Cognitive priming in sung and instrumental music: activation of inferior frontal cortex

Neural correlates of the processing of musical syntax-like structures have been investigated via expectancy violation due to musically unrelated (i.e., unexpected) events in musical contexts. Previous studies reported the implication of inferior frontal cortex in musical structure processing. However - due to the strong musical manipulations - activations might be explained by sensory deviance detection or repetition priming. Our present study investigated neural correlates of musical structure processing with subtle musical violations in a musical priming paradigm. Instrumental and sung sequences ended on related and less-related musical targets. The material controlled sensory priming components, and differences in target processing required listeners' knowledge on musical structures. Participants were scanned with functional Magnetic Resonance Imaging (fMRI) while performing speeded phoneme and timbre identification judgments on the targets. Behavioral results acquired in the scanner replicated the facilitation effect of related over less-related targets. The blood oxygen level-dependent (BOLD) signal linked to target processing revealed activation of right inferior frontal areas (i.e., inferior frontal gyrus, frontal operculum, anterior insula) that was stronger for less-related than for related targets, and this was independent of the material carrying the musical structures. This outcome points to the implication of inferior frontal cortex in the processing of syntactic relations also for musical material and to its role in the processing and integration of sequential information over time. In addition to inferior frontal activation, increased activation was observed in orbital gyrus, temporal areas (anterior superior temporal gyrus, posterior superior temporal gyrus and sulcus, posterior middle temporal gyrus) and supramarginal gyrus.     

Differentiating maturational and training influences on fMRI activation during music processing

Two major influences on how the brain processes music are maturational development and active musical training. Previous functional neuroimaging studies investigating music processing have typically focused on either categorical differences between "musicians versus nonmusicians" or "children versus adults." In the present study, we explored a cross-sectional data set (n=84) using multiple linear regression to isolate the performance-independent effects of age (5 to 33 years) and cumulative duration of musical training (0 to 21,000 practice hours) on fMRI activation similarities and differences between melodic discrimination (MD) and rhythmic discrimination (RD). Age-related effects common to MD and RD were present in three left hemisphere regions: temporofrontal junction, ventral premotor cortex, and the inferior part of the intraparietal sulcus, regions involved in active attending to auditory rhythms, sensorimotor integration, and working memory transformations of pitch and rhythmic patterns. By contrast, training-related effects common to MD and RD were localized to the posterior portion of the left superior temporal gyrus/planum temporale, an area implicated in spectrotemporal pattern matching and auditory-motor coordinate transformations. A single cluster in right superior temporal gyrus showed significantly greater activation during MD than RD. This is the first fMRI which has distinguished maturational from training effects during music processing.     

Dissociation between melodic and rhythmic processing during piano performance from musical scores

When performing or perceiving music, we experience the melodic (spatial) and rhythmic aspects as a unified whole. Moreover, the motor program theory stipulates that the relative timing and the serial order of the movement are invariant features of a motor program. Still, clinical and psychophysical observations suggest independent processing of these two aspects, in both production and perception. Here, we used functional magnetic resonance imaging to dissociate between brain areas processing the melodic and the rhythmic aspects during piano playing from musical scores. This behavior requires that the pianist decodes two types of information from the score in order to produce the desired piece of music. The spatial location of a note head determines which piano key to strike, and the various features of the note, such as the stem and flags determine the timing of each key stroke. We found that the medial occipital lobe, the superior temporal lobe, the rostral cingulate cortex, the putamen and the cerebellum process the melodic information, whereas the lateral occipital and the inferior temporal cortex, the left supramarginal gyrus, the left inferior and ventral frontal gyri, the caudate nucleus, and the cerebellum process the rhythmic information. Thus, we suggest a dissociate involvement of the dorsal visual stream in the spatial pitch processing and the ventral visual stream in temporal movement preparation. We propose that this dissociate organization may be important for fast learning and flexibility in motor control.     

Neural correlates of timbre change in harmonic sounds

Timbre is a major structuring force in music and one of the most important and ecologically relevant features of auditory events. We used sound stimuli selected on the basis of previous psychophysiological studies to investigate the neural correlates of timbre perception. Our results indicate that both the left and right hemispheres are involved in timbre processing, challenging the conventional notion that the elementary attributes of musical perception are predominantly lateralized to the right hemisphere. Significant timbre-related brain activation was found in well-defined regions of posterior Heschl's gyrus and superior temporal sulcus, extending into the circular insular sulcus. Although the extent of activation was not significantly different between left and right hemispheres, temporal lobe activations were significantly posterior in the left, compared to the right, hemisphere, suggesting a functional asymmetry in their respective contributions to timbre processing. The implications of our findings for music processing in particular and auditory processing in general are discussed.     

Sentence complexity and input modality effects in sentence comprehension: an fMRI study

Cortical regions engaged by sentence processing were mapped using functional MRI. The influence of input modality (spoken word vs. print input) and parsing difficulty (sentences containing subject-relative vs. object-relative clauses) was assessed. Auditory presentation was associated with pronounced activity at primary auditory cortex and across the superior temporal gyrus bilaterally. Printed sentences by contrast evoked major activity at several posterior sites in the left hemisphere, including the angular gyrus, supramarginal gyrus, and the fusiform gyrus in the occipitotemporal region. In addition, modality-independent regions were isolated, with greatest overlap seen in the inferior frontal gyrus (IFG). With respect to sentence complexity, object-relative sentences evoked heightened responses in comparison to subject-relative sentences at several left hemisphere sites, including IFG, the middle/superior temporal gyrus, and the angular gyrus. These sites showing modulation of activity as a function of sentence type, independent of input mode, arguably form the core of a cortical system essential to sentence parsing.     

The Effect of Task Relevance on the Cortical Response to Changes in Visual and Auditory Stimuli: An Event-Related fMRI Study

Attention is, in part, a mechanism for identifying features of the sensory environment of potential relevance to behavior. The network of brain areas sensitive to the behavioral relevance of multimodal sensory events has not been fully characterized. We used event-related fMRI to identify brain regions responsive to changes in both visual and auditory stimuli when those changes were either behaviorally relevant or behaviorally irrelevant. A widespread network of "context-dependent" activations responded to both task-irrelevant and task-relevant events but responded more strongly to task-relevant events. The most extensive activations in this network were located in right and left temporoparietal junction (TPJ), with smaller activations in left precuneus, left anterior insula, left anterior cingulate cortex, and right thalamus. Another network of "context-independent" activations responded similarly to all events, regardless of task relevance. This network featured a large activation encompassing left supplementary and cingulate motor areas (SMA/CMA) as well as right IFG, right/left precuneus, and right anterior insula, with smaller activations in right/left inferior temporal gyrus and left posterior cingulate cortex. Distinct context-dependent and context-independent subregions of activation were also found within the left and right TPJ, left anterior insula, and left SMA/CMA. In the right TPJ, a subregion in the supramarginal gyrus showed sensitivity to the behavioral context (i.e., relevance) of stimulus changes, while two subregions in the superior temporal gyrus did not. The results indicate a role for the TPJ in detecting behaviorally relevant events in the sensory environment. The TPJ may serve to identify salient events in the sensory environment both within and independent of the current behavioral context.     

Dissociable systems of working memory for rhythm and melody

Specialized neural systems are engaged by the rhythmic and melodic components of music. Here, we used PET to measure regional cerebral blood flow (rCBF) in a working memory task for sequences of rhythms and melodies, which were presented in separate blocks. Healthy subjects, without musical training, judged whether a target rhythm or melody was identical to a series of subsequently presented rhythms or melodies. When contrasted with passive listening to rhythms, working memory for rhythm activated the cerebellar hemispheres and vermis, right anterior insular cortex, and left anterior cingulate gyrus. These areas were not activated in a contrast between passive listening to rhythms and a non-auditory control, indicating their role in the temporal processing that was specific to working memory for rhythm. The contrast between working memory for melody and passive listening to melodies activated mainly a right-hemisphere network of frontal, parietal, and temporal cortices: areas involved in pitch processing and auditory working memory. Overall, these results demonstrate that rhythm and melody have unique neural signatures not only in the early stages of auditory processing, but also at the higher cognitive level of working memory.     

Semantic and episodic memory of music are subserved by distinct neural networks

Numerous functional imaging studies have shown that retrieval from semantic and episodic memory is subserved by distinct neural networks. However, these results were essentially obtained with verbal and visuospatial material. The aim of this work was to determine the neural substrates underlying the semantic and episodic components of music using familiar and nonfamiliar melodic tunes. To study musical semantic memory, we designed a task in which the instruction was to judge whether or not the musical extract was felt as "familiar." To study musical episodic memory, we constructed two delayed recognition tasks, one containing only familiar and the other only nonfamiliar items. For each recognition task, half of the extracts (targets) were presented in the prior semantic task. The episodic and semantic tasks were to be contrasted by a comparison to two perceptive control tasks and to one another. Cerebral blood flow was assessed by means of the oxygen-15-labeled water injection method, using high-resolution PET. Distinct patterns of activations were found. First, regarding the episodic memory condition, bilateral activations of the middle and superior frontal gyri and precuneus (more prominent on the right side) were observed. Second, the semantic memory condition disclosed extensive activations in the medial and orbital frontal cortex bilaterally, the left angular gyrus, and predominantly the left anterior part of the middle temporal gyri. The findings from this study are discussed in light of the available neuropsychological data obtained in brain-damaged subjects and functional neuroimaging studies.     

Dynamics of brain activity in motor and frontal cortical areas during music listening: a magnetoencephalographic study

There are formidable problems in studying how 'real' music engages the brain over wide ranges of temporal scales extending from milliseconds to a lifetime. In this work, we recorded the magnetoencephalographic signal while subjects listened to music as it unfolded over long periods of time (seconds), and we developed and applied methods to correlate the time course of the regional brain activations with the dynamic aspects of the musical sound. We showed that frontal areas generally respond with slow time constants to the music, reflecting their more integrative mode; motor-related areas showed transient-mode responses to fine temporal scale structures of the sound. The study combined novel analysis techniques designed to capture and quantify fine temporal sequencing from the authentic musical piece (characterized by a clearly defined rhythm and melodic structure) with the extraction of relevant features from the dynamics of the regional brain activations. The results demonstrated that activity in motor-related structures, specifically in lateral premotor areas, supplementary motor areas, and somatomotor areas, correlated with measures of rhythmicity derived from the music. These correlations showed distinct laterality depending on how the musical performance deviated from the strict tempo of the music score, that is, depending on the musical expression.     

Incidental effects of emotional valence in single word processing: an fMRI study

The present study aimed at identifying the neural responses associated with the incidental processing of the emotional valence of single words using event-related functional magnetic resonance imaging (fMRI). Twenty right-handed participants performed a visual lexical decision task, discriminating between nouns and orthographically and phonologically legal nonwords. Positive, neutral and negative word categories were matched for frequency, number and frequency of orthographic neighbors, number of letters and imageability. Response times and accuracy data differed significantly between positive and neutral, and positive and negative words respectively, thus, replicating the findings of a pilot study. Words showed distributed, mainly left hemisphere activations, indicating involvement of a neural network responsible for semantic word knowledge. The neuroimaging data further revealed areas in left orbitofrontal gyrus and bilateral inferior frontal gyrus with greater activation to emotional than to neutral words. These brain regions are known to be involved in processing semantic and emotional information. Furthermore, distinct activations associated with positive words were observed in bilateral middle temporal and superior frontal gyrus, known to support semantic retrieval, and a distributed network, namely anterior and posterior cingulate gyrus, lingual gyrus and hippocampus when comparing positive and negative words. The latter areas were previously associated with explicit and not incidental processing of the emotional meaning of words and emotional memory retrieval. Thus, the results are discussed in relation to models of processing semantic and episodic emotional information.     

Self, mother and abstract other: an fMRI study of reflective social processing

Using fMRI, we studied the neural correlates of self-referential processing by comparing BOLD signal changes during self and mother conditions of a self-reference effect (SRE) task. Conjunction analysis of these two conditions showed several common areas of significant activation, including the medial aspects of the superior frontal gyri, left inferior frontal gyrus, bilateral temporal poles, left superior temporal sulcus and left precuneus. The locations of the 7 strongest peak activations for the self condition and for the mother condition were compared on a subject-by-subject basis in native space. Of the 119 pairs of peaks explored, 87% were located within 2 voxels of each other, demonstrating the commonality of the brain regions subserving both self- and mother-referential processing within an individual subject. In group analyses of the self-referential vs. mother-referential contrast, small differences in activation strength were observed in the left superior frontal sulcus, right cingulate gyrus and the left fusiform gyrus. Greater activation for mother than for self was observed in bilateral temporal lobes. Subjects also performed a social attribution task (SAT) in which they inferred mental states about interacting geometric shapes. Activations from this visual theory of mind task were compared with the activations demonstrated during self-referential processing. Striking similarities were found, including overlapping activations in bilateral medial prefrontal cortices, left inferior frontal gyrus and precuneus. These data suggest that reflective analysis of self, mother and abstract others relies predominantly on the same neural architecture.     

A common language network for comprehension and production: a contribution to the definition of language epicenters with PET

In this paper, we report on a PET activation study designed to assess whether functional neuroimaging would help to uncover essential language areas in normal volunteers and to provide a more accurate definition of their localization. Regional cerebral blood flow was repeatedly monitored in eight right-handed male volunteers, while performing a language comprehension task (listening to factual stories) and a language production task (covert generation of verbs semantically related to heard nouns), using silent resting as a control condition. The conjunction analysis, conducted with SPM, was used to uncover the network of activations common to both task that included three left hemisphere areas, namely (1) the pars opercularis and triangularis of the inferior frontal gyrus, (2) the posterior part of the superior temporal cortex centered around the superior temporal sulcus, extending to the planum temporale posterior part but sparing the supramarginalis and angular gyri, and (3) the most anterior part of the left inferior temporal gyrus at the junction with the anterior fusiform gyrus. The inferior and lateral parts of the right cerebellar cortex were also included in the conjunction network. Each of the three cortical areas, when they are site of lesion or electrical stimulation, elicit impairment in both language comprehension and production and can thus be considered as essential to language. Accordingly, the present results provide conservative anatomofunctional definitions of the Broca, Wernicke, and basal language areas. Interestingly, contralateral homologues of Broca's and Wernicke's areas also lighted up in the conjunction analysis that could be related to the interindividual variability of hemispheric language dominance.     

Temporal and topographical characterization of language cortices using intraoperative optical intrinsic signals

We used intraoperative optical imaging of intrinsic signals (iOIS) and electrocortical stimulation mapping (ESM) to compare functionally active brain regions in 10 awake patients undergoing neurosurgical resection. Patients performed two to four tasks, including visual and auditory naming, word discrimination, and/or orofacial movements. All iOIS maps included areas identified by ESM mapping. However, iOIS also revealed topographical specificity dependent on language task. In Broca's area, naming paradigms activated both anterior and posterior inferior frontal gyrus (IFG), while the word discrimination paradigm activated only posterior IFG. In Wernicke's area, object naming produced activations localizing over the inferior and anterior/posterior regions, while the word discrimination task activated superior and anterior cortices. These results may suggest more posterior phonological activation and more anterior semantic activations in Broca's area, and more anterior/superior phonological activation and more posterior/inferior semantic activations in Wernicke's area. Although similar response onset was observed in Broca's and Wernicke's areas, temporal differences were revealed during block paradigm (20-s) activations. In Broca's area, block paradigms yielded a boxcar temporal activation profile (in all tasks) that resembled response profiles observed in motor cortex (with orofacial movements). In contrast, activations in Wernicke's area responded with a more dynamic profile (including early and late peaks) which varied with paradigm performance. Wernicke's area profiles were very similar to response profiles observed in sensory and visual cortex. The differing temporal patterns may therefore reflect unique processing performed by receptive (Wernicke's) and productive (Broca's) language centers. This study is consistent with task-specific semantic and phonologic regions within Broca's and Wernicke's areas and also is the first report of response profile differences dependent on cortical region and language task.     

Neuronal correlates of theory of mind and empathy: a functional magnetic resonance imaging study in a nonverbal task

Theory of Mind (ToM), the ability to attribute mental states to others, and empathy, the ability to infer emotional experiences, are important processes in social cognition. Brain imaging studies in healthy subjects have described a brain system involving medial prefrontal cortex, superior temporal sulcus and temporal pole in ToM processing. Studies investigating networks associated with empathic responding also suggest involvement of temporal and frontal lobe regions. In this fMRI study, we used a cartoon task derived from Sarfati et al. (1997) [Sarfati, Y., Hardy-Bayle, M.C., Besche, C., Widlocher, D. 1997. Attribution of intentions to others in people with schizophrenia: a non-verbal exploration with comic strips. Schizophrenia Research 25, 199-209.]with both ToM and empathy stimuli in order to allow comparison of brain activations in these two processes. Results of 13 right-handed, healthy, male volunteers were included. Functional images were acquired using a 1.5 T Phillips Gyroscan. Our results confirmed that ToM and empathy stimuli are associated with overlapping but distinct neuronal networks. Common areas of activation included the medial prefrontal cortex, temporoparietal junction and temporal poles. Compared to the empathy condition, ToM stimuli revealed increased activations in lateral orbitofrontal cortex, middle frontal gyrus, cuneus and superior temporal gyrus. Empathy, on the other hand, was associated with enhanced activations of paracingulate, anterior and posterior cingulate and amygdala. We therefore suggest that ToM and empathy both rely on networks associated with making inferences about mental states of others. However, empathic responding requires the additional recruitment of networks involved in emotional processing. These results have implications for our understanding of disorders characterized by impairments of social cognition, such as autism and psychopathy.     

fMRI investigation of unexpected somatosensory feedback perturbation during speech

Somatosensory feedback plays a critical role in the coordination of articulator movements for speech production. In response to unexpected resistance to lip or jaw movements during speech, fluent speakers can use the difference between the somatosensory expectations of a speech sound and the actual somatosensory feedback to adjust the trajectories of functionally relevant but unimpeded articulators. In an effort to investigate the neural substrates underlying the somatosensory feedback control of speech, we used an event-related sparse sampling functional magnetic resonance imaging paradigm and a novel pneumatic device that unpredictably blocked subjects' jaw movements. In comparison to speech, perturbed speech, in which jaw perturbation prompted the generation of compensatory speech motor commands, demonstrated increased effects in bilateral ventral motor cortex, right-lateralized anterior supramarginal gyrus, inferior frontal gyrus pars triangularis and ventral premotor cortex, and bilateral inferior posterior cerebellum (lobule VIII). Structural equation modeling revealed a significant increased influence from left anterior supramarginal gyrus to right anterior supramarginal gyrus and from left anterior supramarginal gyrus to right ventral premotor cortex as well as a significant increased reciprocal influence between right ventral premotor cortex and right ventral motor cortex and right anterior supramarginal gyrus and right inferior frontal gyrus pars triangularis for perturbed speech relative to speech. These results suggest that bilateral anterior supramarginal gyrus, right inferior frontal gyrus pars triangularis, right ventral premotor and motor cortices are functionally coupled and influence speech motor output when somatosensory feedback is unexpectedly perturbed during speech production.     

遗忘型轻度认知障碍患者默认网络改变:基于后扣带回的脑连接研究

目的 探讨遗忘型轻度认知障碍(aMCI)患者后扣带回(PCC)功能连接的变化。 方法 选取性别、年龄和教育程度相匹配的18例aMCI患者(aMCI组)和20名健康老年人(健康老年组)进行GRE-EPI序列的静息状态fMRI检查,以PCC作为种子点,与全脑其他区域进行基于体素的时间序列相关分析。 结果 aMCI组与PCC有显著连接的脑区包括双侧背外侧前额叶(DLPFC)、左侧内侧前额叶(MPFC)及左侧前扣带回(ACC)、双侧顶下小叶(IPL)、双侧颞中回和双侧楔前叶。健康老年组与PCC有显著连接的脑区包括双侧DLPFC、右侧MPFC及ACC、双侧IPL、双侧颞中回、双侧颞下回和双侧楔前叶。健康老年组较aMCI组在右侧额上回、额中回、额下回的盖部及三角部、MPFC及ACC、颞中回及左侧楔前叶连接增强;aMCI组多个脑区功能连接较健康老年组增强,主要包括左侧半球的扣带回中部、顶上小叶(SPL)、IPL、颞中回及右侧IPL。 结论 PCC是aMCI患者脑代谢最先降低的区域,aMCI的PCC与多个功能区的连接破坏是其情节记忆功能减退的生物学基础,而其连接增强提示功能代偿可能。     

Dissociated brain organization for single-digit addition and multiplication

This study compared the patterns of brain activation elicited by single-digit addition and multiplication problems. 20 Chinese undergraduates were asked to verify whether arithmetic equations were true or false during functional magnetic resonance imaging. Results showed that both addition and multiplication were supported by a broad neural system that involved regions within SMA, precentral gyrus, intraparietal sulcus, occipital gyri, superior temporal gyrus, and middle frontal gyrus, as well as some subcortical structures. Nevertheless, addition problems elicited more activation in the intraparietal sulcus and middle occipital gyri at the right hemisphere, and superior occipital gyri at both hemispheres, whereas multiplication had more activation in precentral gyrus, supplementary motor areas, and posterior and anterior superior temporal gyrus at the left hemisphere. This pattern of dissociated activation supports our hypothesis that addition has greater reliance on visuospatial processing and multiplication on verbal processing.     

The effect of appraisal level on processing of emotional prosody in meaningless speech

In visual perception of emotional stimuli, low- and high-level appraisal processes have been found to engage different neural structures. Beyond emotional facial expression, emotional prosody is an important auditory cue for social interaction. Neuroimaging studies have proposed a network for emotional prosody processing that involves a right temporal input region and explicit evaluation in bilateral prefrontal areas. However, the comparison of different appraisal levels has so far relied upon using linguistic instructions during low-level processing, which might confound effects of processing level and linguistic task. In order to circumvent this problem, we examined processing of emotional prosody in meaningless speech during gender labelling (implicit, low-level appraisal) and emotion labelling (explicit, high-level appraisal). While bilateral amygdala, left superior temporal sulcus and right parietal areas showed stronger blood oxygen level-dependent (BOLD) responses during implicit processing, areas with stronger BOLD responses during explicit processing included the left inferior frontal gyrus, bilateral parietal, anterior cingulate and supplemental motor cortex. Emotional versus neutral prosody evoked BOLD responses in right superior temporal gyrus, bilateral anterior cingulate, left inferior frontal gyrus, insula and bilateral putamen. Basal ganglia and right anterior cingulate responses to emotional versus neutral prosody were particularly pronounced during explicit processing. These results are in line with an amygdala-prefrontal-cingulate network controlling different appraisal levels, and suggest a specific role of the left inferior frontal gyrus in explicit evaluation of emotional prosody. In addition to brain areas commonly related to prosody processing, our results suggest specific functions of anterior cingulate and basal ganglia in detecting emotional prosody, particularly when explicit identification is necessary.     

Temporal dynamics of perisylvian activation during language processing in children and adults

The perisylvian region of the human cortex is known to play a major role in language processing. Especially the superior temporal cortex (STC) and the inferior frontal cortex (IFC) have been investigated with respect to their particular involvement in language comprehension. In the present research, the timing of recruitment of these language-related brain areas in both hemispheres was examined as a function of age using functional imaging data of 6-year-old children and adults with a special focus on blood oxygenation level dependent (BOLD) response time courses. The results show that children's activation time courses differ from that of adults. First, children show an overall later peak of BOLD responses. Second, children's IFC responds much later than their STC, while in adults the difference between both regions is less pronounced. Within the STC, both groups show similar regionally U-shaped activation patterns with fastest peaks in voxels at the STC's mid-portion around Heschl's gyrus and longer latencies in anterior and posterior directions, suggesting a coarsely similar information flow in adults and children in the temporal region. Finally, children in contrast to adults, display a temporal primacy of right over left hemispheric activation. The observed overall latency differences between children and adults are in line with the assumption of ongoing maturation in perisylvian brain regions and the connections between them. A functional perspective on BOLD timing argues for a developmental change from higher processing costs in children compared to adults due to slower and less automatic language processes, in particular those located in the IFC. The observed hemispheric differences are discussed in the context of developmental models assuming a high reliance on right-hemisphere-based suprasegmental information processing during language comprehension in childhood.     

Bach speaks: a cortical "language-network" serves the processing of music

The aim of the present study was the investigation of neural correlates of music processing with fMRI. Chord sequences were presented to the participants, infrequently containing unexpected musical events. These events activated the areas of Broca and Wernicke, the superior temporal sulcus, Heschl's gyrus, both planum polare and planum temporale, as well as the anterior superior insular cortices. Some of these brain structures have previously been shown to be involved in music processing, but the cortical network comprising all these structures has up to now been thought to be domain-specific for language processing. To what extent this network might also be activated by the processing of non-linguistic information has remained unknown. The present fMRI-data reveal that the human brain employs this neuronal network also for the processing of musical information, suggesting that the cortical network known to support language processing is less domain-specific than previously believed.