Cross-modal reorganization after sensory deprivation is a model for understanding brain plasticity. Although it is a well-documented phenomenon, we still know little of the mechanisms underlying it or the factors that constrain and promote it. Using fMRI, we identified visual motion-related activity in 17 early-deaf and 17 hearing adults. We found that, in the deaf, the posterior superior temporal gyrus (STG) was responsive to visual motion. We compared functional connectivity of this reorganized cortex between groups to identify differences in functional networks associated with reorganization. In the deaf more than the hearing, the STG displayed increased functional connectivity with a region in the calcarine fissure. We also explored the role of hearing aid use, a factor that may contribute to variability in cross-modal reorganization. We found that both the cross-modal activity in STG and the functional connectivity between STG and calcarine cortex correlated with duration of hearing aid use, supporting the hypothesis that residual hearing affects cross-modal reorganization. We conclude that early auditory deprivation alters not only the organization of auditory regions but also the interactions between auditory and primary visual cortex and that auditory input, as indexed by hearing aid use, may inhibit cross-modal reorganization in early-deaf people.

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.

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.

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.

We used fMRI to investigate the neuronal correlates of encoding and recognizing heard and imagined melodies. Ten participants were shown lyrics of familiar verbal tunes; they either heard the tune along with the lyrics, or they had to imagine it. In a subsequent surprise recognition test, they had to identify the titles of tunes that they had heard or imagined earlier. The functional data showed substantial overlap during melody perception and imagery, including secondary auditory areas. During imagery compared with perception, an extended network including pFC, SMA, intraparietal sulcus, and cerebellum showed increased activity, in line with the increased processing demands of imagery. Functional connectivity of anterior right temporal cortex with frontal areas was increased during imagery compared with perception, indicating that these areas form an imagery-related network. Activity in right superior temporal gyrus and pFC was correlated with the subjective rating of imagery vividness. Similar to the encoding phase, the recognition task recruited overlapping areas, including inferior frontal cortex associated with memory retrieval, as well as left middle temporal gyrus. The results present new evidence for the cortical network underlying goal-directed auditory imagery, with a prominent role of the right pFC both for the subjective impression of imagery vividness and for on-line mental monitoring of imagery-related activity in auditory areas.

Two fMRI experiments explored the neural substrates of a musical imagery task that required manipulation of the imagined sounds: temporal reversal of a melody. Musicians were presented with the first few notes of a familiar tune (Experiment 1) or its title (Experiment 2), followed by a string of notes that was either an exact or an inexact reversal. The task was to judge whether the second string was correct or not by mentally reversing all its notes, thus requiring both maintenance and manipulation of the represented string. Both experiments showed considerable activation of the superior parietal lobe (intraparietal sulcus) during the reversal process. Ventrolateral and dorsolateral frontal cortices were also activated, consistent with the memory load required during the task. We also found weaker evidence for some activation of right auditory cortex in both studies, congruent with results from previous simpler music imagery tasks. We interpret these results in the context of other mental transformation tasks, such as mental rotation in the visual domain, which are known to recruit the intraparietal sulcus region, and we propose that this region subserves general computations that require transformations of a sensory input. Mental imagery tasks may thus have both task or modality-specific components as well as components that supersede any specific codes and instead represent amodal mental manipulation.

Much is known about the motor system and its role in simple movement execution. However, little is understood about the neural systems underlying auditory-motor integration in the context of musical rhythm, or the enhanced ability of musicians to execute precisely timed sequences. Using functional magnetic resonance imaging, we investigated how performance and neural activity were modulated as musicians and nonmusicians tapped in synchrony with progressively more complex and less metrically structured auditory rhythms. A functionally connected network was implicated in extracting higher-order features of a rhythm's temporal structure, with the dorsal premotor cortex mediating these auditory-motor interactions. In contrast to past studies, musicians recruited the prefrontal cortex to a greater degree than nonmusicians, whereas secondary motor regions were recruited to the same extent. We argue that the superior ability of musicians to deconstruct and organize a rhythm's temporal structure relates to the greater involvement of the prefrontal cortex mediating working memory.

The neural processes underlying the ability to divide attention between multiple sensory modalities remain poorly understood. To investigate the role of the dorsolateral prefrontal cortex (DLPFC) in bimodal divided attention, we completed two repetitive transcranial magnetic stimulation (rTMS) studies. We tested the hypothesis that the DLPFC is necessary in the ability to divide attention across modalities. This hypothesis originated as a result of a previous fMRI study in which the posterior DLPFC was active during a bimodal divided attention condition [Johnson, J. A., & Zatorre, R. J. Neural substrates for dividing and focusing attention between simultaneous auditory and visual events. Neuroimage , 2006]. In the current experiments, two separate groups of subjects underwent 10 min of slow rTMS to temporarily disrupt function of the DLPFC. In both groups, the ability to divide attention between unrelated auditory and visual stimuli decreased following DLPFC disruption compared to control site stimulation. Specifically, the ability to divide attention between modalities was hindered, leading to a pattern of behavior similar to bimodal selective attention (ability to attend to one or the other modality but not both). We discuss possible roles of the posterior DLPFC in bimodal divided attention and conclude that the area may be functioning to support the increased working memory load associated with divided, compared to selective attention.

Vigilance behavior, or watch keeping, involves the focusing of attention on the detection of subtle changes in the environment that occur over a long period of time. We investigated the time course of changes in brain activity during the continuous performance of a 60-min auditory vigilance task. The task required the detection of an intensity drop that occurred in 5% of the auditory stimuli. Six 1-min samples of cerebral blood flow (CBF) and electroencephalographic (EEG) activity were obtained at l0-min intervals during the vigilance performance. Changes in CBF were measured by means of positron emission tomography (PET). Performance data (hits, false alarms, reaction time) were analyzed across six 10-min blocks. Eight healthy male volunteers participated in the study. During the 60-min test, the number of correct detections (hits) did not change, but both the reaction time and EEG activity in the theta (4 to 7 Hz) range progressively increased across testing. CBF in several subcortical structures (thalamus, substantia innominata, and putamen) and cortical areas (ventrolateral, dorsolateral, and orbital frontal cortex; parietal cortex; and temporal cortex) decreased as a function of time-on-task; changes in the cortical regions were limited to the right hemisphere. Blood flow also decreased in the temporalis muscles. At the same time, CBF increased in several visual cortical areas including the left and right fusiform gyri. Furthermore, the thalamic blood-flow response co-varied with that in the substantia innominata, the ponto-mesencephalic tegmentum, and the anterior cingulate cortex. The right ventrolateral-frontal blood-flow response covaried with that in the right parietal, orbitofrontal, and dorsolateral frontal cortex. 'Iko main conclusions are drawn from the obtained data. First, we suggest that the observed time-related changes in reaction time, EEG activity, and blood flow in the temporalis muscles are related to changes in the level of arousal (alertness) and that CBF changes in the thalamus-related neural circuitry represent a brain correlate of such changes. Second, we speculate that time-related CBF decreases in cortical regions of the right hemisphere underlie a shift from controlled to automatic attentional processing of the auditory stimuli.

Neuropsychological studies have suggested that imagery processes may be mediated by neuronal mechanisms similar to those used in perception. To test this hypothesis, and to explore the neural basis for song imagery, 12 normal subjects were scanned using the water bolus method to measure cerebral blood flow (CBF) during the performance of three tasks. In the control condition subjects saw pairs of words on each trial and judged which word was longer. In the perceptual condition subjects also viewed pairs of words, this time drawn from a familiar song; simultaneously they heard the corresponding song, and their task was to judge the change in pitch of the two cued words within the song. In the imagery condition, subjects performed precisely the same judgment as in the perceptual condition, but with no auditory input. Thus, to perform the imagery task correctly an internal auditory representation must be accessed. Paired-image subtraction of the resulting pattern of CBF, together with matched MRI for anatomical localization, revealed that both perceptual and imagery. tasks produced similar patterns of CBF changes, as compared to the control condition, in keeping with the hypothesis. More specifically, both perceiving and imagining songs are associated with bilateral neuronal activity in the secondary auditory cortices, suggesting that processes within these regions underlie the phenomenological impression of imagined sounds. Other CBF foci elicited in both tasks include areas in the left and right frontal lobes and in the left parietal lobe, as well as the supplementary motor area. This latter region implicates covert vocalization as one component of musical imagery. Direct comparison of imagery and perceptual tasks revealed CBF increases in the inferior frontal polar cortex and right thalamus. We speculate that this network of regions may be specifically associated with retrieval and/or generation of auditory information from memory.