Not all errors are alike: modulation of error-related neural responses in musical joint action.

During joint action, the sense of agency enables interaction partners to implement corrective and adaptive behaviour in response to performance errors. When agency becomes ambiguous (e.g. when action similarity encourages perceptual self-other overlap), confusion as to who produced what may disrupt this process. The current experiment investigated how ambiguity of agency affects behavioural and neural responses to errors in a joint action domain where self-other overlap is common: musical duos. Pairs of pianists performed piano pieces in synchrony, playing either the same pitches (ambiguous agency) or different pitches (unambiguous agency) while electroencephalography (EEG) was recorded for each individual. Behavioural and event-related potential results showed no effects of the agency manipulation but revealed differences in how distinct error types are processed. Self-produced 'wrong note' errors (substitutions) were left uncorrected, showed post-error slowing and elicited an error-related negativity (ERN) peaking before erroneous keystrokes (pre-ERN). In contrast, self-produced 'extra note' errors (additions) exhibited pre-error slowing, error and post-error speeding, were rapidly corrected and elicited the ERN. Other-produced errors evoked a feedback-related negativity but no behavioural effects. Overall findings shed light upon how the nervous system supports fluent interpersonal coordination in real-time joint action by employing distinct mechanisms to manage different types of errors.

The influence of pitch feedback on learning of motor -timing and sequencing: A piano study with novices.

Audio-motor coordination is a fundamental requirement in the learning and execution of sequential actions such as music performance. Predictive motor control mechanisms determine the sequential content and timing of upcoming tones and thereby facilitate accurate performance. To study the role of auditory-motor predictions at early stages of acquiring piano performance skills, we conducted an experiment in which non-musicians learned to play a musical sequence on the piano in synchrony with a metronome. Three experimental conditions compared errors and timing. The first consisted of normal auditory feedback using conventional piano key-to-tone mappings. The second employed fixed-pitch auditory feedback consisting of a single tone that was given with each key stroke. In the third condition, for each key stroke a tone was randomly drawn from the set of tones associated with the normal sequence. The results showed that when auditory feedback tones were randomly assigned, participants produced more sequencing errors (i.e., a higher percentage of incorrect key strokes) compared to when auditory feedback was normal or consisted of a single tone of fixed pitch. Furthermore, synchronization with the metronome was most accurate in the fixed-pitch single-tone condition. These findings suggest that predictive motor control mechanisms support sequencing and timing, and that these sensorimotor processes are dissociable even at early stages of acquiring complex motor skills such as music performance.

Shared Neural Mechanisms for the Prediction of Own and Partner Musical Sequences after Short-term Piano Duet Training.

Predictive mechanisms in the human brain can be investigated using markers for prediction violations like the mismatch negativity (MMN). Short-term piano training increases the MMN for melodic and rhythmic deviations in the training material. This increase occurs only when the material is actually played, not when it is only perceived through listening, suggesting that learning predictions about upcoming musical events are derived from motor involvement. However, music is often performed in concert with others. In this case, predictions about upcoming actions from a partner are a crucial part of the performance. In the present experiment, we use magnetoencephalography (MEG) to measure MMNs to deviations in one's own and a partner's musical material after both engaged in musical duet training. Event-related field (ERF) results revealed that the MMN increased significantly for own and partner material suggesting a neural representation of the partner's part in a duet situation. Source analysis using beamforming revealed common activations in auditory, inferior frontal, and parietal areas, similar to previous results for single players, but also a pronounced contribution from the cerebellum. In addition, activation of the precuneus and the medial frontal cortex was observed, presumably related to the need to distinguish between own and partner material.

Differential processing of melodic, rhythmic and simple tone deviations in musicians--an MEG study.

Rhythm and melody are two basic characteristics of music. Performing musicians have to pay attention to both, and avoid errors in either aspect of their performance. To investigate the neural processes involved in detecting melodic and rhythmic errors from auditory input we tested musicians on both kinds of deviations in a mismatch negativity (MMN) design. We found that MMN responses to a rhythmic deviation occurred at shorter latencies than MMN responses to a melodic deviation. Beamformer source analysis showed that the melodic deviation activated superior temporal, inferior frontal and superior frontal areas whereas the activation pattern of the rhythmic deviation focused more strongly on inferior and superior parietal areas, in addition to superior temporal cortex. Activation in the supplementary motor area occurred for both types of deviations. We also recorded responses to similar pitch and tempo deviations in a simple, non-musical repetitive tone pattern. In this case, there was no latency difference between the MMNs and cortical activation was smaller and mostly limited to auditory cortex. The results suggest that prediction and error detection of musical stimuli in trained musicians involve a broad cortical network and that rhythmic and melodic errors are processed in partially different cortical streams.

Clinical investigations of receptive and expressive musical functions after stroke.

There is a long tradition of investigating various disorders of musical abilities after stroke. These impairments, associated with acquired amusia, can be highly selective, affecting only music perception (i.e., receptive abilities/functions) or expression (music production abilities), and some patients report that these may dramatically influence their emotional state. The aim of this study was to systematically test both the melodic and rhythmic domains of music perception and expression in left- and right-sided stroke patients compared to healthy subjects. Music perception was assessed using rhythmic and melodic discrimination tasks, while tests of expressive function involved the vocal or instrumental reproduction of rhythms and melodies. Our approach revealed deficits in receptive and expressive functions in stroke patients, mediated by musical expertise. Those patients who had experienced a short period of musical training in childhood and adolescence performed better in the receptive and expressive subtests compared to those without any previous musical training. While discrimination of specific musical patterns was unimpaired after a left-sided stroke, patients with a right-sided stroke had worse results for fine melodic and rhythmic analysis. In terms of expressive testing, the most consistent results were obtained from a test that required patients to reproduce sung melodies. This implies that the means of investigating production abilities can impact the identification of deficits.

Tic-reducing effects of music in patients with Tourette's syndrome: Self-reported and objective analysis.

BACKGROUND: Self-reports by musicians affected with Tourette's syndrome and other sources of anecdotal evidence suggest that tics stop when subjects are involved in musical activity. For the first time, we studied this effect systematically using a questionnaire design to investigate the subjectively assessed impact of musical activity on tic frequency (study 1) and an experimental design to confirm these results (study 2). METHODS: A questionnaire was sent to 29 patients assessing whether listening to music and musical performance would lead to a tic frequency reduction or increase. Then, a within-subject repeated measures design was conducted with eight patients. Five experimental conditions were tested: baseline, musical performance, short time period after musical performance, listening to music and music imagery. Tics were counted based on videotapes. RESULTS: Analysis of the self-reports (study 1) yielded in a significant tic reduction both by listening to music and musical performance. In study 2, musical performance, listening to music and mental imagery of musical performance reduced tic frequency significantly. We found the largest reduction in the condition of musical performance, when tics almost completely stopped. Furthermore, we could find a short-term tic decreasing effect after musical performance. CONCLUSIONS: Self-report assessment revealed that active and passive participation in musical activity can significantly reduce tic frequency. Experimental testing confirmed patients' perception. Active and passive participation in musical activity reduces tic frequency including a short-term lasting tic decreasing effect. Fine motor control, focused attention and goal directed behavior are believed to be relevant factors for this observation.

Rhythmic and melodic deviations in musical sequences recruit different cortical areas for mismatch detection.

The mismatch negativity (MMN), an event-related potential (ERP) representing the violation of an acoustic regularity, is considered as a pre-attentive change detection mechanism at the sensory level on the one hand and as a prediction error signal on the other hand, suggesting that bottom-up as well as top-down processes are involved in its generation. Rhythmic and melodic deviations within a musical sequence elicit a MMN in musically trained subjects, indicating that acquired musical expertise leads to better discrimination accuracy of musical material and better predictions about upcoming musical events. Expectation violations to musical material could therefore recruit neural generators that reflect top-down processes that are based on musical knowledge. We describe the neural generators of the musical MMN for rhythmic and melodic material after a short-term sensorimotor-auditory (SA) training. We compare the localization of musical MMN data from two previous MEG studies by applying beamformer analysis. One study focused on the melodic harmonic progression whereas the other study focused on rhythmic progression. The MMN to melodic deviations revealed significant right hemispheric neural activation in the superior temporal gyrus (STG), inferior frontal cortex (IFC), and the superior frontal (SFG) and orbitofrontal (OFG) gyri. IFC and SFG activation was also observed in the left hemisphere. In contrast, beamformer analysis of the data from the rhythm study revealed bilateral activation within the vicinity of auditory cortices and in the inferior parietal lobule (IPL), an area that has recently been implied in temporal processing. We conclude that different cortical networks are activated in the analysis of the temporal and the melodic content of musical material, and discuss these networks in the context of the dual-pathway model of auditory processing.

A beamformer analysis of MEG data reveals frontal generators of the musically elicited mismatch negativity.

To localize the neural generators of the musically elicited mismatch negativity with high temporal resolution we conducted a beamformer analysis (Synthetic Aperture Magnetometry, SAM) on magnetoencephalography (MEG) data from a previous musical mismatch study. The stimuli consisted of a six-tone melodic sequence comprising broken chords in C- and G-major. The musical sequence was presented within an oddball paradigm in which the last tone was lowered occasionally (20%) by a minor third. The beamforming analysis revealed significant right hemispheric neural activation in the superior temporal (STC), inferior frontal (IFC), superior frontal (SFC) and orbitofrontal (OFC) cortices within a time window of 100-200 ms after the occurrence of a deviant tone. IFC and SFC activation was also observed in the left hemisphere. The pronounced early right inferior frontal activation of the auditory mismatch negativity has not been shown in MEG studies so far. The activation in STC and IFC is consistent with earlier electroencephalography (EEG), optical imaging and functional magnetic resonance imaging (fMRI) studies that reveal the auditory and inferior frontal cortices as main generators of the auditory MMN. The observed right hemispheric IFC is also in line with some previous music studies showing similar activation patterns after harmonic syntactic violations. The results demonstrate that a deviant tone within a musical sequence recruits immediately a distributed neural network in frontal and prefrontal areas suggesting that top-down processes are involved when expectation violation occurs within well-known stimuli.

Cortical plasticity induced by short-term multimodal musical rhythm training.

Performing music is a multimodal experience involving the visual, auditory, and somatosensory modalities as well as the motor system. Therefore, musical training is an excellent model to study multimodal brain plasticity. Indeed, we have previously shown that short-term piano practice increase the magnetoencephalographic (MEG) response to melodic material in novice players. Here we investigate the impact of piano training using a rhythmic-focused exercise on responses to rhythmic musical material. Musical training with non musicians was conducted over a period of two weeks. One group (sensorimotor-auditory, SA) learned to play a piano sequence with a distinct musical rhythm, another group (auditory, A) listened to, and evaluated the rhythmic accuracy of the performances of the SA-group. Training-induced cortical plasticity was evaluated using MEG, comparing the mismatch negativity (MMN) in response to occasional rhythmic deviants in a repeating rhythm pattern before and after training. The SA-group showed a significantly greater enlargement of MMN and P2 to deviants after training compared to the A- group. The training-induced increase of the rhythm MMN was bilaterally expressed in contrast to our previous finding where the MMN for deviants in the pitch domain showed a larger right than left increase. The results indicate that when auditory experience is strictly controlled during training, involvement of the sensorimotor system and perhaps increased attentional recources that are needed in producing rhythms lead to more robust plastic changes in the auditory cortex compared to when rhythms are simply attended to in the auditory domain in the absence of motor production.

Processing of complex auditory patterns in musicians and nonmusicians.

In the present study we investigated the capacity of the memory store underlying the mismatch negativity (MMN) response in musicians and nonmusicians for complex tone patterns. While previous studies have focused either on the kind of information that can be encoded or on the decay of the memory trace over time, we studied capacity in terms of the length of tone sequences, i.e., the number of individual tones that can be fully encoded and maintained. By means of magnetoencephalography (MEG) we recorded MMN responses to deviant tones that could occur at any position of standard tone patterns composed of four, six or eight tones during passive, distracted listening. Whereas there was a reliable MMN response to deviant tones in the four-tone pattern in both musicians and nonmusicians, only some individuals showed MMN responses to the longer patterns. This finding of a reliable capacity of the short-term auditory store underlying the MMN response is in line with estimates of a three to five item capacity of the short-term memory trace from behavioural studies, although pitch and contour complexity covaried with sequence length, which might have led to an understatement of the reported capacity. Whereas there was a tendency for an enhancement of the pattern MMN in musicians compared to nonmusicians, a strong advantage for musicians could be shown in an accompanying behavioural task of detecting the deviants while attending to the stimuli for all pattern lengths, indicating that long-term musical training differentially affects the memory capacity of auditory short-term memory for complex tone patterns with and without attention. Also, a left-hemispheric lateralization of MMN responses in the six-tone pattern suggests that additional networks that help structuring the patterns in the temporal domain might be recruited for demanding auditory processing in the pitch domain.

Auditory-somatosensory integration and cortical plasticity in musical training.

Learning to play a musical instrument requires complex multimodal skills involving simultaneous perception of several sensory modalities: auditory, visual, and somatosensory as well as the motor system. Musical training thus provides an adequate neuroscientific model to study multimodal integration and plasticity in musical training. The aim of this study was to investigate the impact of short-term uni- and multimodal musical training on auditory-somatosensory integration and plasticity. Two groups of nonmusicians were musically trained. The first group (sensorimotor-auditory group, SA) learned to play a musical sequence on the piano, whereas the second one (auditory group, A) actively listened to and made judgments about the correctness of the music. The training-induced cortical plasticity effect was assessed by recording musically elicited mismatch negativity (MMN) from magnetoencephalographic (MEG) measurements before and after training. The SA group showed significant enlargement of MMN after training compared to the A group, reflecting greater enhancement of musical representations in auditory cortex after sensorimotor-auditory training compared to mere auditory training. This study demonstrates that the sensorimotor and auditory systems integrate and that this multimodal training causes cortical reorganizational changes in the auditory cortex over and above the changes introduced by auditory training alone.

Imagery mismatch negativity in musicians.

The present study investigated musical imagery in musicians and nonmusicians by means of magnetoencephalography (MEG). We used a new paradigm in which subjects had to continue familiar melodies in their mind and then judged if a further presented tone was a correct continuation of the melody. Incorrect tones elicited an imagery mismatch negativity (iMMN) in musicians but not in nonmusicians. This finding suggests that the MMN component can be based on an imagined instead of a sensory memory trace and that imagery of music is modulated by musical expertise.

Looking for a pattern: an MEG study on the abstract mismatch negativity in musicians and nonmusicians.

BACKGROUND: The mismatch negativity (MMN) is an early component of event-related potentials/fields, which can be observed in response to violations of regularities in sound sequences. The MMN can be elicited by simple feature (e.g. pitch) deviations in standard oddball paradigms as well as by violations of more complex sequential patterns. By means of magnetoencephalography (MEG) we investigated if a pattern MMN could be elicited based on global rather than local probabilities and if the underlying ability to integrate long sequences of tones is enhanced in musicians compared to nonmusicians. RESULTS: A pattern MMN was observed in response to violations of a predominant sequential pattern (AAAB) within a standard oddball tone sequence consisting of only two different tones. This pattern MMN was elicited even though the probability of pattern deviants in the sequence was as high as 0.5. Musicians showed more leftward-lateralized pattern MMN responses, which might be due to a stronger specialization of the ability to integrate information in a sequence of tones over a long time range. CONCLUSION: The results indicate that auditory grouping and the probability distribution of possible patterns within a sequence influence the expectations about upcoming tones, and that the MMN might also be based on global statistical knowledge instead of a local memory trace. The results also show that auditory grouping based on sequential regularities can occur at a much slower presentation rate than previously presumed, and that probability distributions of possible patterns should be taken into account even for the construction of simple oddball sequences.

Neural basis of music imagery and the effect of musical expertise.

Although the influence of long-term musical training on the processing of heard music has been the subject of many studies, the neural basis of music imagery and the effect of musical expertise remain insufficiently understood. By means of magnetoencephalography (MEG) we compared musicians and nonmusicians in a musical imagery task with familiar melodies. Subjects listened to the beginnings of the melodies, continued them in their imagination and then heard a tone which was either a correct or an incorrect further continuation of the melody. Only in musicians was the imagery of these melodies strong enough to elicit an early preattentive brain response to unexpected incorrect continuations of the imagined melodies; this response, the imagery mismatch negativity (iMMN), peaked approximately 175 ms after tone onset and was right-lateralized. In contrast to previous studies the iMMN was not based on a heard but on a purely imagined memory trace. Our results suggest that in trained musicians imagery and perception rely on similar neuronal correlates, and that the musicians' intense musical training has modified this network to achieve a superior ability for imagery and preattentive processing of music.

Cortical plasticity induced by short-term unimodal and multimodal musical training.

Learning to play a musical instrument requires complex multimodal skills involving simultaneous perception of several sensory modalities: auditory, visual, somatosensory, as well as the motor system. Therefore, musical training provides a good and adequate neuroscientific model to study multimodal brain plasticity effects in humans. Here, we investigated the impact of short-term unimodal and multimodal musical training on brain plasticity. Two groups of nonmusicians were musically trained over the course of 2 weeks. One group [sensorimotor-auditory (SA)] learned to play a musical sequence on the piano, whereas the other group [auditory (A)] listened to and made judgments about the music that had been played by participants of the sensorimotor-auditory group. Training-induced cortical plasticity was assessed by recording the musically elicited mismatch negativity (MMNm) from magnetoencephalographic measurements before and after training. SA and A groups showed significantly different cortical responses after training. Specifically, the SA group showed significant enlargement of MMNm after training compared with the A group, reflecting greater enhancement of musical representations in auditory cortex after sensorimotor-auditory training compared with after mere auditory training. Thus, we have experimentally demonstrated that not only are sensorimotor and auditory systems connected, but also that sensorimotor-auditory training causes plastic reorganizational changes in the auditory cortex over and above changes introduced by auditory training alone.