Tracking Training-Related Plasticity by Combining fMRI and DTI: The Right Hemisphere Ventral Stream Mediates Musical Syntax Processing.

As a functional homolog for left-hemispheric syntax processing in language, neuroimaging studies evidenced involvement of right prefrontal regions in musical syntax processing, of which underlying white matter connectivity remains unexplored so far. In the current experiment, we investigated the underlying pathway architecture in subjects with 3 levels of musical expertise. Employing diffusion tensor imaging tractography, departing from seeds from our previous functional magnetic resonance imaging study on music syntax processing in the same participants, we identified a pathway in the right ventral stream that connects the middle temporal lobe with the inferior frontal cortex via the extreme capsule, and corresponds to the left hemisphere ventral stream, classically attributed to syntax processing in language comprehension. Additional morphometric consistency analyses allowed dissociating tract core from more dispersed fiber portions. Musical expertise related to higher tract consistency of the right ventral stream pathway. Specifically, tract consistency in this pathway predicted the sensitivity for musical syntax violations. We conclude that enduring musical practice sculpts ventral stream architecture. Our results suggest that training-related pathway plasticity facilitates the right hemisphere ventral stream information transfer, supporting an improved sound-to-meaning mapping in music.

Electrical Neuroimaging of Music Processing Reveals Mid-Latency Changes with Level of Musical Expertise.

This original research focused on the effect of musical training intensity on cerebral and behavioral processing of complex music using high-density event-related potential (ERP) approaches. Recently we have been able to show progressive changes with training in gray and white matter, and higher order brain functioning using (f)MRI [(functional) Magnetic Resonance Imaging], as well as changes in musical and general cognitive functioning. The current study investigated the same population of non-musicians, amateur pianists and expert pianists using spatio-temporal ERP analysis, by means of microstate analysis, and ERP source imaging. The stimuli consisted of complex musical compositions containing three levels of transgression of musical syntax at closure that participants appraised. ERP waveforms, microstates and underlying brain sources revealed gradual differences according to musical expertise in a 300-500 ms window after the onset of the terminal chords of the pieces. Within this time-window, processing seemed to concern context-based memory updating, indicated by a P3b-like component or microstate for which underlying sources were localized in the right middle temporal gyrus, anterior cingulate and right parahippocampal areas. Given that the 3 expertise groups were carefully matched for demographic factors, these results provide evidence of the progressive impact of training on brain and behavior.

Impact of major and minor mode on EEG frequency range activities of music processing as a function of expertise.

Processing western tonal music may yield distinct brain responses depending on the mode of the musical compositions. Although subjective feelings in response to major and minor mode are well described, the underlying brain mechanisms and their development with increasing expertise have not been thoroughly examined. Using high-density electroencephalography, the present study investigated neuronal activities in the frequency domain in response to polyphone musical compositions in major and minor mode in non-musicians, amateurs and experts. During active listening decrease of theta- and gamma-frequency range activities occurred with increasing expertise in right posterior regions, possibly reflecting enhanced processing efficiency. Moreover, minor and major compositions distinctively modulated synchronization of neuronal activities in high frequency ranges (beta and gamma) in frontal regions, with increased activity in response to minor compositions in musicians and in experts in particular. These results suggest that high-frequency electroencephalographic (EEG) activities carry information about musical mode, showing gradual increase of processing efficiency and sensitivity with musical expertise.

Transcranial Alternating Current Stimulation (tACS) differentially modulates speech perception in young and older adults.

BACKGROUND: Normal aging is accompanied by a functional decline in processing temporal features of spoken language, such as voice onset time (VOT). On an electrophysiological level, this finding is paralleled by altered patterns of gamma oscillations. OBJECTIVE: Using 40 Hz transcranial alternating current stimulation (tACS) over the bilateral auditory cortex, this study aims to compare the effect of tACS to modulate VOT-processing in samples of healthy young and older adults. METHODS: Twenty-five healthy young (age 20-35 years) and 20 older adults (age 60-75 years) participated in this study. Presented with an auditory phoneme categorization task participants received 40 Hz and 6 Hz tACS on two consecutive sessions. RESULTS: While 40 Hz tACS diminished task accuracy in young adults, older adults benefitted from this stimulation resulting in a more precise phonetic categorization. CONCLUSION: The results of the study are discussed with respect to the non-linear relationship between gamma oscillations in the vicinity of the auditory cortex and VOT-processing. The present findings are promising in the context of an intervention for subjects with impaired ability to process temporal acoustic features in the speech signal.

40Hz-Transcranial alternating current stimulation (tACS) selectively modulates speech perception.

The present study investigated the functional relevance of gamma oscillations for the processing of rapidly changing acoustic features in speech signals. For this purpose we analyzed repetition-induced perceptual learning effects in 18 healthy adult participants. The participants received either 6Hz or 40Hz tACS over the bilateral auditory cortex, while repeatedly performing a phoneme categorization task. In result, we found that 40Hz tACS led to a specific alteration in repetition-induced perceptual learning. While participants in the non-stimulated control group as well as those in the experimental group receiving 6Hz tACS considerably improved their perceptual performance, the application of 40Hz tACS selectively attenuated the repetition-induced improvement in phoneme categorization abilities. Our data provide causal evidence for a functional relevance of gamma oscillations during the perceptual learning of acoustic speech features. Moreover, we demonstrate that even less than twenty minutes of alternating current stimulation below the individual perceptual threshold is sufficient to affect speech perception. This finding is relevant in that this novel approach might have implications with respect to impaired speech processing in dyslexics and older adults.

Age-Related Neural Oscillation Patterns During the Processing of Temporally Manipulated Speech.

This EEG-study aims to investigate age-related differences in the neural oscillation patterns during the processing of temporally modulated speech. Viewing from a lifespan perspective, we recorded the electroencephalogram (EEG) data of three age samples: young adults, middle-aged adults and older adults. Stimuli consisted of temporally degraded sentences in Swedish-a language unfamiliar to all participants. We found age-related differences in phonetic pattern matching when participants were presented with envelope-degraded sentences, whereas no such age-effect was observed in the processing of fine-structure-degraded sentences. Irrespective of age, during speech processing the EEG data revealed a relationship between envelope information and the theta band (4-8 Hz) activity. Additionally, an association between fine-structure information and the gamma band (30-48 Hz) activity was found. No interaction, however, was found between acoustic manipulation of stimuli and age. Importantly, our main finding was paralleled by an overall enhanced power in older adults in high frequencies (gamma: 30-48 Hz). This occurred irrespective of condition. For the most part, this result is in line with the Asymmetric Sampling in Time framework (Poeppel in Speech Commun 41:245-255, 2003), which assumes an isomorphic correspondence between frequency modulations in neurophysiological patterns and acoustic oscillations in spoken language. We conclude that speech-specific neural networks show strong stability over adulthood, despite initial processes of cortical degeneration indicated by enhanced gamma power. The results of our study therefore confirm the concept that sensory and cognitive processes undergo multidirectional trajectories within the context of healthy aging.

Electrophysiological evidence for a specific neural correlate of musical violation expectation in primary-school children.

The majority of studies on music processing in children used simple musical stimuli. Here, primary schoolchildren judged the appropriateness of musical closure in expressive polyphone music, while high-density electroencephalography was recorded. Stimuli ended either regularly or contained refined in-key harmonic transgressions at closure. The children discriminated the transgressions well above chance. Regular and transgressed endings evoked opposite scalp voltage configurations peaking around 400ms after stimulus onset with bilateral frontal negativity for regular and centro-posterior negativity (CPN) for transgressed endings. A positive correlation could be established between strength of the CPN response and rater sensitivity (d-prime). We also investigated whether the capacity to discriminate the transgressions was supported by auditory domain specific or general cognitive mechanisms, and found that working memory capacity predicted transgression discrimination. Latency and distribution of the CPN are reminiscent of the N400, typically observed in response to semantic incongruities in language. Therefore our observation is intriguing, as the CPN occurred here within an intra-musical context, without any symbols referring to the external world. Moreover, the harmonic in-key transgressions that we implemented may be considered syntactical as they transgress structural rules. Such structural incongruities in music are typically followed by an early right anterior negativity (ERAN) and an N5, but not so here. Putative contributive sources of the CPN were localized in left pre-motor, mid-posterior cingulate and superior parietal regions of the brain that can be linked to integration processing. These results suggest that, at least in children, processing of syntax and meaning may coincide in complex intra-musical contexts.

Musical training intensity yields opposite effects on grey matter density in cognitive versus sensorimotor networks.

Using optimized voxel-based morphometry, we performed grey matter density analyses on 59 age-, sex- and intelligence-matched young adults with three distinct, progressive levels of musical training intensity or expertise. Structural brain adaptations in musicians have been repeatedly demonstrated in areas involved in auditory perception and motor skills. However, musical activities are not confined to auditory perception and motor performance, but are entangled with higher-order cognitive processes. In consequence, neuronal systems involved in such higher-order processing may also be shaped by experience-driven plasticity. We modelled expertise as a three-level regressor to study possible linear relationships of expertise with grey matter density. The key finding of this study resides in a functional dissimilarity between areas exhibiting increase versus decrease of grey matter as a function of musical expertise. Grey matter density increased with expertise in areas known for their involvement in higher-order cognitive processing: right fusiform gyrus (visual pattern recognition), right mid orbital gyrus (tonal sensitivity), left inferior frontal gyrus (syntactic processing, executive function, working memory), left intraparietal sulcus (visuo-motor coordination) and bilateral posterior cerebellar Crus II (executive function, working memory) and in auditory processing: left Heschl's gyrus. Conversely, grey matter density decreased with expertise in bilateral perirolandic and striatal areas that are related to sensorimotor function, possibly reflecting high automation of motor skills. Moreover, a multiple regression analysis evidenced that grey matter density in the right mid orbital area and the inferior frontal gyrus predicted accuracy in detecting fine-grained incongruities in tonal music.

Effects of prior information on decoding degraded speech: an fMRI study.

Expectations and prior knowledge are thought to support the perceptual analysis of incoming sensory stimuli, as proposed by the predictive-coding framework. The current fMRI study investigated the effect of prior information on brain activity during the decoding of degraded speech stimuli. When prior information enabled the comprehension of the degraded sentences, the left middle temporal gyrus and the left angular gyrus were activated, highlighting a role of these areas in meaning extraction. In contrast, the activation of the left inferior frontal gyrus (area 44/45) appeared to reflect the search for meaningful information in degraded speech material that could not be decoded because of mismatches with the prior information. Our results show that degraded sentences evoke instantaneously different percepts and activation patterns depending on the type of prior information, in line with prediction-based accounts of perception.

Hippocampal volume predicts fluid intelligence in musically trained people.

Recently, age-related hippocampal (HP) volume loss could be associated with a decrease in general fluid intelligence (gF). In the present study we investigated whether and how extensive musical training modulates human HP volume and gF performance. Previously, some studies demonstrated positive effects of musical training on higher cognitive functions such as learning and memory, associated with neural adaptations beyond the auditory domain. In order to detect possible associations between musical training and gF, we bilaterally segmented the HP formation and assessed the individual gF performance of people with different levels of musical expertise. Multiple regression analyses revealed that HP volume predicts gF in musicians but not in nonmusicians; in particular, bilaterally enhanced HP volume is associated with increased gF exclusively in musically trained people (amateurs and experts). This result suggests that musical training facilitates the recruitment of cognitive resources, which are essential for gF and linked to HP functioning. Musical training, even at a moderate level of intensity, can thus be considered as a potential strategy to decelerate age-related effects of cognitive decline.

Degree of musical expertise modulates higher order brain functioning.

Using functional magnetic resonance imaging, we show for the first time that levels of musical expertise stepwise modulate higher order brain functioning. This suggests that degree of training intensity drives such cerebral plasticity. Participants (non-musicians, amateurs, and expert musicians) listened to a comprehensive set of specifically composed string quartets with hierarchically manipulated endings. In particular, we implemented 2 irregularities at musical closure that differed in salience but were both within the tonality of the piece (in-key). Behavioral sensitivity scores (d') of both transgressions perfectly separated participants according to their level of musical expertise. By contrasting brain responses to harmonic transgressions against regular endings, functional brain imaging data showed compelling evidence for stepwise modulation of brain responses by both violation strength and expertise level in a fronto-temporal network hosting universal functions of working memory and attention. Additional independent testing evidenced an advantage in visual working memory for the professionals, which could be predicted by musical training intensity. The here introduced findings of brain plasticity demonstrate the progressive impact of musical training on cognitive brain functions that may manifest well beyond the field of music processing.

Training of tonal similarity ratings in non-musicians: a "rapid learning" approach.

Although cognitive music psychology has a long tradition of expert-novice comparisons, experimental training studies are rare. Studies on the learning progress of trained novices in hearing harmonic relationships are still largely lacking. This paper presents a simple training concept using the example of tone/triad similarity ratings, demonstrating the gradual progress of non-musicians compared to musical experts: In a feedback-based "rapid learning" paradigm, participants had to decide for single tones and chords whether paired sounds matched each other well. Before and after the training sessions, they provided similarity judgments for a complete set of sound pairs. From these similarity matrices, individual relational sound maps, intended to display mental representations, were calculated by means of non-metric multidimensional scaling (NMDS), and were compared to an expert model through procrustean transformation. Approximately half of the novices showed substantial learning success, with some participants even reaching the level of professional musicians. Results speak for a fundamental ability to quickly train an understanding of harmony, show inter-individual differences in learning success, and demonstrate the suitability of the scaling method used for learning research in music and other domains. Results are discussed in the context of the "giftedness" debate.

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.

Absolute pitch--functional evidence of speech-relevant auditory acuity.

Absolute pitch (AP) has been shown to be associated with morphological changes and neurophysiological adaptations in the planum temporale, a cortical area involved in higher-order auditory and speech perception processes. The direct link between speech processing and AP has hitherto not been addressed. We provide first evidence that AP compared with relative pitch (RP) ability is associated with significantly different hemodynamic responses to complex speech sounds. By systematically varying the lexical and/or prosodic information of speech stimuli, we demonstrated consistent activation differences in AP musicians compared with RP musicians and nonmusicians. These differences relate to stronger activations in the posterior part of the middle temporal gyrus and weaker activations in the anterior mid-part of the superior temporal gyrus. Furthermore, this pattern is considerably modulated by the auditory acuity of AP. Our results suggest that the neural underpinnings of pitch processing expertise exercise a strong influence on propositional speech perception (sentence meaning).

White matter plasticity in the corticospinal tract of musicians: a diffusion tensor imaging study.

With the advent of diffusion tensor imaging (DTI), the study of plastic changes in white matter architecture due to long-term practice has attracted increasing interest. Professional musicians provide an ideal model for investigating white matter plasticity because of their early onset of extensive auditory and sensorimotor training. We performed fiber tractography and subsequent voxelwise analysis, region of interest (ROI) analysis, and detailed slicewise analysis of diffusion parameters in the corticospinal tract (CST) on 26 professional musicians and a control group of 13 participants. All analyses resulted in significantly lower fractional anisotropy (FA) values in both the left and the right CST in the musician group. Furthermore, a right-greater-than-left asymmetry of FA was observed regardless of group. In the musician group, diffusivity was negatively correlated with the onset of musical training in childhood. A subsequent median split into an early and a late onset musician group (median=7 years) revealed increased diffusivity in the CST of the early onset group as compared to both the late onset group and the controls. In conclusion, these DTI-based findings might indicate plastic changes in white matter architecture of the CST in professional musicians. Our results imply that training-induced changes in diffusion characteristics of the axonal membrane may lead to increased radial diffusivity as reflected in decreased FA values.

The plasticity of the superior longitudinal fasciculus as a function of musical expertise: a diffusion tensor imaging study.

Previous neuroimaging studies have demonstrated that musical expertise leads to functional alterations in language processing. We utilized diffusion tensor imaging (DTI) to investigate white matter plasticity in musicians with absolute pitch (AP), relative pitch and non-musicians. Using DTI, we analysed the fractional anisotropy (FA) of the superior longitudinal fasciculus (SLF), which is considered the most primary pathway for processing and production of speech and music. In association with different levels of musical expertise, we found that AP is characterized by a greater left than right asymmetry of FA in core fibres of the SLF. A voxel-based analysis revealed three clusters within the left hemisphere SLF that showed significant positive correlations with error rates only for AP-musicians in an AP-test, but not for musicians without AP. We therefore conclude that the SLF architecture in AP musicians is related to AP acuity. In order to reconcile our observations with general aspects of development of fibre bundles, we introduce the Pioneer Axon Thesis, a theoretical approach to formalize axonal arrangements of major white matter pathways.

The multiple synaesthete E.S.: neuroanatomical basis of interval-taste and tone-colour synaesthesia.

Synaesthesia is the involuntary physical experience of a crossmodal linkage such as when hearing a tone evokes the additional sensation of seeing a colour. We previously described a professional musician with absolute pitch perception who experiences both different tastes in response to hearing different tone intervals (e.g., major third and sweet) and the more common tone-colour synaesthesia in which each particular tone is linked to a specific colour (e.g., C and red). One of the current theories of synaesthesia proposes that local crossactivation or disinhibition of feedback occurs because of increased connectivity between relevant brain areas. Based on diffusion tensor and T1-weighted magnetic resonance imaging we performed fractional anisotropy (FA) analysis, probabilistic fibre tractography, and voxel-based morphometry in the synaesthete E.S. compared with 17 professional musicians and 20 normal control subjects using voxel-wise z-score transformations. We report increased FA and volumetric white (WM) and grey matter (GM) peculiarities in E.S.'s auditory and gustatory areas, hence explaining the interval-taste synaesthesia. Probabilistic fibre tractography revealed hyperconnectivity in bilateral perisylvian-insular regions in the synaesthete E.S. Differences in FA and volumetric WM and GM alterations in visual areas might represent the neuroarchitectural foundation of the tone-colour synaesthesia. Still unknown are the causes of the structural alterations, although an X-chromosomal linked dominant trait has been suggested. Whether hyperconnectivity occurs due to a failure in neural pruning or even synaptic sprouting remains to be shown. Our findings might have implications for the understanding of multimodal integration and may encourage similar research into dysfunctional perceptual phenomenon such as hallucinations in schizophrenics or in Charles Bonnet syndrome.