coupling of the high-band alpha might be due to the fact that the two auditory cortices do not receive the same information from the ears during DL. These results suggest that functional coupling of alpha rhythms can constitute a neural substrate for the lateralization of auditory stimuli during DL.     

Psychoacoustic and electrophysiologic correlates of central hearing disorders in man

Summary Evaluation of central hearing disorders in neuropsychologic patients is handicapped by their insufficient ability to describe auditory deficits and by the lack of easily applicable audiological tests. A novel psychoacoustic discrimination test (PDT) was developed to determine ear asymmetries in the discrimination of changes in intensity, frequency, or temporal structure of regularly presented dichotic stimuli. In 19 of 21 patients with lesions of the auditory cortex or the acoustic radiation according to CT scan evaluation a higher error score was observed for target stimuli presented at the ear contralateral to the side of brain infarction (6 right, 15 left). In the remaining 2 and in 3 other patients with lesions sparing auditory structures no significant ear asymmetries were seen. This may indicate that auditory perception is reduced in patients with only one intact auditory cortex or one intact acoustic radiation, possibly because of a limitation in information processing capacity. Auditory evoked potential results are presented for a normal subject and two patients to illustrate electrophysiologic correlates of central hearing disorders. Using a transformation of scalp into dipole source activity (Scherg and von Cramon 1986), a unilateral loss of middle latency activity was found in case A, who had a lesion of the left acoustic radiation. The extended lesion of the right auditory cortex in case B resulted in a loss of both middle and late latency dipole source potentials of the right temporal lobe. In both cases a corresponding increase in the PDT error score on the contralateral ear was found.     

EFFECT OF ATTENTION ON CENTRAL AUDITORY PROCESSING: AN fMRI STUDY

Functional magnetic resonance imaging was used to investigate preattentive and attentional processing of auditory stimuli in 18 right-handed normal volunteers. Responses to trains of 1000-Hz pure tones and infrequent (15%) deviant 1300-Hz tones were characterized while subjects ignored all tones; listened for deviants in the left ear; or listened for deviants in the right ear. Preattentive detection of deviants, associated with the mismatch negativity in electrophysiology, was associated with bilateral temporal lobe activation, with a rightward predominance. Processing of deviant stimuli while attending to either ear produced a more robust and widespread activation of these temporal regions, again with a rightward predominance. Thus, preattentive tone processing appears to be linked to asymmetric activation of a core set of temporal regions in which activity is significantly amplified by selective attention. Extratemporal regions activated by attending to targets in either ear included the anterior cingulate cortex, supramarginal gyrus, and dorsolateral prefrontal cortex.     

Effect of attention on central auditory processing: an fMRI study

Functional magnetic resonance imaging was used to investigate preattentive and attentional processing of auditory stimuli in 18 right-handed normal volunteers. Responses to trains of 1000-Hz pure tones and infrequent (15%) deviant 1300-Hz tones were characterized while subjects ignored all tones; listened for deviants in the left ear; or listened for deviants in the right ear. Preattentive detection of deviants, associated with the mismatch negativity in electrophysiology, was associated with bilateral temporal lobe activation, with a rightward predominance. Processing of deviant stimuli while attending to either ear produced a more robust and widespread activation of these temporal regions, again with a rightward predominance. Thus, preattentive tone processing appears to be linked to asymmetric activation of a core set of temporal regions in which activity is significantly amplified by selective attention. Extratemporal regions activated by attending to targets in either ear included the anterior cingulate cortex, supramarginal gyrus, and dorsolateral prefrontal cortex.     

Incidence of laterality effects in mandibular and manual performance of dichoptic visual pursuit tracking.

In dichotic pursuit auditory tracking tasks (PAT) subjects match a continuously varying pure tone presented to one ear with a second tone presented to the other ear and controlled by unidimensional movements of part of their motor system. In previous studies in which tonal frequency was varied, performance was significantly better when the tone controlled by a speech articulator (tongue, jaw) was presented to the right ear, rather than the left, but not if the tone was hand controlled. This study examined a visual analog of the PAT in which subjects matched the vertical position of a continually moving horizontal line (target) presented on one side of their point of fixation, with a second line (cursor) presented on the other side of their fixation point. Two predictions were confirmed for 12 right handed subjects: that there would be no significant laterality effect for articulatory (jaw) tracking because previous auditory tracking findings were speech related; and, that there would be a significant laterality effect (cursor right field-left hemisphere) for right hand tracking because of the development of a specialized sensorimotor integration mechanism for eye-right hand coordination in the left hemisphere. Alternative explanations for the right hand tracking results, and for the nonsignificant trend towards a laterality effect (cursor left field-right hemisphere) for left hand tracking, were discussed.     

Laterality of basic auditory perception

Laterality (left-right ear differences) of auditory processing was assessed using basic auditory skills: (1) gap detection, (2) frequency discrimination, and (3) intensity discrimination. Stimuli included tones (500, 1000, and 4000 Hz) and wide-band noise presented monaurally to each ear of typical adult listeners. The hypothesis tested was that processing of tonal stimuli would be enhanced by left ear (LE) stimulation and noise by right ear (RE) presentations. To investigate the limits of laterality by (1) spectral width, a narrow-band noise (NBN) of 450-Hz bandwidth was evaluated using intensity discrimination, and (2) stimulus duration, 200, 500, and 1000 ms duration tones were evaluated using frequency discrimination. A left ear advantage (LEA) was demonstrated with tonal stimuli in all experiments, but an expected REA for noise stimuli was not found. The NBN stimulus demonstrated no LEA and was characterised as a noise. No change in laterality was found with changes in stimulus durations. The LEA for tonal stimuli is felt to be due to more direct connections between the left ear and the right auditory cortex, which has been shown to be primary for spectral analysis and tonal processing. The lack of a REA for noise stimuli is unexplained. Sex differences in laterality for noise stimuli were noted but were not statistically significant. This study did establish a subtle but clear pattern of LEA for processing of tonal stimuli.     

Effects of attention on dichotic listening: an 15O-PET study

The present study investigated the effect of attention on brain activation in a dichotic listening situation. Dichotic listening is a technique to study laterality effects in the auditory sensory modality. Two different stimuli were presented simultaneously, one in each ear. Twelve subjects listened to lists of consonant-vowel syllables, or short musical instrument passages, with the task of detecting a "target" syllable or musical instrument by pressing a button. The target stimulus appeared an equal number of times in the left and right ear. The subjects were instructed to either concentrate on the stimuli presented in both ears, or only on the left or right ear stimulus. Brain activation was measured with 15O-PET, and significant changes in regional normalized counts (rNC) were evaluated using statistical parametric mapping (SPM96) software. Concentrating on either the right or left ear stimulus significantly decreased activity bilaterally in the temporal lobes compared to concentrating on both ear stimuli, at the expense of an increased activation in the right posterior and inferior superior parietal lobe. The CV-syllables activated areas corresponding to the classic language areas of Broca and Wernicke. The musical instrument stimuli mainly activated areas in visual association cortex, cerebellum, and the hippocampus. An interpretation of the findings is that attention has a facilitating effect for auditory processing, causing reduced activation in the primary auditory cortex when attention is explicitly recruited. The observed activations in the parietal lobe during the focused attention conditions could be part of a modality non-specific "attentional network".     

MEG auditory evoked fields suggest altered structural/functional asymmetry in primary but not secondary auditory cortex in bipolar disorder

Objectives: Objective physiological indices independently characterizing affective and schizophreniform psychoses would contribute to our understanding of the nature of their relationships. Magnetoencephalography (MEG)‐based metrics of altered structural/functional asymmetry in the superior temporal gyrus have previously been found to characterize schizophrenia at the level of both the primary auditory (AI) and the secondary auditory (AII) cortex. This study examines these markers in patients with bipolar disorder, with the goal of improved understanding of the patterns of brain asymmetry that may independently characterize affective and schizophreniform psychosis. Methods: We studied 17 euthymic bipolar subjects and 17 matched controls. Auditory evoked fields were generated by both 40 Hz auditory stimuli eliciting steady state gamma band (SSR), activating the AI cortex, and discrete 1 kHz tone pips, activating the AII cortex. MEG was recorded from the hemisphere contralateral to the ear stimulated using a 37‐channel MEG system. Source location estimates were calculated in both left and right hemispheres. Neuroanatomical location estimates for medial Heschl’s gyri were determined from magnetic resonance images for correlation with MEG source locations. Results: Bipolar subjects failed to demonstrate normal laterality of SSR AI responses, indicating altered patterns of asymmetry at the level of AI cortex, but demonstrated normal asymmetry of AII responses (right anterior to left). Medial Heschl’s gyri centroids were similarly lateralized in both groups, however (right anterior to left), dissociating function from structure in the AI cortex in the bipolar group. Conclusions: The findings are compatible with altered functional/structural relationships, including diminished left‐right hemisphere asymmetry of the AI, but not the AII cortex in bipolar disorder. In schizophrenia, both the AI and AII cortices exhibit such derangements; thus, the findings support both shared and nonshared features of auditory cortical disruption between the two disorders. This functional disorganization may help explain previously reported decreases in amplitude and phase synchrony of SSR gamma band responses in bipolar subjects, suggesting impaired neocortical synchrony in AI, possibly at a cortico‐thalamic level, but perhaps not extending to heteromodal association cortex, and may relate to the cognitive impairments found in bipolar disorder.