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We applied fMRI and diffusion-weighted MRI to study the segregation of cognitive and motor functions in the human cerebro-cerebellar system. Our fMRI results show that a load increase in a nonverbal auditory working memory task is associated with enhanced brain activity in the parietal, dorsal premotor, and lateral prefrontal cortices and in lobules VII-VIII of the posterior cerebellum, whereas a sensory-motor control task activated the motor/somatosensory, medial prefrontal, and posterior cingulate cortices and lobules V/VI of the anterior cerebellum. The load-dependent activity in the crus I/II had a specific relationship with cognitive performance: This activity correlated negatively with load-dependent increase in RTs. This correlation between brain activity and RTs was not observed in the sensory-motor task in the activated cerebellar regions. Furthermore, probabilistic tractography analysis of the diffusion-weighted MRI data suggests that the tracts between the cerebral and the cerebellar areas exhibiting cognitive load-dependent and sensory-motor activity are mainly projected via separated pontine (feed-forward tracts) and thalamic (feedback tracts) nuclei. The tractography results also indicate that the crus I/II in the posterior cerebellum is linked with the lateral prefrontal areas activated by cognitive load increase, whereas the anterior cerebellar lobe is not. The current results support the view that cognitive and motor functions are segregated in the cerebellum. On the basis of these results and theories of the function of the cerebellum, we suggest that the posterior cerebellar activity during a demanding cognitive task is involved with optimization of the response speed.

Musical competence may confer cognitive advantages that extend beyond processing of familiar musical sounds. Behavioural evidence indicates a general enhancement of both working memory and attention in musicians. It is possible that musicians, due to their training, are better able to maintain focus on task-relevant stimuli, a skill which is crucial to working memory. We measured the blood oxygenation-level dependent (BOLD) activation signal in musicians and non-musicians during working memory of musical sounds to determine the relation among performance, musical competence and generally enhanced cognition. All participants easily distinguished the stimuli. We tested the hypothesis that musicians nonetheless would perform better, and that differential brain activity would mainly be present in cortical areas involved in cognitive control such as the lateral prefrontal cortex. The musicians performed better as reflected in reaction times and error rates. Musicians also had larger BOLD responses than non-musicians in neuronal networks that sustain attention and cognitive control, including regions of the lateral prefrontal cortex, lateral parietal cortex, insula, and putamen in the right hemisphere, and bilaterally in the posterior dorsal prefrontal cortex and anterior cingulate gyrus. The relationship between the task performance and the magnitude of the BOLD response was more positive in musicians than in non-musicians, particularly during the most difficult working memory task. The results confirm previous findings that neural activity increases during enhanced working memory performance. The results also suggest that superior working memory task performance in musicians rely on an enhanced ability to exert sustained cognitive control. This cognitive benefit in musicians may be a consequence of focused musical training.

We investigated whether music experts and laypersons differ with regard to aesthetic evaluation of musical sequences. 16 music experts and 16 music laypersons judged the aesthetic value (beauty judgment task) as well as the harmonic correctness (correctness judgment task) of chord sequences. The sequences consisted of five chords with the final chord sounding congruous, ambiguous or incongruous relative to the harmonic context established by the preceding four chords. On behavioural measures, few differences were observed between experts and laypersons. However, several differences in event-related potential (ERP) parameters were observed in auditory, cognitive and aesthetic processing of chord cadences between experts and laypersons. First, established ERP effects known to reflect the processing of harmonic rule violation were investigated. Here, differences between the groups were observed in the processing of the mild violation - experts and laypersons differed in their early brain responses to the beginning of the chord sequence. Furthermore, ERP data indicated distinctions between experts and laypersons in aesthetic evaluation at three different stages. Firstly, during the interval of task-cue presentation, a stronger contingent negative variation (CNV) to the beauty judgment task was observed for experts, indicating that experts invest more effort into preparation for aesthetic processes than into correctness judgments. Secondly, during the first four chords, preparation for the correctness judgment required more exertion on the laypersons' side. Thirdly, during the last chord, laypersons showed a larger late and widespread positivity for the beauty compared to the correctness judgment, indicating a stronger reliance on internal affective states while forming a judgment., (Copyright 2010 Elsevier B.V. All rights reserved.)

At the level of the auditory cortex, musicians discriminate pitch changes more accurately than nonmusicians. However, it is not agreed upon how sound familiarity and musical expertise interact in the formation of pitch-change discrimination skills, that is, whether musicians possess musical pitch discrimination abilities that are generally more accurate than in nonmusicians or, alternatively, whether they may be distinguished from nonmusicians particularly with respect to the discrimination of nonprototypical sounds that do not play a reference role in Western tonal music. To resolve this, we used magnetoencephalography (MEG) to measure the change-related magnetic mismatch response (MMNm) in musicians and nonmusicians to two nonprototypical chords, a "dissonant" chord containing a highly unpleasant interval and a "mistuned" chord including a mistuned pitch, and a minor chord, all inserted in a context of major chords. Major and minor are the most frequently used chords in Western tonal music which both musicians and nonmusicians are most familiar with, whereas the other chords are more rarely encountered in tonal music. The MMNm was stronger in musicians than in nonmusicians in response to the dissonant and mistuned chords, whereas no group difference was found in the MMNm strength to minor chords. Correspondingly, the length of musical training correlated with the MMNm strength for the dissonant and mistuned chords only. Our findings provide evidence for superior automatic discrimination of nonprototypical chords in musicians. Most likely, this results from a highly sophisticated auditory system in musicians allowing a more efficient discrimination of chords deviating from the conventional categories of tonal music.

In the neurosciences of music, a consensus on the nature of affective states during music listening has not been reached. What is undeniable is that subjective affective states can be triggered by various and even opposite musical events. Here we review the few recent studies on the neural determinants of subjective affective processes of music, contrasted with early automatic neural processes linked to the objective universal properties of music. In particular, we focus on the evaluative judgments of music by subjects according to its aesthetic and structural values, on music-specific emotions felt by listeners, and on conscious liking. We then discuss and seek to stimulate further research on the interplay between the emotional attributes of music and the subjective cognitive, psychological, and biographic factors, such as personality traits and cognitive strategies of listening. We finally draw the neuroscientist's attention to the sociocultural context as a relevant variable to study when considering music as an aesthetic domain.

Background noise has become part of our everyday life in modern societies. Its presence affects both the ability to concentrate and communicate. Some individuals, like children, the elderly, and non-native speakers have pronounced problems in noisy environments. Here we review evidence suggesting that background noise has both transient and sustained detrimental effects on central speech processing. Studies on the effects of noise on neural processes have demonstrated hemispheric reorganization in speech processing in adult individuals during background noise. During noise, the well-known left hemisphere dominance in speech discrimination became right hemisphere preponderant. Furthermore, long-term exposure to noise has a persistent effect on the brain organization of speech processing and attention control. These results both stress the importance to re-evaluate which noise levels can be considered safe for brain functions and raise concerns on the speech and cognitive abilities of individuals living in noisy environments.

We analyzed the processes of making aesthetic judgments of music, focusing on the differences between music experts and laypersons. Sixteen students of musicology and 16 control subjects (also students) judged the aesthetic value as well as the harmonic correctness of chord sequences. Event-related potential (ERP) data indicate differences between experts and laypersons in making aesthetic judgments at three different processing stages. Additionally, effects of expertise on ERP components that have previously been proven to be sensitive to musical training were replicated. The study thus provides insights into the effects of musical expertise on neural correlates of aesthetic music processing.

Goal-directed behavior lowers activity in brain areas that include the medial frontal cortex, the medial and lateral parietal cortex, and limbic and paralimbic brain regions, commonly referred to as the "default network." These activity decreases are believed to reflect the interruption of processes that are ongoing when the mind is in a restful state. Previously, the nature of these processes was probed by varying cognitive task parameters, but the presence of emotional processes, while often assumed, was little investigated. With fMRI, we studied the effect of systematic variations of both cognitive load and emotional stimulus connotation on task-related decreases in the default network by employing an auditory working memory (WM) task with musical sounds. The performance of the WM task, compared to passive listening, lowered the activity in medial and lateral, prefrontal, parietal, temporal, and limbic regions. In a subset of these regions, the magnitude of decrease depended on the memory load; the greater the cognitive load, the larger the magnitude of the observed decrease. Furthermore, in the right amygdala and the left precuneus, areas previously associated with processing of unpleasant dissonant musical sounds, there was an interaction between the experimental condition and the stimulus type. The current results are consistent with the previously reported effect of task difficulty on task-related brain activation decreases. The results also indicate that task-related decreases may be further modulated by the emotional stimulus connotation.

Like language, music engagement is universal, complex and present early in life. However, approximately 4% of the general population experiences a lifelong deficit in music perception that cannot be explained by hearing loss, brain damage, intellectual deficiencies or lack of exposure. This musical disorder, commonly known as tone-deafness and now termed congenital amusia, affects mostly the melodic pitch dimension. Congenital amusia is hereditary and is associated with abnormal grey and white matter in the auditory cortex and the inferior frontal cortex. In order to relate these anatomical anomalies to the behavioural expression of the disorder, we measured the electrical brain activity of amusic subjects and matched controls while they monitored melodies for the presence of pitch anomalies. Contrary to current reports, we show that the amusic brain can track quarter-tone pitch differences, exhibiting an early right-lateralized negative brain response. This suggests near-normal neural processing of musical pitch incongruities in congenital amusia. It is important because it reveals that the amusic brain is equipped with the essential neural circuitry to perceive fine-grained pitch differences. What distinguishes the amusic from the normal brain is the limited awareness of this ability and the lack of responsiveness to the semitone changes that violate musical keys. These findings suggest that, in the amusic brain, the neural pitch representation cannot make contact with musical pitch knowledge along the auditory-frontal neural pathway.

In Western tonal music, the rules of harmony determine the order and music-structural importance of events in a musical piece: for instance, the tonic chord, built on the first note of the diatonic scale, is usually placed at the end of chord sequences. A brain response termed the early right anterior negativity (ERAN) is elicited when a harmonically incongruous chord is inserted within or at the end of a musical sequence. The present study was conducted to test whether the ERAN reflects the processing of harmony rather than the building of a tonal context and whether the ERAN is also elicited by violations of the tuning of the sounds upon which harmony is based. To this aim, ten subjects listened to musical sequences containing either expected chords only, a harmonically incongruous chord in one of three positions within the cadence, or a harmonically congruous but mistuned chord in one of the three positions. Simultaneously, the electroencephalograph (EEG) was recorded. Incongruous chords violating the rules of harmony elicited a bilateral early anterior negativity, the amplitude of which depended on the degree of the harmony violation. On the contrary, mistuned chords, violating the rule of relations between all the sounds in the sequences, elicited a bilateral fronto-central negativity (the mismatch negativity, or MMN). The MMN was not modulated by the position of the violation within the musical sequence and had a longer peak latency than the anterior negativity elicited by the harmony rule violations. In conclusion, violations of the harmony and tuning rules of Western tonal music were found to generate specific and distinct electric responses in the human brain.

Timbre is a multidimensional perceptual attribute of complex tones that characterizes the identity of a sound source. Our study explores the representation in auditory sensory memory of three timbre dimensions (acoustically related to attack time, spectral centroid, and spectrum fine structure), using the mismatch negativity (MMN) component of the auditory event-related potential. MMN is elicited by a discriminable change in a sound sequence and reflects the detection of the discrepancy between the current stimulus and traces in auditory sensory memory. The stimuli used in the present study were carefully controlled synthetic tones. MMNs were recorded after changes along each of the three timbre dimensions and their combinations. Additivity of unidimensional MMNs and dipole modeling results suggest partially separate MMN generators for different timbre dimensions, reflecting their mainly separate processing in auditory sensory memory. The results expand to timbre dimensions a property of separation of the representation in sensory memory that has already been reported between basic perceptual attributes (pitch, loudness, duration, and location) of sound sources.

While listening to music, we immediately detect 'wrong' tones that do not match our expectations based on the prior context. This study aimed to determine whether such expectations can occur preattentively, as indexed by event-related potentials (ERPs), and whether these are modulated by attentional processes. To this end, we recorded ERPs in nonmusicians while they were presented with unfamiliar melodies, containing either a pitch deviating from the equal-tempered chromatic scale (out-of-tune) or a pitch deviating from the diatonic scale (out-of-key). ERPs were recorded in a passive experiment in which subjects were distracted from the sounds and in an active experiment in which they were judging how incongruous each melody was. In both the experiments, pitch incongruities elicited an early frontal negativity that was not modulated by attentional focus. This early negativity, closely corresponding to the mismatch negativity (MMN) of the ERPs, was mainly originated in the auditory cortex and occurred in response to both pitch violations but with larger amplitude for the more salient out-of-tune pitch than the less salient out-of-key pitch. Attentional processes leading to the conscious access of musical scale information were indexed by the late parietal positivity (resembling the P600 of the ERPs) elicited in response to both incongruous pitches in the active experiment only. Our results indicate that the relational properties of the musical scale are quickly and automatically extracted by the auditory cortex even before the intervention of focused attention.

Musicians and nonmusicians listened to major, minor, and dissonant musical chords while their BOLD brain responses were registered with functional magnetic resonance imaging. In both groups of listeners, minor and dissonant chords, compared with major chords, elicited enhanced responses in several brain areas, including the amygdala, retrosplenial cortex, brain stem, and cerebellum, during passive listening but not during memorization of the chords. The results indicate that (1) neural processing in emotion-related brain areas is activated even by single chords, (2) emotion processing is enhanced in the absence of cognitive requirements, and (3) musicians and nonmusicians do not differ in their neural responses to single musical chords during passive listening.

The role of attention in conjoining features of an object has been a topic of much debate. Studies using the mismatch negativity (MMN), an index of detecting acoustic deviance, suggested that the conjunctions of auditory features are preattentively represented in the brain. These studies, however, used sequentially presented sounds and thus are not directly comparable with visual studies of feature integration. Therefore, the current study presented an array of spatially distributed sounds to determine whether the auditory features of concurrent sounds are correctly conjoined without focal attention directed to the sounds. Two types of sounds differing from each other in timbre and pitch were repeatedly presented together while subjects were engaged in a visual n-back working-memory task and ignored the sounds. Occasional reversals of the frequent pitch-timbre combinations elicited MMNs of a very similar amplitude and latency irrespective of the task load. This result suggested preattentive integration of auditory features. However, performance in a subsequent target-search task with the same stimuli indicated the occurrence of illusory conjunctions. The discrepancy between the results obtained with and without focal attention suggests that illusory conjunctions may occur during voluntary access to the preattentively encoded object representations.

Objective: Long-term exposure to noise may cause an altered hemispheric lateralization of speech processing even in silent conditions. We examined whether this lateralization shift is speech specific or occurs also for other sounds., Methods: Brain responses from 10 healthy noise-exposed workers (>5 years) and 10 matched controls were recorded with a 32-channel electroencephalogram in two conditions, one including standard and deviant speech sounds, the other non-speech sounds, with novel sounds in both., Results: The deviant-sound elicited mismatch negativity (MMN) was larger to non-speech than speech sounds in control subjects, while it did not differ between the sound types in the noise-exposed subjects. Moreover, the MMN to speech sounds was lateralized to the right hemisphere in exposed workers, while it was left-hemisphere predominant in control subjects. No group topography difference was found for non-speech sounds. The deviant sounds that were close in formant space to the standards elicited a longer MMN latency in both speech and non-speech conditions in exposed subjects than controls. No group differences were found for cortical responses to novel sounds., Conclusions: Long-term noise exposure altered the strength and the hemispheric organization of speech-sound discrimination and decreased the speed of sound-change processing., Significance: Subpathological changes in cortical responses to sounds may occur even in subjects without a peripheral damage but continuously exposed to noisy auditory environments.

This study aimed at determining how the human brain automatically processes phoneme categories irrespective of the large acoustic inter-speaker variability. Subjects were presented with 450 different speech stimuli, equally distributed across the [a], [i], and [u] vowel categories, and each uttered by a different male speaker. A 306-channel magnetoencephalogram (MEG) was used to record N1m, the magnetic counterpart of the N1 component of the auditory event-related potential (ERP). The N1m amplitude and source locations differed between vowel categories. We also found that the spectrum dissimilarities were reproduced in the cortical representations of the large set of the phonemes used in this study: vowels with similar spectral envelopes had closer cortical representations than those whose spectral differences were the largest. Our data further extend the notion of differential cortical representations in response to vowel categories, previously demonstrated by using only one or a few tokens representing each category.

We determined whether the human N1 evoked by tones with different frequencies might be affected by a brief discrimination-training at one specific frequency. During 1 h training, subjects learned to discriminate a 1062 Hz tone from another tone. Before and after training, subjects heard for 26 min tones with a frequency of 1000 Hz, replaced every sixth one by test tones with frequencies randomly and equiprobably chosen as 835, 886, 941, 1000, 1062, 1128 or 1198 Hz. The N1 to the test tone was larger when its frequency was further from the repeating frequency. After training N1 s were attenuated to all tones except the trained and repeated ones, indicating a refractory frequency effect, long-term habituation, and sensitization to the repeated and trained tones.

So far studies about noise effects on speech perception have been focused on the development of hearing loss and other related pathologies. However, behavioral findings indicate that speech perception and intelligibility are disrupted in presence of noise in certain clinical groups in a way that is unpredictable on the basis of the audiogram performed in silence. This observation suggests that even soft noise can modify the cerebral mechanisms underlying speech information processing. Confirming the behavioral results, electric and magnetic brain measurements showed that noise presented to healthy subjects decreases the amplitude and increases the latency of brain responses to speech sounds. Recently, further experimental data allowed us to have a deeper knowledge of the neural mechanisms interventing in difficult listening conditions. In particolar, speech sounds presented in noisy background evoke bioelectric responses in neural populations located in the right cerebral hemisphere, that otherwise would not be activated. These findings can in future contribute to unveal new aspects of the noise pathology and its assessment in an early pre-clinical stage.

We wished to determine whether multiple sound patterns can be simultaneously represented in the temporary auditory buffer (auditory sensory memory), when subjects have no task related to the sounds. To this end we used the mismatch negativity (MMN) event-related potential, an electric brain response elicited when a frequent sound is infrequently replaced by a different sound. The MMN response is based on the presence of the auditory sensory memory trace of the frequent sounds, which exists whether or not these sounds are in the focus of the subject's attention. Subjects watching a movie were presented with sound sequences consisting of two frequent sound patterns, each formed of four different tones and an infrequent pattern consisting of the first two tones of one of the frequent sound pattern and the last two tones of the other frequent sound pattern. The infrequent sound pattern elicited an MMN, indicating that multiple sound patterns are formed at an early, largely automatic stage of auditory processing., (Copyright 2002 Lippincott Williams & Wilkins)

We investigated the brain mechanisms enabling one automatically discriminate phoneme category irrespective of the large inter-speaker variability in the acoustic features of the voices. For this purpose, subjects were presented with 450 different speech stimuli, each uttered by a different speaker, belonging to three vowel categories, while a 306-channel magnetoencephalogram (MEG) was obtained to record the magnetic counterpart of the mismatch negativity (MMNm), elicited only when sensory memory traces for repetitive sounds are formed in the auditory cortex. Despite this wide acoustic variation, category changes elicited prominent MMNm responses, which were considerably stronger in the left than in the right hemisphere in the right-handed subjects. These results implicate the presence of long-term memory traces for vowels, which can recognize the vowel-specific invariant code enabling correct vowel percept even in the presence of realistic acoustic variation., (Copyright 2002 Lippincott Williams & Wilkins)