A multidimensional characterization of the neurocognitive architecture underlying age-related temporal speech processing.

Healthy aging is often associated with speech comprehension difficulties in everyday life situations despite a pure-tone hearing threshold in the normative range. Drawing on this background, we used a multidimensional approach to assess the functional and structural neural correlates underlying age-related temporal speech processing while controlling for pure-tone hearing acuity. Accordingly, we combined structural magnetic resonance imaging and electroencephalography, and collected behavioral data while younger and older adults completed a phonetic categorization and discrimination task with consonant-vowel syllables varying along a voice-onset time continuum. The behavioral results confirmed age-related temporal speech processing singularities which were reflected in a shift of the boundary of the psychometric categorization function, with older adults perceiving more syllable characterized by a short voice-onset time as /ta/ compared to younger adults. Furthermore, despite the absence of any between-group differences in phonetic discrimination abilities, older adults demonstrated longer N100/P200 latencies as well as increased P200 amplitudes while processing the consonant-vowel syllables varying in voice-onset time. Finally, older adults also exhibited a divergent anatomical gray matter infrastructure in bilateral auditory-related and frontal brain regions, as manifested in reduced cortical thickness and surface area. Notably, in the younger adults but not in the older adult cohort, cortical surface area in these two gross anatomical clusters correlated with the categorization of consonant-vowel syllables characterized by a short voice-onset time, suggesting the existence of a critical gray matter threshold that is crucial for consistent mapping of phonetic categories varying along the temporal dimension. Taken together, our results highlight the multifaceted dimensions of age-related temporal speech processing characteristics, and pave the way toward a better understanding of the relationships between hearing, speech and the brain in older age.

Early tone categorization in absolute pitch musicians is subserved by the right-sided perisylvian brain.

Absolute pitch (AP) is defined as the ability to identify and label tones without reference to keyality. In this context, the main question is whether early or late processing stages are responsible for this ability. We investigated the electrophysiological responses to tones in AP and relative pitch (RP) possessors while participants listened attentively to sine tones. Since event-related potentials are particularly suited for tracking tone encoding (N100 and P200), categorization (N200), and mnemonic functions (N400), we hypothesized that differences in early pitch processing stages would be reflected by increased N100 and P200-related areas in AP musicians. Otherwise, differences in later cognitive stages of tone processing should be mirrored by increased N200 and/or N400 areas in AP musicians. AP possessors exhibited larger N100 areas and a tendency towards enhanced P200 areas. Furthermore, the sources of these components were estimated and statistically compared between the two groups for a set of a priori defined regions of interest. AP musicians demonstrated increased N100-related current densities in the right superior temporal sulcus, middle temporal gyrus, and Heschl's gyrus. Results are interpreted as indicating that early between-group differences in right-sided perisylvian brain regions might reflect auditory tone categorization rather than labelling mechanisms.

Theta Coherence Asymmetry in the Dorsal Stream of Musicians Facilitates Word Learning.

Word learning constitutes a human faculty which is dependent upon two anatomically distinct processing streams projecting from posterior superior temporal (pST) and inferior parietal (IP) brain regions toward the prefrontal cortex (dorsal stream) and the temporal pole (ventral stream). The ventral stream is involved in mapping sensory and phonological information onto lexical-semantic representations, whereas the dorsal stream contributes to sound-to-motor mapping, articulation, complex sequencing in the verbal domain, and to how verbal information is encoded, stored, and rehearsed from memory. In the present source-based EEG study, we evaluated functional connectivity between the IP lobe and Broca's area while musicians and non-musicians learned pseudowords presented in the form of concatenated auditory streams. Behavioral results demonstrated that musicians outperformed non-musicians, as reflected by a higher sensitivity index (d'). This behavioral superiority was paralleled by increased left-hemispheric theta coherence in the dorsal stream, whereas non-musicians showed stronger functional connectivity in the right hemisphere. Since no between-group differences were observed in a passive listening control condition nor during rest, results point to a task-specific intertwining between musical expertise, functional connectivity, and word learning.

Electrophysiological Correlates of Absolute Pitch in a Passive Auditory Oddball Paradigm: a Direct Replication Attempt.

Humans with absolute pitch (AP) are able to effortlessly name the pitch class of a sound without an external reference. The association of labels with pitches cannot be entirely suppressed even if it interferes with task demands. This suggests a high level of automaticity of pitch labeling in AP. The automatic nature of AP was further investigated in a study by Rogenmoser et al. (2015). Using a passive auditory oddball paradigm in combination with electroencephalography, they observed electrophysiological differences between musicians with and without AP in response to piano tones. Specifically, the AP musicians showed a smaller P3a, an event-related potential (ERP) component presumably reflecting early attentional processes. In contrast, they did not find group differences in the mismatch negativity (MMN), an ERP component associated with auditory memory processes. They concluded that early cognitive processes are facilitated in AP during passive listening and are more important for AP than the preceding sensory processes. In our direct replication study on a larger sample of musicians with (n = 54, 27 females, 27 males) and without (n = 50, 24 females, 26 males) AP, we successfully replicated the non-significant effects of AP on the MMN. However, we could not replicate the significant effects for the P3a. Additional Bayes factor analyses revealed moderate to strong evidence (Bayes factor > 3) for the null hypothesis for both MMN and P3a. Therefore, the results of this replication study do not support the postulated importance of cognitive facilitation in AP during passive tone listening.

Human Brainstem Exhibits higher Sensitivity and Specificity than Auditory-Related Cortex to Short-Term Phonetic Discrimination Learning.

Phonetic discrimination learning is an active perceptual process that operates under the influence of cognitive control mechanisms by increasing the sensitivity of the auditory system to the trained stimulus attributes. It is assumed that the auditory cortex and the brainstem interact in order to refine how sounds are transcribed into neural codes. Here, we evaluated whether these two computational entities are prone to short-term functional changes, whether there is a chronological difference in malleability, and whether short-term training suffices to alter reciprocal interactions. We performed repeated cortical (i.e., mismatch negativity responses, MMN) and subcortical (i.e., frequency-following response, FFR) EEG measurements in two groups of participants who underwent one hour of phonetic discrimination training or were passively exposed to the same stimulus material. The training group showed a distinctive brainstem energy reduction in the trained frequency-range (i.e., first formant), whereas the passive group did not show any response modulation. Notably, brainstem signal change correlated with the behavioral improvement during training, this result indicating a close relationship between behavior and underlying brainstem physiology. Since we did not reveal group differences in MMN responses, results point to specific short-term brainstem changes that precede functional alterations in the auditory cortex.

Faster native vowel discrimination learning in musicians is mediated by an optimization of mnemonic functions.

The ability to discriminate phonemes varying in spectral and temporal attributes constitutes one of the most basic intrinsic elements underlying language learning mechanisms. Since previous work has consistently shown that professional musicians are characterized by perceptual and cognitive advantages in a variety of language-related tasks, and since vowels can be considered musical sounds within the domain of speech, here we investigated the behavioral and electrophysiological correlates of native vowel discrimination learning in a sample of professional musicians and non-musicians. We evaluated the contribution of both the neurophysiological underpinnings of perceptual (i.e., N1/P2 complex) and mnemonic functions (i.e., N400 and P600 responses) while the participants were instructed to judge whether pairs of native consonant-vowel (CV) syllables manipulated in the first formant transition of the vowel (i.e., from /tu/ to /to/) were identical or not. Results clearly demonstrated faster learning in musicians, compared to non-musicians, as reflected by shorter reaction times and higher accuracy. Most notably, in terms of morphology, time course, and voltage strength, this steeper learning curve was accompanied by distinctive N400 and P600 manifestations between the two groups. In contrast, we did not reveal any group differences during the early stages of auditory processing (i.e., N1/P2 complex), suggesting that faster learning was mediated by an optimization of mnemonic but not perceptual functions. Based on a clear taxonomy of the mnemonic functions involved in the task, results are interpreted as pointing to a relationship between faster learning mechanisms in musicians and an optimization of echoic (i.e., N400 component) and working memory (i.e., P600 component) functions.

Independent component processes underlying emotions during natural music listening.

The aim of this study was to investigate the brain processes underlying emotions during natural music listening. To address this, we recorded high-density electroencephalography (EEG) from 22 subjects while presenting a set of individually matched whole musical excerpts varying in valence and arousal. Independent component analysis was applied to decompose the EEG data into functionally distinct brain processes. A k-means cluster analysis calculated on the basis of a combination of spatial (scalp topography and dipole location mapped onto the Montreal Neurological Institute brain template) and functional (spectra) characteristics revealed 10 clusters referring to brain areas typically involved in music and emotion processing, namely in the proximity of thalamic-limbic and orbitofrontal regions as well as at frontal, fronto-parietal, parietal, parieto-occipital, temporo-occipital and occipital areas. This analysis revealed that arousal was associated with a suppression of power in the alpha frequency range. On the other hand, valence was associated with an increase in theta frequency power in response to excerpts inducing happiness compared to sadness. These findings are partly compatible with the model proposed by Heller, arguing that the frontal lobe is involved in modulating valenced experiences (the left frontal hemisphere for positive emotions) whereas the right parieto-temporal region contributes to the emotional arousal.

The Influence of Pre-stimulus EEG Activity on Reaction Time During a Verbal Sternberg Task is Related to Musical Expertise.

Previous work highlighted the possibility that musical training has an influence on cognitive functioning. The suggested reason for this influence is the strong recruitment of attention, planning, and working memory functions during playing a musical instrument. The purpose of the present work was twofold, namely to evaluate the general relationship between pre-stimulus electrophysiological activity and cognition, and more specifically the influence of musical expertise on working memory functions. With this purpose in mind, we used covariance mapping analyses to evaluate whether pre-stimulus electroencephalographic activity is predictive for reaction time during a visual working memory task (Sternberg paradigm) in musicians and non-musicians. In line with our hypothesis, we replicated previous findings pointing to a general predictive value of pre-stimulus activity for working memory performance. Most importantly, we also provide first evidence for an influence of musical expertise on working memory performance that could distinctively be predicted by pre-stimulus spectral power. Our results open novel perspectives for better comprehending the vast influences of musical expertise on cognition.

Interhemispheric transcallosal connectivity between the left and right planum temporale predicts musicianship, performance in temporal speech processing, and functional specialization.

Currently, there is strong evidence showing that musicianship favours functional and structural changes of the left planum temporale (PT), and that these cortical reorganizations facilitate the discrimination of temporal speech cues. Based on the proposition of a division of labour between the left and right PT, here we postulated that the musicians' advantage in processing temporal speech cues and PT specialization origin, at least in part, from increased white matter connectivity between the two auditory-related cortices. In particular, we assume that increased transcallosal PT connectivity might promote functional specialization and asymmetry of homotopic brain regions. With this purpose in mind, we applied diffusion tensor imaging and compared axial diffusivity (AD), radial diffusivity (RD), and fractional anisotropy (FA) of the interhemispheric connection between the left and right PT in thirteen musicians and 13 nonmusicians. Furthermore, in the form of an addendum, we integrated cortical surface area values and blood oxygenation level dependent (BOLD) responses of the left PT that were collected in the context of two previous studies conducted with the same sample of subjects. Our results indicate increased connectivity between the left and right PT in musicians compared to nonmusicians, as indexed by reduced mean RD. We did not find significant between-group differences in FA and AD. Most notably, RD was related to the performance in the phonetic categorization task, musical aptitudes, as well as to BOLD responses in the left PT. Hence, we provide first evidence for a relationship between PT connectivity, musicianship, and phonetic categorization.

Bridging the gap between perceptual and cognitive perspectives on absolute pitch.

Absolute pitch (AP) refers to the rare ability to identify the chroma of a tone or to produce a specific pitch without reference to keyality (e.g., G or C). Previously, AP has been proposed to rely on the distinctive functional-anatomical architecture of the left auditory-related cortex (ARC), this specific trait possibly enabling an optimized early "categorical perception". In contrast, currently prevailing models of AP postulate that cognitive rather than perceptual processes, namely "pitch labeling" mechanisms, more likely constitute the bearing skeleton of AP. This associative memory component has previously been proposed to be dependent, among other mechanisms, on the recruitment of the left dorsolateral prefrontal cortex (DLPFC) as well as on the integrity of the left arcuate fasciculus, a fiber bundle linking the posterior supratemporal plane with the DLPFC. Here, we attempted to integrate these two apparently conflicting perspectives on AP, namely early "categorical perception" and "pitch labeling". We used electroencephalography and evaluated resting-state intracranial functional connectivity between the left ARC and DLPFC in a sample of musicians with and without AP. Results demonstrate significantly increased left-hemispheric theta phase synchronization in AP compared with non-AP musicians. Within the AP group, this specific electrophysiological marker was predictive of absolute-hearing behavior and explained ∼30% of variance. Thus, we propose that in AP subjects the tonal inputs and the corresponding mnemonic representations are tightly coupled in such a manner that the distinctive electrophysiological signature of AP can saliently be detected in only 3 min of resting-state measurements.

Auditory evoked responses in musicians during passive vowel listening are modulated by functional connectivity between bilateral auditory-related brain regions.

Currently, there is striking evidence showing that professional musical training can substantially alter the response properties of auditory-related cortical fields. Such plastic changes have previously been shown not only to abet the processing of musical sounds, but likewise spectral and temporal aspects of speech. Therefore, here we used the EEG technique and measured a sample of musicians and nonmusicians while the participants were passively exposed to artificial vowels in the context of an oddball paradigm. Thereby, we evaluated whether increased intracerebral functional connectivity between bilateral auditory-related brain regions may promote sensory specialization in musicians, as reflected by altered cortical N1 and P2 responses. This assumption builds on the reasoning that sensory specialization is dependent, at least in part, on the amount of synchronization between the two auditory-related cortices. Results clearly revealed that auditory-evoked N1 responses were shaped by musical expertise. In addition, in line with our reasoning musicians showed an overall increased intracerebral functional connectivity (as indexed by lagged phase synchronization) in theta, alpha, and beta bands. Finally, within-group correlative analyses indicated a relationship between intracerebral beta band connectivity and cortical N1 responses, however only within the musicians' group. Taken together, we provide first electrophysiological evidence for a relationship between musical expertise, auditory-evoked brain responses, and intracerebral functional connectivity among auditory-related brain regions.

Music and language expertise influence the categorization of speech and musical sounds: behavioral and electrophysiological measurements.

In this study, we used high-density EEG to evaluate whether speech and music expertise has an influence on the categorization of expertise-related and unrelated sounds. With this purpose in mind, we compared the categorization of speech, music, and neutral sounds between professional musicians, simultaneous interpreters (SIs), and controls in response to morphed speech-noise, music-noise, and speech-music continua. Our hypothesis was that music and language expertise will strengthen the memory representations of prototypical sounds, which act as a perceptual magnet for morphed variants. This means that the prototype would "attract" variants. This so-called magnet effect should be manifested by an increased assignment of morphed items to the trained category, by a reduced maximal slope of the psychometric function, as well as by differential event-related brain responses reflecting memory comparison processes (i.e., N400 and P600 responses). As a main result, we provide first evidence for a domain-specific behavioral bias of musicians and SIs toward the trained categories, namely music and speech. In addition, SIs showed a bias toward musical items, indicating that interpreting training has a generic influence on the cognitive representation of spectrotemporal signals with similar acoustic properties to speech sounds. Notably, EEG measurements revealed clear distinct N400 and P600 responses to both prototypical and ambiguous items between the three groups at anterior, central, and posterior scalp sites. These differential N400 and P600 responses represent synchronous activity occurring across widely distributed brain networks, and indicate a dynamical recruitment of memory processes that vary as a function of training and expertise.

An empirical reevaluation of absolute pitch: behavioral and electrophysiological measurements.

Here, we reevaluated the "two-component" model of absolute pitch (AP) by combining behavioral and electrophysiological measurements. This specific model postulates that AP is driven by a perceptual encoding ability (i.e., pitch memory) plus an associative memory component (i.e., pitch labeling). To test these predictions, during EEG measurements AP and non-AP (NAP) musicians were passively exposed to piano tones (first component of the model) and additionally instructed to judge whether combinations of tones and labels were conceptually associated or not (second component of the model). Auditory-evoked N1/P2 potentials did not reveal differences between the two groups, thus indicating that AP is not necessarily driven by a differential pitch encoding ability at the processing level of the auditory cortex. Otherwise, AP musicians performed the conceptual association task with an order of magnitude better accuracy and shorter RTs than NAP musicians did, this result clearly pointing to distinctive conceptual associations in AP possessors. Most notably, this behavioral superiority was reflected by an increased N400 effect and accompanied by a subsequent late positive component, the latter not being distinguishable in NAP musicians.

The encoding of vowels and temporal speech cues in the auditory cortex of professional musicians: an EEG study.

Here, we applied a multi-feature mismatch negativity (MMN) paradigm in order to systematically investigate the neuronal representation of vowels and temporally manipulated CV syllables in a homogeneous sample of string players and non-musicians. Based on previous work indicating an increased sensitivity of the musicians' auditory system, we expected to find that musically trained subjects will elicit increased MMN amplitudes in response to temporal variations in CV syllables, namely voice-onset time (VOT) and duration. In addition, since different vowels are principally distinguished by means of frequency information and musicians are superior in extracting tonal (and thus frequency) information from an acoustic stream, we also expected to provide evidence for an increased auditory representation of vowels in the experts. In line with our hypothesis, we could show that musicians are not only advantaged in the pre-attentive encoding of temporal speech cues, but most notably also in processing vowels. Additional "just noticeable difference" measurements suggested that the musicians' perceptual advantage in encoding speech sounds was more likely driven by the generic constitutional properties of a highly trained auditory system, rather than by its specialisation for speech representations per se. These results shed light on the origin of the often reported advantage of musicians in processing a variety of speech sounds.

Increased cortical surface area of the left planum temporale in musicians facilitates the categorization of phonetic and temporal speech sounds.

We measured musicians and non-musicians by using structural magnetic resonance imaging to investigate relationships between cortical features of the left planum temporale (PT) and the categorization of consonant-vowel (CV) syllables and their reduced-spectrum analogues. The present work is based on previous functional studies consistently showing that the left PT is particularly responsive to transient acoustic features in CV syllables and their reduced-spectrum analogues, and on striking evidence pointing to structural alterations of the left PT as a function of musicianship. By combining these two observations, we hypothesized to find that differences in cortical surface area (SA) and cortical thickness (CT) of the left PT in musicians may facilitate the categorization of fast-changing phonetic cues. Behavioural results indicated that musicians and non-musicians achieved a comparable performance in the categorization of CV syllables, whereas the musicians performed significantly better than the controls in the more demanding reduced-spectrum condition. This better behavioural performance corresponds to an increased cortical SA of the left PT in musicians compared to non-musicians. No differences in CT of the left PT were found between groups. In line with our predictions, we revealed a positive correlation between cortical SA of the left PT in musicians and the behavioural performance during the acoustically more demanding reduced-spectrum condition. Hence, we provide first evidence for a relationship between musical expertise, cortical SA of the left PT, and the processing of fast-changing phonetic cues.

Musicianship boosts perceptual learning of pseudoword-chimeras: an electrophysiological approach.

A vast amount of previous work has consistently revealed that professional music training is associated with functional and structural alterations of auditory-related brain regions. Meanwhile, there is also an increasing array of evidence, which shows that musicianship facilitates segmental, as well as supra-segmental aspects of speech processing. Based on this evidence, we addressed a novel research question, namely whether professional music training has an influence on the perceptual learning of speech sounds. In the context of an EEG experiment, we presented auditory pseudoword-chimeras, manipulated in terms of spectral- or envelope-related acoustic information, to a group of professional musicians and non-musicians. During EEG measurements, participants were requested to assign the auditory-presented pseudoword-chimeras to one out of four visually presented templates. As expected, both groups showed behavioural learning effects during the time course of the experiment. These learning effects were associated with an increase in accuracy, a decrease in reaction time, as well as a decrease in the P2-like microstate duration in both groups. Notably, the musicians showed an increased learning performance compared to the controls during the first two runs of the spectral condition. This perceptual learning effect, which varies as a function of musical expertise, was reflected by a reduction of the P2-like microstate duration. Results may mirror transfer effects from musical training to the processing of spectral information in speech sounds. Hence, this study provides first evidence for a relationship between changes in microstates, musical expertise, and perceptual verbal learning mechanisms.

Pre-attentive modulation of brain responses to tones in coloured-hearing synesthetes.

BACKGROUND: Coloured-hearing (CH) synesthesia is a perceptual phenomenon in which an acoustic stimulus (the inducer) initiates a concurrent colour perception (the concurrent). Individuals with CH synesthesia "see" colours when hearing tones, words, or music; this specific phenomenon suggesting a close relationship between auditory and visual representations. To date, it is still unknown whether the perception of colours is associated with a modulation of brain functions in the inducing brain area, namely in the auditory-related cortex and associated brain areas. In addition, there is an on-going debate as to whether attention to the inducer is necessarily required for eliciting a visual concurrent, or whether the latter can emerge in a pre-attentive fashion. RESULTS: By using the EEG technique in the context of a pre-attentive mismatch negativity (MMN) paradigm, we show that the binding of tones and colours in CH synesthetes is associated with increased MMN amplitudes in response to deviant tones supposed to induce novel concurrent colour perceptions. Most notably, the increased MMN amplitudes we revealed in the CH synesthetes were associated with stronger intracerebral current densities originating from the auditory cortex, parietal cortex, and ventral visual areas. CONCLUSIONS: The automatic binding of tones and colours in CH synesthetes is accompanied by an early pre-attentive process recruiting the auditory cortex, inferior and superior parietal lobules, as well as ventral occipital areas.

Neurofunctional and behavioral correlates of phonetic and temporal categorization in musically trained and untrained subjects.

The perception of rapidly changing verbal and nonverbal auditory patterns is a fundamental prerequisite for speech and music processing. Previously, the left planum temporale (PT) has been consistently shown to support the discrimination of fast changing verbal and nonverbal sounds. Furthermore, it has been repeatedly shown that the functional and structural architecture of this supratemporal brain region differs as a function of musical training. In the present study, we used the functional magnetic resonance imaging technique, in a sample of professional musicians and nonmusicians, in order to examine the functional contribution of the left PT to the categorization of consonant-vowel syllables and their reduced-spectrum analogues. In line with our hypothesis, the musicians showed enhanced brain responses in the left PT and superior discrimination abilities in the reduced-spectrum condition. Moreover, we found a positive correlation between the responsiveness of the left PT and the performance in the reduced-spectrum condition across all subjects irrespective of musical expertise. These results have implications for our understanding of musical expertise in relation to segmental speech processing.

The spatiotemporal characteristics of elementary audiovisual speech and music processing in musically untrained subjects.

Previously, the EEG technique has been used to investigate the spatiotemporal properties of audiovisual (AV) processing by taking advantage of the violation of the "additive model", which is considered to be a very conservative approach. In the present work, we used a less conservative and novel approach than the criterion of superadditivity for estimating AV interactions. Hence, we estimated AV interaction patterns by comparing the responses to AV stimuli with the averaged responses to the unimodal visual and auditory stimuli in musically untrained subjects and by presenting syllables and piano tones coupled with flashlights. Our results suggest that the two AV objects elicited consistent interaction patterns within the time course of unisensory processing in the time range between 80 and 250ms post stimulus onset. The scalp topographies, as well as the source estimation approach we adopted, indicate that the first interaction pattern at around 100ms was partially driven by auditory-related cortical regions. Additionally, we found evidence for a second interaction pattern at around 200ms that was mainly associated with the responsiveness of extra-sensory brain regions. During this later processing stage, only the music condition was associated with putative responses that originated from auditory-related cortical fields. This study provides a novel approach to investigate the basic principles underlying elementary AV speech and music processing in subjects without formal musical education.

Long-term exposure to music enhances the sensitivity of the auditory system in children.

This event-related brain potential study aims to contribute to the present debate regarding the effect of musical training on the maturation of the human auditory nervous system. To address this issue, we recorded the mismatch negativity (MMN) evoked by violin and pure sine-wave tones in a group of 7.5- to 12-year-old children who had either several years of musical experience with Suzuki violin lessons, or no musical training. The strength of the MMN responses to violin tones evident in the Suzuki students clearly surpassed responses in controls; the reverse pattern was observed for sine-wave tones. Suzuki students showed significantly shorter MMN latencies to violin tones than to pure tones; the MMN latency did not differ significantly between pure tones and violin sounds in the control group. Thus, our data provide general evidence of how and to what extent extensive musical experience affects the maturation of human auditory function at multiple levels, namely, accuracy and speed of auditory discrimination processing. Our findings add to the present understanding of neuroplastic organization and function of the mammalian nervous system. Furthermore, behavioural recordings obtained from the participating children provide corroborating evidence for a relationship between the duration and intensity of training, the specific sensitivity to instrumental timbre, and pitch recognition abilities.