Dopamine dysregulation in Parkinson's disease flattens the pleasurable urge to move to musical rhythms.

The pleasurable urge to move to music (PLUMM) activates motor and reward areas of the brain and is thought to be driven by predictive processes. Dopamine in motor and limbic networks is implicated in beat-based timing and music-induced pleasure, suggesting a central role of basal ganglia (BG) dopaminergic systems in PLUMM. This study tested this hypothesis by comparing PLUMM in participants with Parkinson's disease (PD), age-matched controls, and young controls. Participants listened to musical sequences with varying rhythmic and harmonic complexity (low, medium and high), and rated their experienced pleasure and urge to move to the rhythm. In line with previous results, healthy younger participants showed an inverted U-shaped relationship between rhythmic complexity and ratings, with preference for medium complexity rhythms, while age-matched controls showed a similar, but weaker, inverted U-shaped response. Conversely, PD showed a significantly flattened response for both the urge to move and pleasure. Crucially, this flattened response could not be attributed to differences in rhythm discrimination and did not reflect an overall decrease in ratings. For harmonic complexity, PD showed a negative linear pattern for both the urge to move and pleasure while healthy age-matched controls showed the same pattern for pleasure and an inverted U for the urge to move. This contrasts with the pattern observed in young healthy controls in previous studies, suggesting that both healthy aging and PD also influence affective responses to harmonic complexity. Together, these results support the role of dopamine within cortico-striatal circuits in the predictive processes that form the link between the perceptual processing of rhythmic patterns and the affective and motor responses to rhythmic music.

Using cortico-cerebellar structural patterns to classify early- and late-trained musicians.

A body of current evidence suggests that there is a sensitive period for musical training: people who begin training before the age of seven show better performance on tests of musical skill, and also show differences in brain structure-especially in motor cortical and cerebellar regions-compared with those who start later. We used support vector machine models-a subtype of supervised machine learning-to investigate distributed patterns of structural differences between early-trained (ET) and late-trained (LT) musicians and to better understand the age boundaries of the sensitive period for early musicianship. After selecting regions of interest from the cerebellum and cortical sensorimotor regions, we applied recursive feature elimination with cross-validation to produce a model which optimally and accurately classified ET and LT musicians. This model identified a combination of 17 regions, including 9 cerebellar and 8 sensorimotor regions, and maintained a high accuracy and sensitivity (true positives, i.e., ET musicians) without sacrificing specificity (true negatives, i.e., LT musicians). Critically, this model-which defined ET musicians as those who began their training before the age of 7-outperformed all other models in which age of start was earlier or later (between ages 5-10). Our model's ability to accurately classify ET and LT musicians provides additional evidence that musical training before age 7 affects cortico-cerebellar structure in adulthood, and is consistent with the hypothesis that connected brain regions interact during development to reciprocally influence brain and behavioral maturation.

Rhythm and time in the premotor cortex.

Many animals can encode temporal intervals and use them to plan their actions, but only humans can flexibly extract a regular beat from complex patterns, such as musical rhythms. Beat-based timing is hypothesized to rely on the integration of sensory information with temporal information encoded in motor regions such as the medial premotor cortex (MPC), but how beat-based timing might be encoded in neuronal populations is mostly unknown. Gámez and colleagues show that the MPC encodes temporal information via a population code visible as circular trajectories in state space; these patterns may represent precursors to more-complex skills such as beat-based timing.

Effector-independent brain network for auditory-motor integration: fMRI evidence from singing and cello playing.

Many everyday tasks share high-level sensory goals but differ in the movements used to accomplish them. One example of this is musical pitch regulation, where the same notes can be produced using the vocal system or a musical instrument controlled by the hands. Cello playing has previously been shown to rely on brain structures within the singing network for performance of single notes, except in areas related to primary motor control, suggesting that the brain networks for auditory feedback processing and sensorimotor integration may be shared (Segado et al. 2018). However, research has shown that singers and cellists alike can continue singing/playing in tune even in the absence of auditory feedback (Chen et al. 2013, Kleber et al. 2013), so different paradigms are required to test feedback monitoring and control mechanisms. In singing, auditory pitch feedback perturbation paradigms have been used to show that singers engage a network of brain regions including anterior cingulate cortex (ACC), anterior insula (aINS), and intraparietal sulcus (IPS) when compensating for altered pitch feedback, and posterior superior temporal gyrus (pSTG) and supramarginal gyrus (SMG) when ignoring it (Zarate et al. 2005, 2008). To determine whether the brain networks for cello playing and singing directly overlap in these sensory-motor integration areas, in the present study expert cellists were asked to compensate for or ignore introduced pitch perturbations when singing/playing during fMRI scanning. We found that cellists were able to sing/play target tones, and compensate for and ignore introduced feedback perturbations equally well. Brain activity overlapped for singing and playing in IPS and SMG when compensating, and pSTG and dPMC when ignoring; differences between singing/playing across all three conditions were most prominent in M1, centered on the relevant motor effectors (hand, larynx). These findings support the hypothesis that pitch regulation during cello playing relies on structures within the singing network and suggests that differences arise primarily at the level of forward motor control.

Arcuate fasciculus architecture is associated with individual differences in pre-attentive detection of unpredicted music changes.

The mismatch negativity (MMN) is an event related brain potential (ERP) elicited by unpredicted sounds presented in a sequence of repeated auditory stimuli. The neural sources of the MMN have been previously attributed to a fronto-temporo-parietal network which crucially overlaps with the so-called auditory dorsal stream, involving inferior and middle frontal, inferior parietal, and superior and middle temporal regions. These cortical areas are structurally connected by the arcuate fasciculus (AF), a three-branch pathway supporting the feedback-feedforward loop involved in auditory-motor integration, auditory working memory, storage of acoustic templates, as well as comparison and update of those templates. Here, we characterized the individual differences in the white-matter macrostructural properties of the AF and explored their link to the electrophysiological marker of passive change detection gathered in a melodic multifeature MMN-EEG paradigm in 26 healthy young adults without musical training. Our results show that left fronto-temporal white-matter connectivity plays an important role in the pre-attentive detection of rhythm modulations within a melody. Previous studies have shown that this AF segment is also critical for language processing and learning. This strong coupling between structure and function in auditory change detection might be related to life-time linguistic (and possibly musical) exposure and experiences, as well as to timing processing specialization of the left auditory cortex. To the best of our knowledge, this is the first time in which the relationship between neurophysiological (EEG) and brain white-matter connectivity indexes using DTI-tractography are studied together. Thus, the present results, although still exploratory, add to the existing evidence on the importance of studying the constraints imposed on cognitive functions by the underlying structural connectivity.

Neural network retuning and neural predictors of learning success associated with cello training.

The auditory and motor neural systems are closely intertwined, enabling people to carry out tasks such as playing a musical instrument whose mapping between action and sound is extremely sophisticated. While the dorsal auditory stream has been shown to mediate these audio-motor transformations, little is known about how such mapping emerges with training. Here, we use longitudinal training on a cello as a model for brain plasticity during the acquisition of specific complex skills, including continuous and many-to-one audio-motor mapping, and we investigate individual differences in learning. We trained participants with no musical background to play on a specially designed MRI-compatible cello and scanned them before and after 1 and 4 wk of training. Activation of the auditory-to-motor dorsal cortical stream emerged rapidly during the training and was similarly activated during passive listening and cello performance of trained melodies. This network activation was independent of performance accuracy and therefore appears to be a prerequisite of music playing. In contrast, greater recruitment of regions involved in auditory encoding and motor control over the training was related to better musical proficiency. Additionally, pre-supplementary motor area activity and its connectivity with the auditory cortex during passive listening before training was predictive of final training success, revealing the integrative function of this network in auditory-motor information processing. Together, these results clarify the critical role of the dorsal stream and its interaction with auditory areas in complex audio-motor learning.

What you learn & when you learn it: Impact of early bilingual & music experience on the structural characteristics of auditory-motor pathways.

Music and language engage the dorsal auditory pathway, linked by the arcuate fasciculus (AF). Sustained practice in these activities can modify brain structure, depending on length of experience but also age of onset (AoO). To study the impact of early experience on brain structure we manually dissected the AF in bilinguals with and without music training (MT) who differed in the AoO of their second language (L2), or MT. We found the usual left-greater-than-right asymmetry in the volume of the long segment (LS) of the AF across all groups. However, simultaneous exposure to two languages from birth enhanced this leftward asymmetry, while early start of MT (≤7) enhanced the right LS macrostructure, reducing the normative asymmetry. Thus, immersive exposure to an L2 in the first year of life can produce long-term plastic effects on the left LS, which is considered to be largely under genetic control, while deliberate music training in early childhood alters the right LS, whose structure appears more open to experience. These findings show that AoO of specific experience plays a key role in a complex gene-environment interaction model where normative brain maturation is differentially impacted by diverse intensive auditory-motor experiences at different points during development.

The sensation of groove engages motor and reward networks.

The sensation of groove has been defined as the pleasurable desire to move to music, suggesting that both motor timing and reward processes are involved in this experience. Although many studies have investigated rhythmic timing and musical reward separately, none have examined whether the associated cortical and subcortical networks are engaged while participants listen to groove-based music. In the current study, musicians and non-musicians listened to and rated experimentally controlled groove-based stimuli while undergoing functional magnetic resonance imaging. Medium complexity rhythms elicited higher ratings of pleasure and wanting to move and were associated with activity in regions linked to beat perception and reward, as well as prefrontal and parietal regions implicated in generating and updating stimuli-based expectations. Activity in basal ganglia regions of interest, including the nucleus accumbens, caudate and putamen, was associated with ratings of pleasure and wanting to move, supporting their important role in the sensation of groove. We propose a model in which different cortico-striatal circuits interact to support the mechanisms underlying groove, including internal generation of the beat, beat-based expectations, and expectation-based affect. These results show that the sensation of groove is supported by motor and reward networks in the brain and, along with our proposed model, suggest that the basal ganglia are crucial nodes in networks that interact to generate this powerful response to music.

Efficacy of Auditory versus Motor Learning for Skilled and Novice Performers.

Humans must learn a variety of sensorimotor skills, yet the relative contributions of sensory and motor information to skill acquisition remain unclear. Here we compare the behavioral and neural contributions of perceptual learning to that of motor learning, and we test whether these contributions depend on the expertise of the learner. Pianists and nonmusicians learned to perform novel melodies on a piano during fMRI scanning in four learning conditions: listening (auditory learning), performing without auditory feedback (motor learning), performing with auditory feedback (auditory-motor learning), or observing visual cues without performing or listening (cue-only learning). Visual cues were present in every learning condition and consisted of musical notation for pianists and spatial cues for nonmusicians. Melodies were performed from memory with no visual cues and with auditory feedback (recall) five times during learning. Pianists showed greater improvements in pitch and rhythm accuracy at recall during auditory learning compared with motor learning. Nonmusicians demonstrated greater rhythm improvements at recall during auditory learning compared with all other learning conditions. Pianists showed greater primary motor response at recall during auditory learning compared with motor learning, and response in this region during auditory learning correlated with pitch accuracy at recall and with auditory-premotor network response during auditory learning. Nonmusicians showed greater inferior parietal response during auditory compared with auditory-motor learning, and response in this region correlated with pitch accuracy at recall. Results suggest an advantage for perceptual learning compared with motor learning that is both general and expertise-dependent. This advantage is hypothesized to depend on feedforward motor control systems that can be used during learning to transform sensory information into motor production.

Auditory prediction cues motor preparation in the absence of movements.

There is increasing evidence for integrated representation of sensory and motor information in the brain, and that seeing or hearing action-related stimuli may automatically cue the movements required to respond to or produce them. In this study we tested whether anticipation of tones in a known melody automatically activates corresponding motor representations in a predictive way, in preparation for potential upcoming movements. Therefore, we trained 20 non-musicians (8 men, 12 women) to play a simple melody. Then, while they passively listened to the learned or unlearned melodies, we applied single pulse transcranial magnetic stimulation (TMS) over M1 to measure motor evoked potentials from the associated finger muscle either preceding or following the onset of individual tones. Our results show that listening to the learned melody increased corticospinal excitability for specific finger muscles before tone onset. This demonstrates that predictable auditory information can activate motor representations in an anticipatory muscle-specific manner, even in the absence of intention to move. This suggests that the motor system is involved in the prediction of sensory events, likely based on auditory-parietal-prefrontal feedforward/feedback loops that automatically prepare predictable sound-related actions independent of actual execution and the associated auditory feedback. Overall, we propose that multimodal forward models of upcoming sounds and actions support motor preparation, facilitate error detection and correction, and guide perception.

White-matter structural connectivity predicts short-term melody and rhythm learning in non-musicians.

Music learning has received increasing attention in the last decades due to the variety of functions and brain plasticity effects involved during its practice. Most previous reports interpreted the differences between music experts and laymen as the result of training. However, recent investigations suggest that these differences are due to a combination of genetic predispositions with the effect of music training. Here, we tested the relationship of the dorsal auditory-motor pathway with individual behavioural differences in short-term music learning. We gathered structural neuroimaging data from 44 healthy non-musicians (28 females) before they performed a rhythm- and a melody-learning task during a single behavioural session, and manually dissected the arcuate fasciculus (AF) in both hemispheres. The macro- and microstructural organization of the AF (i.e., volume and FA) predicted the learning rate and learning speed in the musical tasks, but only in the right hemisphere. Specifically, the volume of the right anterior segment predicted the synchronization improvement during the rhythm task, the FA in the right long segment was correlated with the learning rate in the melody task, and the volume and FA of the right whole AF predicted the learning speed during the melody task. This is the first study finding a specific relation between different branches within the AF and rhythmic and melodic materials. Our results support the relevant function of the AF as the structural correlate of both auditory-motor transformations and the feedback-feedforward loop, and suggest a crucial involvement of the anterior segment in error-monitoring processes related to auditory-motor learning. These findings have implications for both the neuroscience of music field and second-language learning investigations.

Sensorimotor integration is enhanced in dancers and musicians.

Studying individuals with specialized training, such as dancers and musicians, provides an opportunity to investigate how intensive practice of sensorimotor skills affects behavioural performance across various domains. While several studies have found that musicians have improved motor, perceptual and sensorimotor integration skills compared to untrained controls, fewer studies have examined the effect of dance training on such skills. Moreover, no study has specifically compared the effects of dance versus music training on perceptual or sensorimotor performance. To this aim, in the present study, expert dancers, expert musicians and untrained controls were tested on a range of perceptual and sensorimotor tasks designed to discriminate performance profiles across groups. Dancers performed better than musicians and controls on a dance imitation task (involving whole-body movement), but musicians performed better than dancers and controls on a musical melody discrimination task as well as on a rhythm synchronization task (involving finger tapping). These results indicate that long-term intensive dance and music training are associated with distinct enhancements in sensorimotor skills. This novel work advances knowledge of the effects of long-term dance versus music training and has potential applications in therapies for motor disorders.

Effects of age and cognitive load on response reprogramming.

A dual-task paradigm was used to examine the effect of cognitive load on motor reprogramming. We propose that in the face of conflict, both executive control and motor control mechanisms become more interconnected in the process of reprogramming motor behaviors. If so, one would expect a concurrent cognitive load to compromise younger adults' (YAs) motor reprogramming ability and further exacerbate the response reprogramming ability of older adults (OAs). Nineteen YAs and 14 OAs overlearned a sequence of key presses. Deviations of the practiced sequence were introduced to assess motor reprogramming ability. A Serial Sevens Test was used as the cognitive load. A 3D motion capture system was used to parse finger movements into planning and motor execution times. Global response time analysis revealed that under single-task conditions, during prepotent transitions, OAs responded as quickly as YAs, but they were disproportionately worse than YAs during conflict transitions. Under dual-task conditions, YAs performance became more similar to that of OAs. Movement data were decomposed into planning and movement time, revealing that under single-task conditions, when responding to conflicting stimuli YAs reduced their movement time in order to compensate for delayed planning time; however, additional cognitive load prevented them from exhibiting this compensatory hastening on conflict transitions. We propose that age-related declines in response reprogramming may be linked to reduced cognitive capacity. Current findings suggest that cognitive capacity, reduced in the case of OAs or YAs under divided attention conditions, influences the ability to flexibly adapt to conflicting conditions.

Early musical training and white-matter plasticity in the corpus callosum: evidence for a sensitive period.

Training during a sensitive period in development may have greater effects on brain structure and behavior than training later in life. Musicians are an excellent model for investigating sensitive periods because training starts early and can be quantified. Previous studies suggested that early training might be related to greater amounts of white matter in the corpus callosum, but did not control for length of training or identify behavioral correlates of structural change. The current study compared white-matter organization using diffusion tensor imaging in early- and late-trained musicians matched for years of training and experience. We found that early-trained musicians had greater connectivity in the posterior midbody/isthmus of the corpus callosum and that fractional anisotropy in this region was related to age of onset of training and sensorimotor synchronization performance. We propose that training before the age of 7 years results in changes in white-matter connectivity that may serve as a scaffold upon which ongoing experience can build.

Early musical training is linked to gray matter structure in the ventral premotor cortex and auditory-motor rhythm synchronization performance.

Evidence in animals and humans indicates that there are sensitive periods during development, times when experience or stimulation has a greater influence on behavior and brain structure. Sensitive periods are the result of an interaction between maturational processes and experience-dependent plasticity mechanisms. Previous work from our laboratory has shown that adult musicians who begin training before the age of 7 show enhancements in behavior and white matter structure compared with those who begin later. Plastic changes in white matter and gray matter are hypothesized to co-occur; therefore, the current study investigated possible differences in gray matter structure between early-trained (ET; <7) and late-trained (LT; >7) musicians, matched for years of experience. Gray matter structure was assessed using voxel-wise analysis techniques (optimized voxel-based morphometry, traditional voxel-based morphometry, and deformation-based morphometry) and surface-based measures (cortical thickness, surface area and mean curvature). Deformation-based morphometry analyses identified group differences between ET and LT musicians in right ventral premotor cortex (vPMC), which correlated with performance on an auditory motor synchronization task and with age of onset of musical training. In addition, cortical surface area in vPMC was greater for ET musicians. These results are consistent with evidence that premotor cortex shows greatest maturational change between the ages of 6-9 years and that this region is important for integrating auditory and motor information. We propose that the auditory and motor interactions required by musical practice drive plasticity in vPMC and that this plasticity is greatest when maturation is near its peak.

ERP evidence of adaptive changes in error processing and attentional control during rhythm synchronization learning.

The ability to detect and use information from errors is essential during the acquisition of new skills. There is now a wealth of evidence about the brain mechanisms involved in error processing. However, the extent to which those mechanisms are engaged during the acquisition of new motor skills remains elusive. Here we examined rhythm synchronization learning across 12 blocks of practice in musically naïve individuals and tracked changes in ERP signals associated with error-monitoring and error-awareness across distinct learning stages. Synchronization performance improved with practice, and performance improvements were accompanied by dynamic changes in ERP components related to error-monitoring and error-awareness. Early in learning, when performance was poor and the internal representations of the rhythms were weaker we observed a larger error-related negativity (ERN) following errors compared to later learning. The larger ERN during early learning likely results from greater conflict between competing motor responses, leading to greater engagement of medial-frontal conflict monitoring processes and attentional control. Later in learning, when performance had improved, we observed a smaller ERN accompanied by an enhancement of a centroparietal positive component resembling the P3. This centroparietal positive component was predictive of participant's performance accuracy, suggesting a relation between error saliency, error awareness and the consolidation of internal templates of the practiced rhythms. Moreover, we showed that during rhythm learning errors led to larger auditory evoked responses related to attention orientation which were triggered automatically and which were independent of the learning stage. The present study provides crucial new information about how the electrophysiological signatures related to error-monitoring and error-awareness change during the acquisition of new skills, extending previous work on error processing and cognitive control mechanisms to a more ecologically valid context.

Attention, working memory, and inhibitory control in aging: Comparing amateur singers, instrumentalists, and active controls.

Despite the ubiquity of musical activities, little is known about the specificity of their association with executive functions. In this cross-sectional study, we examined this relationship as a function of age. Our main hypotheses were that executive functions would decline in older age, that this relationship would be reduced in singers and instrumentalists compared to nonmusician active controls, and that the amount of musical experience would be more strongly associated with executive functions compared to the specific type of activity. A sample of 122 cognitively healthy adults aged 20-88 years was recruited, consisting of 39 amateur singers, 43 amateur instrumentalists, and 40 nonmusician controls. Tests of auditory processing speed, auditory selective attention, auditory and visual inhibitory control, and auditory working memory were administered. The results confirm a negative relationship between age and executive functions. While musicians' advantages were found in selective attention, inhibitory control, and auditory working memory, these advantages were specific rather than global. Furthermore, most of these advantages were independent of age and experience. Finally, there were only limited differences between instrumentalists and singers, suggesting that the relationship between music-making activities and executive functions may be, at least in part, general as opposed to activity-specific.

Music reward sensitivity is associated with greater information transfer capacity within dorsal and motor white matter networks in musicians.

There are pronounced differences in the degree to which individuals experience music-induced pleasure which are linked to variations in structural connectivity between auditory and reward areas. However, previous studies exploring the link between white matter structure and music reward sensitivity (MRS) have relied on standard diffusion tensor imaging methods, which present challenges in terms of anatomical accuracy and interpretability. Further, the link between MRS and connectivity in regions outside of auditory-reward networks, as well as the role of musical training, have yet to be investigated. Therefore, we investigated the relation between MRS and structural connectivity in a large number of directly segmented and anatomically verified white matter tracts in musicians (n = 24) and non-musicians (n = 23) using state-of-the-art tract reconstruction and fixel-based analysis. Using a manual tract-of-interest approach, we additionally tested MRS-white matter associations in auditory-reward networks seen in previous studies. Within the musician group, there was a significant positive relation between MRS and fiber density and cross section in the right middle longitudinal fascicle connecting auditory and inferior parietal cortices. There were also positive relations between MRS and fiber-bundle cross-section in tracts connecting the left thalamus to the ventral precentral gyrus and connecting the right thalamus to the right supplementary motor area, however, these did not survive FDR correction. These results suggest that, within musicians, dorsal auditory and motor networks are crucial to MRS, possibly via their roles in top-down predictive processing and auditory-motor transformations.

Interacting cortical and basal ganglia networks underlying finding and tapping to the musical beat.

Humans are able to find and tap to the beat of musical rhythms varying in complexity from children's songs to modern jazz. Musical beat has no one-to-one relationship with auditory features-it is an abstract perceptual representation that emerges from the interaction between sensory cues and higher-level cognitive organization. Previous investigations have examined the neural basis of beat processing but have not tested the core phenomenon of finding and tapping to the musical beat. To test this, we used fMRI and had musicians find and tap to the beat of rhythms that varied from metrically simple to metrically complex-thus from a strong to a weak beat. Unlike most previous studies, we measured beat tapping performance during scanning and controlled for possible effects of scanner noise on beat perception. Results showed that beat finding and tapping recruited largely overlapping brain regions, including the superior temporal gyrus (STG), premotor cortex, and ventrolateral PFC (VLPFC). Beat tapping activity in STG and VLPFC was correlated with both perception and performance, suggesting that they are important for retrieving, selecting, and maintaining the musical beat. In contrast BG activity was similar in all conditions and was not correlated with either perception or production, suggesting that it may be involved in detecting auditory temporal regularity or in associating auditory stimuli with a motor response. Importantly, functional connectivity analyses showed that these systems interact, indicating that more basic sensorimotor mechanisms instantiated in the BG work in tandem with higher-order cognitive mechanisms in PFC.

Repetition suppression in auditory-motor regions to pitch and temporal structure in music.

Music performance requires control of two sequential structures: the ordering of pitches and the temporal intervals between successive pitches. Whether pitch and temporal structures are processed as separate or integrated features remains unclear. A repetition suppression paradigm compared neural and behavioral correlates of mapping pitch sequences and temporal sequences to motor movements in music performance. Fourteen pianists listened to and performed novel melodies on an MR-compatible piano keyboard during fMRI scanning. The pitch or temporal patterns in the melodies either changed or repeated (remained the same) across consecutive trials. We expected decreased neural response to the patterns (pitch or temporal) that repeated across trials relative to patterns that changed. Pitch and temporal accuracy were high, and pitch accuracy improved when either pitch or temporal sequences repeated over trials. Repetition of either pitch or temporal sequences was associated with linear BOLD decrease in frontal-parietal brain regions including dorsal and ventral premotor cortex, pre-SMA, and superior parietal cortex. Pitch sequence repetition (in contrast to temporal sequence repetition) was associated with linear BOLD decrease in the intraparietal sulcus (IPS) while pianists listened to melodies they were about to perform. Decreased BOLD response in IPS also predicted increase in pitch accuracy only when pitch sequences repeated. Thus, behavioral performance and neural response in sensorimotor mapping networks were sensitive to both pitch and temporal structure, suggesting that pitch and temporal structure are largely integrated in auditory-motor transformations. IPS may be involved in transforming pitch sequences into spatial coordinates for accurate piano performance.