Degree of musical expertise modulates higher order brain functioning.

Using functional magnetic resonance imaging, we show for the first time that levels of musical expertise stepwise modulate higher order brain functioning. This suggests that degree of training intensity drives such cerebral plasticity. Participants (non-musicians, amateurs, and expert musicians) listened to a comprehensive set of specifically composed string quartets with hierarchically manipulated endings. In particular, we implemented 2 irregularities at musical closure that differed in salience but were both within the tonality of the piece (in-key). Behavioral sensitivity scores (d') of both transgressions perfectly separated participants according to their level of musical expertise. By contrasting brain responses to harmonic transgressions against regular endings, functional brain imaging data showed compelling evidence for stepwise modulation of brain responses by both violation strength and expertise level in a fronto-temporal network hosting universal functions of working memory and attention. Additional independent testing evidenced an advantage in visual working memory for the professionals, which could be predicted by musical training intensity. The here introduced findings of brain plasticity demonstrate the progressive impact of musical training on cognitive brain functions that may manifest well beyond the field of music processing.

Rhythm evokes action: early processing of metric deviances in expressive music by experts and laymen revealed by ERP source imaging.

To examine how musical expertise tunes the brain to subtle metric anomalies in an ecological musical context, we presented piano compositions ending on standard and deviant cadences (endings) to expert pianists and musical laymen, while high-density EEG was recorded. Temporal expectancies were manipulated by substituting standard "masculine" cadences at metrically strong positions with deviant, metrically unaccented, "feminine" cadences. Experts detected metrically deviant cadences better than laymen. Analyses of event-related potentials demonstrated that an early P3a-like component (~150-300 ms), elicited by musical closure, was significantly enhanced at frontal and parietal electrodes in response to deviant endings in experts, whereas a reduced response to deviance occurred in laymen. Putative neuronal sources contributing to the modulation of this component were localized in a network of brain regions including bilateral supplementary motor areas, middle and posterior cingulate cortex, precuneus, associative visual areas, as well as in the right amygdala and insula. In all these regions, experts showed enhanced responses to metric deviance. Later effects demonstrated enhanced activations within the same brain network, as well as higher processing speed for experts. These results suggest that early brain responses to metric deviance in experts may rely on motor representations mediated by the supplementary motor area and motor cingulate regions, in addition to areas involved in self-referential imagery and relevance detection. Such motor representations could play a role in temporal sensory prediction evolved from musical training and suggests that rhythm evokes action more strongly in highly trained instrumentalists.

Neuroanatomy of hemispatial neglect and its functional components: a study using voxel-based lesion-symptom mapping.

Spatial neglect is a perplexing neuropsychological syndrome, in which patients fail to detect (and/or respond to) stimuli located contralaterally to their (most often right) hemispheric lesion. Neglect is characterized by a wide heterogeneity, and a role for multiple components has been suggested, but the exact nature of the critical components remains unclear. Moreover, many different lesion sites have been reported, leading to enduring controversies about the relative contribution of different cortical and/or subcortical brain regions. Here we report a systematic anatomo-functional study of 80 patients with a focal right hemisphere stroke, who were examined by a series of neuropsychological tests assessing different clinical manifestations of neglect. We first performed a statistical factorial analysis of their behavioural performance across all tests, in order to break down neglect symptoms into coherent profiles of co-varying deficits. We then examined the neural correlates of these distinct neglect profiles using a statistical voxel-based lesion-symptom mapping method that correlated the anatomical extent of brain damage with the relative severity of deficits along the different profiles in each patient. Our factorial analysis revealed three main factors explaining 82% of the total variance across all neglect tests, which suggested distinct components related to perceptive/visuo-spatial, exploratory/visuo-motor, and allocentric/object-centred aspects of spatial neglect. Our anatomical voxel-based lesion-symptom mapping analysis pointed to specific neural correlates for each of these components, including the right inferior parietal lobule for the perceptive/visuo-spatial component, the right dorsolateral prefrontal cortex for the exploratory/visuo-motor component, and deep temporal lobe regions for the allocentric/object-centred component. By contrast, standard anatomical overlap analysis indicated that subcortical damage to paraventricular white matter tracts was associated with severe neglect encompassing several tests. Taken together, our results provide new support to the view that the clinical manifestations of hemispatial neglect might reflect a combination of distinct components affecting different domains of spatial cognition, and that intra-hemispheric disconnection due to white matter lesions might produce severe neglect by impacting on more than one functional domain.

Biofeedback training for partial weight bearing in patients after total hip arthroplasty.

OBJECTIVE: To evaluate a new biofeedback training method based on visual delivery of information in patients after total hip arthroplasty (THA). DESIGN: Intervention study with prepost design. SETTING: Hospitalized care in a university referral center. PARTICIPANTS: Patients (N=11) (age 56.1+/-9.0 y) shortly after THA. INTERVENTION: A mobile system has been used for biofeedback training with the predefined partial weight bearing (PWB) threshold of 20 kg. After the learning period, 4 retention tests, consisting of 3 successive walking cycles without feedback, were recorded for each patient: (1) acquisition test, (2) early retention test (after 30 min), (3) the day after, and (4) after 2 days. MAIN OUTCOME MEASURE: The pressure error and the maximum pressure force at each step before and after biofeedback training. RESULTS: A significant difference of pressure errors between the beginning and the end of the learning period has been measured (42.5+/-22.5 N vs 3.7+/-11.4N, P<.001). However, there was no difference between the beginning of the learning period and different retention tests (after 30 minutes, after 1 day, after 2 days). In terms of maximal pressure force, there was a difference between the beginning and the end of learning (251 N vs 195 N, P<.05). The retention tests did not show significant differences compared with the baseline values. CONCLUSIONS: THA patients were able to use the defined PWB during a short period of time and shortly after stopping the training; both the pressure errors and the maximal pressure force attended the values before training. These results confirm the difficulties to achieve PWB in patients after THA.

Early neuronal responses in right limbic structures mediate harmony incongruity processing in musical experts.

In western tonal music, musical phrases end with an explicit harmonic consequent which is highly expected. As such expectation is a consequence of musical background, cerebral processing of incongruities of musical grammar might be a function of expertise. We hypothesized that a subtle incongruity of standard closure should evoke a profound and rapid reaction in an expert's brain. If such a reaction is due to neuroplasticity as a consequence of musical training, it should be correlated with distinctive activations in sensory, motor and/or cognitive function related brain areas in response to the incongruent closure. Using event related potential (ERP) source imaging, we determined the temporal dynamics of neuronal activity in highly trained pianists and musical laymen in response to syntactic harmonic incongruities in expressive music, which were easily detected by the experts but not by the laymen. Our results revealed that closure incongruity evokes a selective early response in musical experts, characterized by a strong, right lateralized negative ERP component. Statistical source analysis could demonstrate putative contribution to the generation of this component in right temporal-limbic areas, encompassing hippocampal complex and amygdala, and in right insula. Its early onset (approximately 200 ms) preceded responses in frontal areas that may reflect more conscious processing. These results go beyond previous work demonstrating that musical training can change activity of sensory and motor areas during musical or audio-motor tasks, and suggest that functional plasticity in right medial-temporal structures and insula also modulates processing of subtle harmonic incongruities.

Dissociable components of spatial neglect associated with frontal and parietal lesions.

Spatial neglect is a complex neuropsychological disorder, in which patients fail to detect and respond to contralesional stimuli. Recent studies suggest that these symptoms may reflect a combination of different component deficits, associated with different lesion substrates. Thus, damage to right lateral prefrontal and inferior parietal regions produce different degrees of left neglect on cancellation and line bisection tasks, respectively. Here we tested for dissociable behaviors across two tasks designed to assess distinct cognitive processes possibly mediating such components, in 14 patients with right focal lesion in either the frontal or parietal lobe. In the "distractor filtering" task, patients had to respond to a visual target presented centrally, with or without a lateralized distractor. Only frontal-lesioned patients showed a marked slowing of reaction times when a central target appeared with a simultaneous right distractor (compared to center and left distractor). In the "spatial coding" task, patients had to detect a target among successive visual stimuli presented horizontally with three sequence conditions (regular/predictive or irregular/non-predictive). Only parietal-lesioned patients were unable to benefit from the predictability of the target position, with similar reaction times across all sequence conditions. By contrast, frontal patients showed faster reaction times on trials with a regular succession of stimuli (compared to random order). Taken together, these results suggest that frontal damage may contribute to left inattention by disrupting top-down control and resistance to distractors on the ipsilesional side, whereas parietal damage may disrupt the maintenance of stable locations in space across gaze shifts or time. This further supports the notion that left neglect may arise as a combined breakdown or impaired connectivity between frontal and parietal mechanisms involved (respectively) in the selective control and memory storage components of spatial attention.

Clinical assessment of motor function: a processes oriented instrument based on a speed-accuracy trade-off paradigm.

In this study, we developed a digitizing tablet-based instrument for the clinical assessment of human voluntary movements targeting motor processes of planning, programming and execution. The tool was used to investigate an adaptation of Fitts' reciprocal tapping task [10], comprising four conditions, each of them modulated by three indices of difficulty related to the amplitude of movement required. Temporal, spatial and sequential constraints underlying the various conditions allowed the intricate motor processes to be dissociated. Data obtained from a group of elderly healthy subjects (N=50) were in agreement with the literature on motor control, in the temporal and spatial domains. Speed constraints generated gains in the temporal domain and costs in the spatial one, while spatial constraints generated gain in the spatial domain and costs in the temporal one; finally, sequential constraints revealed the integrative nature of the cognitive operations involved in motor production. This versatile instrument proved capable of providing quantitative, accurate and sensitive measures of the various processes sustaining voluntary movement in healthy subjects. Altogether, analyses performed in this study generated a theoretical framework and reference data which could be used in the future for the clinical assessment of patients with various movement disorders, in particular Parkinson's disease.

Dynamic modulation of visual detection by auditory cues in spatial neglect.

One of the most constant findings of studies about selective attention is that detection of visual stimuli is enhanced when a visual cue is presented at the position of the upcoming target. In healthy participants, comparable benefits were reported when the cue was presented in a different modality than the target. The aim of this study was to examine spatial and temporal dynamics of visual attention following auditory cues in patients with spatial neglect. Twelve healthy subjects and five patients with left-sided neglect were asked to react to a small vertical line presented randomly at one of four positions. The target appeared 150 or 1000 ms after an auditory cue that was either static (continuous 380 Hz tone presented to the left or right ear) or dynamic (380 Hz tone moving from the left to the right ear or vice versa). The reaction time pattern of healthy participants was unaffected by the different tones. In contrast, reaction times of neglect patients were significantly faster to left targets following a dynamic tone moving from right to left in comparison to a tone moving from left to right. Interestingly, static unilateral tones modulated visual attention of neglect patients to a lesser degree than dynamic tones. The modulation of visual attention by dynamic auditory cues was of short duration and disappeared after 1000 ms. These results demonstrate a fast automatic shift of spatial attention in the direction of a moving tone, suggesting strong dynamic links between visual and auditory attention in patients with a severe spatial deficit.

Programming effectors and coordination in bimanual in-phase mirror finger movements.

We investigated cerebral activation during programming of in-phase symmetric finger movements in a precued response task. Partial precues provided advance information about either mirror effectors or in-phase coordination of bimanual movements, while full precue specified both response parameters and neutral precue no movement information. Effects of precueing were assessed on reaction time (RT), contingent negative variation (CNV), and alpha and beta event-related desynchronization (ERD). Information on coordination mode induced less efficient preparation than information on effectors, as revealed by longer RT, but paradoxically the CNV was found of larger amplitude for in-phase than for mirror precue. Full and in-phase precues were associated to largest cerebral activation, as reflected by CNV amplitude as well as beta ERD. It is suggested that with in-phase precueing, abstract programming of coordination and concrete preparation of possible effectors overlap, engaging more cerebral resources than when symmetric effectors are pre-specified. Alpha ERD underwent regional modulations dependent on the type of preparation, pointing out the role of the right parietal region in visuomotor transformation with full movement programming, and the preferential implication of the dominant hemisphere and medial brain regions in synchronization of both hand movements. Beta ERD topographical distribution suggested an increased implication of bilateral and medial motor regions in anticipation to the response signal with incomplete movement preparation.

Event-related potentials and time course of the "other-race" face classification advantage.

Other-race faces are less accurately recognized than same race faces but classified faster by race. Using event-related potentials (ERPs), we captured the brain temporal dynamics of face classification by race processing performed by 12 Caucasian participants. As expected, participants were faster to classify by race Asian than Caucasian faces. ERPs results identified the occurrence of the other-race face classification advantage at around 240 ms, in a stage related to the processing of visual information at the semantic level. The elaboration of individual face structural representation, reflected in the N170 face-sensitive component, was insufficient to achieve this process. Altogether, these findings suggest that the lesser experience of other-race faces engender fewer semantic representations, which in turn accelerate their speed of processing.

Musical training intensity yields opposite effects on grey matter density in cognitive versus sensorimotor networks.

Using optimized voxel-based morphometry, we performed grey matter density analyses on 59 age-, sex- and intelligence-matched young adults with three distinct, progressive levels of musical training intensity or expertise. Structural brain adaptations in musicians have been repeatedly demonstrated in areas involved in auditory perception and motor skills. However, musical activities are not confined to auditory perception and motor performance, but are entangled with higher-order cognitive processes. In consequence, neuronal systems involved in such higher-order processing may also be shaped by experience-driven plasticity. We modelled expertise as a three-level regressor to study possible linear relationships of expertise with grey matter density. The key finding of this study resides in a functional dissimilarity between areas exhibiting increase versus decrease of grey matter as a function of musical expertise. Grey matter density increased with expertise in areas known for their involvement in higher-order cognitive processing: right fusiform gyrus (visual pattern recognition), right mid orbital gyrus (tonal sensitivity), left inferior frontal gyrus (syntactic processing, executive function, working memory), left intraparietal sulcus (visuo-motor coordination) and bilateral posterior cerebellar Crus II (executive function, working memory) and in auditory processing: left Heschl's gyrus. Conversely, grey matter density decreased with expertise in bilateral perirolandic and striatal areas that are related to sensorimotor function, possibly reflecting high automation of motor skills. Moreover, a multiple regression analysis evidenced that grey matter density in the right mid orbital area and the inferior frontal gyrus predicted accuracy in detecting fine-grained incongruities in tonal music.

Stability-dependent behavioural and electro-cortical reorganizations during intentional switching between bimanual tapping modes.

This study investigated behavioural and electro-cortical reorganizations accompanying intentional switching between two distinct bimanual coordination tapping modes (In-phase and Anti-phase) that differ in stability when produced at the same movement rate. We expected that switching to a less stable tapping mode (In-to-Anti switching) would lead to larger behavioural perturbations and require supplementary neural resources than switching to a more stable tapping mode (Anti-to-In switching). Behavioural results confirmed that the In-to-Anti switching lasted longer than the Anti-to-In switching. A general increase in attention-related neural activity was found at the moment of switching for both conditions. Additionally, two condition-dependent EEG reorganizations were observed. First, a specific increase in cortico-cortical coherence appeared exclusively during the In-to-Anti switching. This result may reflect a strengthening in inter-regional communication in order to engage in the subsequent, less stable, tapping mode. Second, a decrease in motor-related neural activity (increased beta spectral power) was found for the Anti-to-In switching only. The latter effect may reflect the interruption of the previous, less stable, tapping mode. Given that previous results on spontaneous Anti-to-In switching revealing an inverse pattern of EEG reorganization (decreased beta spectral power), present findings give new insight on the stability-dependent neural correlates of intentional motor switching.

Electro-cortical correlates of motor inhibition: a comparison between selective and non-selective stop tasks.

In this study, we compared a selective stop task (transition from a bimanual in-phase to a unimanual index fingers' tapping), with a non-selective stop task (stopping a bimanual in-phase tapping at all), and with a switching task (transition from in-phase to anti-phase bimanual tapping). The aim was twofold: 1) to identify the electro-cortical correlates of selective and non-selective inhibition processes and 2) to investigate which type of inhibition - selective or not - is required when switching between two bimanual motor patterns. The results revealed that all tasks led to enhanced activation (alpha power) of the left sensorimotor and posterior regions which seems to reflect an overall effort to stop the preferred bimanual in-phase tendency. Each task implied specific functional connectivity reorganizations (beta coherence) between cerebral motor areas, probably reflecting engagement in a new unimanual or bimanual movement.

Time course of brain activity during change blindness and change awareness: performance is predicted by neural events before change onset.

People often remain "blind" to visual changes occurring during a brief interruption of the display. The processing stages responsible for such failure remain unresolved. We used event-related potentials to determine the time course of brain activity during conscious change detection versus change blindness. Participants saw two successive visual displays, each with two faces, and reported whether one of the faces changed between the first and second displays. Relative to blindness, change detection was associated with a distinct pattern of neural activity at several successive processing stages, including an enhanced occipital P1 response and a sustained frontal activity (CNV-like potential) after the first display, before the change itself. The amplitude of the N170 and P3 responses after the second visual display were also modulated by awareness of the face change. Furthermore, a unique topography of event-related potential activity was observed during correct change and correct no-change reports, but not during blindness, with a recurrent time course in the stimulus sequence and simultaneous sources in the parietal and temporo-occipital cortex. These results indicate that awareness of visual changes may depend on the attentional state subserved by coordinated neural activity in a distributed network, before the onset of the change itself.

The fusiform face area is tuned for curvilinear patterns with more high-contrasted elements in the upper part.

The ability to identify conspecifics from the face is of primary interest for human social behavior. Newborns' visual preference for schematic face-like stimuli has been recently related to a sensitivity for visual patterns with a greater number of elements in the upper compared to the lower part. At the adult level, neuroimaging studies have identified a network of cortical areas devoted to the detection and identification of faces. However, whether and how low-level structural properties of face stimuli contribute to the preferential response to faces in these areas remain to be clarified. Using functional magnetic resonance imaging (fMRI), here we investigated whether the adults' face-sensitive cortical areas show a preference for top-heavy patterns, similarly to newborns' preference. Twelve participants were presented with head-shaped and square patterns with either more elements in the upper or the lower vertical part. In the right fusiform gyrus ('fusiform face area', FFA), an area showing a preference for faces over other visual object categories, there was a larger activation for curvilinear patterns with more high-contrast elements in the upper part, even though these patterns were not perceived as face stimuli. These findings provide direct evidence that the FFA is tuned for geometrical properties fitting best with the structure of faces, a computational mechanism that might drive the automatic detection of faces in the visual world.

Actual and mental motor preparation and execution: a spatiotemporal ERP study.

Studies evaluating the role of the executive motor system in motor imagery came to a general agreement in favour of the activation of the primary motor area (M1) during imagery, although in reduced proportion as compared to motor execution. It is still unclear whether this difference occurs within the preparation period or the execution period of the movement, or both. In the present study, EEG was used to investigate separately the preparation and the execution periods of overt and covert movements in adults. We designed a paradigm that randomly mixed actual and kinaesthetic imagined trials of an externally paced sequence of finger key presses. Sixty channel event-related potentials were recorded to capture the cerebral activations underlying the preparation for motor execution and motor imagery, as well as cerebral activations implied in motor execution and motor imagery. Classical waveform analysis was combined with data-driven spatiotemporal segmentation analysis. In addition, a LAURA source localization algorithm was applied to functionally define brain related motor areas. Our results showed first that the difference between actual and mental motor acts takes place at the late stage of the preparation period and consists of a quantitative modulation of the activity of common structures in M1. Second, they showed that primary motor structures are involved to the same extent in the actual or imagined execution of a motor act. These findings reinforce and refine the functional equivalence hypothesis between actual and imagined motor acts.