Music predictability and liking enhance pupil dilation and promote motor learning in non-musicians.

Humans can anticipate music and derive pleasure from it. Expectations facilitate the learning of movements associated with anticipated events, and they are also linked with reward, which may further facilitate learning of the anticipated rewarding events. The present study investigates the synergistic effects of predictability and hedonic responses to music on arousal and motor-learning in a naïve population. Novel melodies were manipulated in their overall predictability (predictable/unpredictable) as objectively defined by a model of music expectation, and ranked as high/medium/low liked based on participants' self-reports collected during an initial listening session. During this session, we also recorded ocular pupil size as an implicit measure of listeners' arousal. During the following motor task, participants learned to play target notes of the melodies on a keyboard (notes were of similar motor and musical complexity across melodies). Pupil dilation was greater for liked melodies, particularly when predictable. Motor performance was facilitated in predictable rather than unpredictable melodies, but liked melodies were learned even in the unpredictable condition. Low-liked melodies also showed learning but mostly in participants with higher scores of task perceived competence. Taken together, these results highlight  the effects of stimuli predictability on learning, which can be however overshadowed by the effects of stimulus liking or task-related intrinsic motivation.

Regional cerebellar volumes are related to early musical training and finger tapping performance.

The cerebellum has been associated with timing on the millisecond scale and with musical rhythm and beat processing. Early musical training (before age 7) is associated with enhanced rhythm synchronization performance and differences in cortical motor areas and the corpus callosum. In the present study, we examined the relationships between regional cerebellar volumes, early musical training, and timing performance. We tested adult musicians and non-musicians on a standard finger tapping task, and extracted cerebellar gray and white matter volumes using a novel multi-atlas automatic segmentation pipeline. We found that early-trained musicians had reduced volume in bilateral cerebellar white matter and right lobules IV, V and VI, compared to late-trained musicians. Strikingly, better timing performance, greater musical experience and an earlier age of start of musical training were associated with smaller cerebellar volumes. Better timing performance was specifically associated with smaller volumes of right lobule VI. Collectively, these findings support the sensitivity of the cerebellum to the age of initiation of musical training and suggest that lobule VI plays a role in timing. The smaller cerebellar volumes associated with musical training and timing performance may be a reflection of more efficiently implemented low-level timing and sensorimotor processes.

The role of musical training in emergent and event-based timing.

INTRODUCTION: Musical performance is thought to rely predominantly on event-based timing involving a clock-like neural process and an explicit internal representation of the time interval. Some aspects of musical performance may rely on emergent timing, which is established through the optimization of movement kinematics, and can be maintained without reference to any explicit representation of the time interval. We predicted that musical training would have its largest effect on event-based timing, supporting the dissociability of these timing processes and the dominance of event-based timing in musical performance. MATERIALS AND METHODS: We compared 22 musicians and 17 non-musicians on the prototypical event-based timing task of finger tapping and on the typically emergently timed task of circle drawing. For each task, participants first responded in synchrony with a metronome (Paced) and then responded at the same rate without the metronome (Unpaced). RESULTS: Analyses of the Unpaced phase revealed that non-musicians were more variable in their inter-response intervals for finger tapping compared to circle drawing. Musicians did not differ between the two tasks. Between groups, non-musicians were more variable than musicians for tapping but not for drawing. We were able to show that the differences were due to less timer variability in musicians on the tapping task. Correlational analyses of movement jerk and inter-response interval variability revealed a negative association for tapping and a positive association for drawing in non-musicians only. DISCUSSION: These results suggest that musical training affects temporal variability in tapping but not drawing. Additionally, musicians and non-musicians may be employing different movement strategies to maintain accurate timing in the two tasks. These findings add to our understanding of how musical training affects timing and support the dissociability of event-based and emergent timing modes.

Cerebellum and M1 interaction during early learning of timed motor sequences.

We used positron emission tomography (PET) to examine within-day learning of timed motor sequences. The results of this experiment are novel in showing an interaction between cerebellum and primary motor cortex (M1) during learning that appears to be mediated by the dentate nucleus (DN) and in demonstrating that activity in these regions is directly related to performance. Subjects were scanned during learning (LRN) across three blocks of practice and during isochronous (ISO) and perceptual (PER) baseline conditions. CBF was compared across blocks of learning and between the LRN and baseline conditions. Results demonstrated an interaction between the cerebellum and M1 such that earlier, poorer performance was associated with greater activity in the cerebellar hemispheres and later, better performance was associated with greater activity in M1. Inter-regional correlation analyses confirmed that as CBF in the cerebellum decreases, blood flow in M1 increases. Importantly, these analyses also revealed that activity in cerebellar cortex was positively correlated with activity in right DN and that DN activity was negatively correlated with blood flow in M1. Activity in the cerebellar hemispheres early in learning is likely related to error correction mechanisms which optimize movement kinematics resulting in improved performance. Concurrent DN activity may be related to encoding of this information and DN output to M1 may play a role in consolidation processes that lay down motor memories. Increased activity in M1 later in learning may reflect strengthening of synaptic connections associated with changes in motor maps that are characteristic of learning in both animals and humans.

Spatial localization after excision of human auditory cortex.

Neurophysiological and animal ablation studies concur that primary auditory cortex is necessary for computation of the spatial coordinates of a sound source. Human studies have reported conflicting findings but have often suffered from inadequate psychophysical measures and/or poor lesion localization. We tested patients with unilateral temporal lobe excisions either encroaching on or sparing Heschl's gyrus (HG), quantifying lesion extent using anatomical magnetic resonance imaging measures. Subjects performed two tasks. In the localization task, they heard single clicks in a free-field spatial array subtending 180 degrees of azimuth and indicated the perceived location with a laser pointer. In the discrimination task, two clicks were presented, and subjects indicated if they were in the same or different position. As a group, patients with right temporal excision, either encroaching onto HG or not, were significantly impaired in both hemifields in both tasks, although this was not true for all individuals. Patients with left temporal resections generally performed normally, although some of the patients with left HG excision showed impaired performance bilaterally, especially in the discrimination task. This pattern stands in marked contrast to previous studies showing significant preservation of localization in hemispherectomized patients. We conclude that (1) contrary to hypotheses derived from animal studies, human auditory spatial processes are dependent primarily on cortical areas within right superior temporal cortex, which encompass both spatial hemifields; (2) functional reorganization may not take place after restricted focal damage but only after more extensive early damage; and (3) the existence of individual differences likely illustrates differential patterns of functional lateralization and/or recovery.

Functional specificity in the right human auditory cortex for perceiving pitch direction.

Previous lesion and functional imaging studies in humans suggest a greater involvement of right rather than left auditory cortical areas in certain aspects of pitch processing. In the present study, adaptive psychophysical procedures were used to determine auditory perceptual thresholds in 14 neurologically normal subjects, and in 31 patients who had undergone surgical resection from either the right or left temporal lobe for the relief of intractable epilepsy. In a subset of the patients, the lesion encroached significantly upon the gyrus of Heschl or its underlying white matter as determined from MRI analysis. Subjects were asked to perform two different perceptual tasks on the same set of stimuli. In a pitch discrimination task, the subject had to decide whether two elements of a pure tone pair were the same or different. In a task requiring the judgement of direction of pitch change, subjects decided whether pitch rose or fell from the first tone to the second. Thresholds were determined by measuring the minimum pitch difference required for correct task performance. Mean thresholds in the pitch discrimination task did not differ between patient groups and control subjects. In contrast, patients with temporal lobe excisions that encroached upon the gyrus of Heschl in the right hemisphere (but not in the left) showed significantly elevated thresholds when judging the direction of pitch change. These findings support a specialization of function linked to right auditory cortical areas for the processing of pitch direction, and specifically suggest a dissociation between simple sensory discrimination and higher order perception.

The role of auditory cortex in retention of rhythmic patterns as studied in patients with temporal lobe removals including Heschl's gyrus.

This experiment examined the participation of the auditory cortex of the temporal lobe in the perception and retention of rhythmic patterns. Four patient groups were tested on a paradigm contrasting reproduction of auditory and visual rhythms: those with right or left anterior temporal lobe removals which included Heschl's gyrus (HG), the region of primary auditory cortex (RT-A and LT-A); and patients with right or left anterior temporal lobe removals which did not include HG (RT-a and LT-a). Estimation of lesion extent in HG using an MRI-based probabilistic map indicated that, in the majority of subjects, the lesion was confined to the anterior secondary auditory cortex located on the anterior-lateral extent of HG. On the rhythm reproduction task, RT-A patients were impaired in retention of auditory but not visual rhythms, particularly when accurate reproduction of stimulus durations was required. In contrast, LT-A patients as well as both RT-a and LT-a patients were relatively unimpaired on this task. None of the patient groups was impaired in the ability to make an adequate motor response. Further, they were unimpaired when using a dichotomous response mode, indicating that they were able to adequately differentiate the stimulus durations and, when given an alternative method of encoding, to retain them. Taken together, these results point to a specific role for the right anterior secondary auditory cortex in the retention of a precise analogue representation of auditory tonal patterns.

Cerebellar contributions to motor timing: a PET study of auditory and visual rhythm reproduction.

The perception and production of temporal patterns, or rhythms, is important for both music and speech. However, the way in which the human brain achieves accurate timing of perceptual input and motor output is as yet little understood. Central control of both motor timing and perceptual timing across modalities has been linked to both the cerebellum and the basal ganglia (BG). The present study was designed to test the hypothesized central control of temporal processing and to examine the roles of the cerebellum, BG, and sensory association areas. In this positron emission tomography (PET) activation paradigm, subjects reproduced rhythms of increasing temporal complexity that were presented separately in the auditory and visual modalities. The results provide support for a supramodal contribution of the lateral cerebellar cortex and cerebellar vermis to the production of a timed motor response, particularly when it is complex and/or novel. The results also give partial support to the involvement of BG structures in motor timing, although this may be more directly related to implementation of the motor response than to timing per se. Finally, sensory association areas and the ventrolateral frontal cortex were found to be involved in modality-specific encoding and retrieval of the temporal stimuli. Taken together, these results point to the participation of a number of neural structures in the production of a timed motor response from an external stimulus. The role of the cerebellum in timing is conceptualized not as a clock or counter but simply as the structure that provides the necessary circuitry for the sensory system to extract temporal information and for the motor system to learn to produce a precisely timed response.

Interhemispheric anatomical differences in human primary auditory cortex: probabilistic mapping and volume measurement from magnetic resonance scans.

The gyral morphology of the region of the primary auditory cortex (PAC) in the human brain is highly variable, and possible asymmetries between the hemispheres have been noted since the beginning of the century. We mapped the location and extent of PAC as identified from gross anatomical landmarks in magnetic resonance scans that had been transformed into Talairach-Tournoux stereotaxic space. Individual maps were averaged to produce a probabilistic map of the region which can be co-registered with any image of brain structure or function that has been similarly transformed. The map can be used to localize a region of interest, such as a lesion, or an activation focus from position omission tomography or functional magnetic resonance imaging, within a specified range of probability. We also measured the total volume of the region and found a significant L > R asymmetry both on average and in the majority of subjects. Automatic segmentation of the volumes into grey and white matter revealed larger volumes of white, but not grey matter on the left. This larger volume of cortical connecting fibres may be related to the known left-hemisphere dominance for speech, and a preferential role for left PAC in processing temporal aspects of auditory stimuli is suggested.