fMRI evidence for cortical modification during learning of Mandarin lexical tone

Functional magnetic resonance imaging was employed before and after six native English speakers completed lexical tone training as part of a program to learn Mandarin as a second language. Language-related areas including Broca's area, Wernicke's area, auditory cortex, and supplementary motor regions were active in all subjects before and after training and did not vary in average location. Across all subjects, improvements in performance were associated with an increase in the spatial extent of activation in left superior temporal gyrus (Brodmann's area 22, putative Wernicke's area), the emergence of activity in adjacent Brodmann's area 42, and the emergence of activity in right inferior frontal gyrus (Brodmann's area 44), a homologue of putative Broca's area. These findings demonstrate a form of enrichment plasticity in which the early cortical effects of learning a tone-based second language involve both expansion of preexisting language-related areas and recruitment of additional cortical regions specialized for functions similar to the new language functions.     

A noninvasive, presurgical expressive and receptive language investigation in a 9-year-old epileptic boy using near-infrared spectroscopy

The intracarotid amobarbital test (IAT) is used for presurgical evaluation of language lateralization. However, this procedure has many limitations, especially in children. As an alternative to IAT, in the case described here, near-infrared spectroscopy (NIRS) was used to investigate expressive and receptive language lateralization as part of the presurgical evaluation of a 9-year-old Yiddish-speaking boy with a probable left temporal epileptic focus. This child could not tolerate IAT or functional MRI. He underwent two NIRS recording sessions while performing expressive and receptive language tasks. Results indicated predominantly left-sided expressive language in Broca's area with ipsilateral cortical recruitment of more posterior regions. Receptive language showed a bilateral cerebral pattern, perhaps as an expression of cerebral plasticity or compensation in this young patient. This case report illustrates that NIRS may contribute to presurgical investigation and could become a noninvasive alternative to IAT and functional MRI in determining speech lateralization in children.     

Activation of language cortex with automatic speech tasks

OBJECTIVE: To identify automatic speech tasks that reliably demonstrate increased regional cerebral blood flow (rCBF) in Broca's and Wernicke's areas of the cortex using PET. BACKGROUND: Localizing language with direct cortical stimulation mapping requires that patients have a stable baseline on tests that engage eloquent cortex. For dysphasic patients or younger children, automatic speech tasks such as counting are often used in lieu of more complex language tests. Evidence from both lesion and neuroimaging studies suggests that these tasks may not adequately engage language cortices. In this study, we examined rCBF during automatic oromotor and speech tasks of varying complexity to identify those eliciting increased CBF in Broca's and Wernicke's areas. METHODS: Eight normal volunteers underwent PET during rest, tongue movements, and three automatic speech tasks: repeating a phoneme sequence, repeating the months of the year, and reciting a memorized prose passage. Images were averaged across subjects and compared across tasks for regional localization and laterality. RESULTS: Whereas all activation tasks produced increased relative CBF in brain regions that correlated with articulation and auditory processing, only the two tasks that used real words (versus phonemes) showed left-lateralized rCBF increases in posterior superior temporal lobe (Wernicke's area), and only the prose repetition task produced left lateralized activity in Broca's area. CONCLUSIONS: Whereas automatic speech typically does not engage language cortex, repeating a memorized prose passage showed unambiguous activation in both Broca's and Wernicke's areas. These results caution against the use of common automatic speech tasks for mapping eloquent cortex and suggest an alternative task for those with poor language abilities or acquired dysphasia who cannot perform standardized language tests reliably.     

Revisiting the role of Broca's area in sentence processing: syntactic integration versus syntactic working memory

Most previous neuroimaging studies of sentence processing have associated Broca's area with syntactic processing; however, the exact nature of the processes subserved by this brain region is yet not well understood. Although some authors suggest that Brodmann area (BA) 44 of the left inferior frontal gyrus (i.e., Broca's area) is relevant for syntactic integration processes, others claim that it is associated with working memory mechanisms relevant for language processing. To dissociate these two possible functions, the present study investigated hemodynamic responses elicited while participants processed German indirect wh-questions. Activation increases were observed in left BA 44 together with superior temporal areas and right hemispheric homologues for sentences with noncanonical word order, in which a verb argument was dislocated from its canonical position over a relatively long distance. In these sentences, syntactic working memory load was assumed to be greatest. In contrast, no activation increase was elicited by object-initial as opposed to subject-initial sentences that did not differ with respect to working memory costs but with respect to syntactic integration costs. These data strongly suggest that Broca's area plays a critical role in syntactic working memory during online sentence comprehension.     

Resting functional connectivity of language networks: characterization and reproducibility

The neural basis of language comprehension and production has been associated with superior temporal (Wernicke's) and inferior frontal (Broca's) cortical areas, respectively. However, recent resting-state functional connectivity (RSFC) and lesion studies have implicated a more extended network in language processing. Using a large RSFC data set from 970 healthy subjects and seed regions in Broca's and Wernicke's, we recapitulate this extended network that includes not only adjoining prefrontal, temporal and parietal regions but also bilateral caudate and left putamen/globus pallidus and subthalamic nucleus. We also show that the language network has predominance of short-range functional connectivity (except posterior Wernicke's area that exhibited predominant long-range connectivity), which is consistent with reliance on local processing. Predominantly, long-range connectivity was left lateralized (except anterior Wernicke's area that exhibited rightward lateralization). The language network also exhibited anti-correlated activity with auditory (only for Wernicke's area) and visual cortices that suggests integrated sequential activity with regions involved with listening or reading words. Assessment of the intra-subject's reproducibility of this network and its characterization in individuals with language dysfunction is required to determine its potential as a biomarker for language disorders.     

The temporal characteristics of functional activation in Broca's area during overt picture naming

The opercular and triangular sections of the inferior frontal gyrus, also known as Broca's area, have been shown to be involved in various language tasks. In the current study we investigated both the functional role, as well as the precise temporal involvement of Broca's area during picture naming. We applied online event-related transcranial magnetic stimulation (TMS) to Broca's area at five different time points after picture presentation, aiming to cover the complete language production process. Applying real TMS at 300 msec after picture presentation led to an increase in picture naming latency, whereas sham stimulation and real stimulation at earlier and later time windows did not result in any changes in reaction time (RT). Our methodological approach enabled us to get insight into the temporal characteristics of the involvement of this brain area during picture naming. Making use of this information and directly relating it to psycholinguistic models, we conclude that Broca's area may be involved in the process of syllabification during overt speech production.     

Evidence of cortical reorganization of language networks after stroke with subacute Broca's aphasia: a blood oxygenation level dependent-functional magnetic resonance imaging study

Aphasia is an acquired language disorder that is a common consequence of stroke.The pathogenesis of the disease is not fully understood,and as a result,current treatment options are not satisfactory.Here,we used blood oxygenation level-dependent functional magnetic resonance imaging to evaluate the activation of bilateral cortices in patients with Broca's aphasia 1 to 3 months after stroke.Our results showed that language expression was associated with multiple brain regions in which the right hemisphere participated in the generation of language.The activation areas in the left hemisphere of aphasia patients were significantly smaller compared with those in healthy adults.The activation frequency,volumes,and intensity in the regions related to language,such as the left inferior frontal gyrus(Broca's area),the left superior temporal gyrus,and the right inferior frontal gyrus(the mirror region of Broca's area),were lower in patients compared with healthy adults.In contrast,activation in the right superior temporal gyrus,the bilateral superior parietal lobule,and the left inferior temporal gyrus was stronger in patients compared with healthy controls.These results suggest that the right inferior frontal gyrus plays a role in the recovery of language function in the subacute stage of stroke-related aphasia by increasing the engagement of related brain areas.     

Second-language instinct and instruction effects: nature and nurture in second-language acquisition

Adults seem to have greater difficulties than children in acquiring a second language (L2) because of the alleged "window of opportunity" around puberty. Postpuberty Japanese participants learned a new English rule with simplex sentences during one month of instruction, and then they were tested on "uninstructed complex sentences" as well as "instructed simplex sentences." The behavioral data show that they can acquire more knowledge than is instructed, suggesting the interweaving of nature (universal principles of grammar, UG) and nurture (instruction) in L2 acquisition. The comparison in the "uninstructed complex sentences" between post-instruction and pre-instruction using functional magnetic resonance imaging reveals a significant activation in Broca's area. Thus, this study provides new insight into Broca's area, where nature and nurture cooperate to produce L2 learners' rich linguistic knowledge. It also shows neural plasticity of adult L2 acquisition, arguing against a critical period hypothesis, at least in the domain of UG.     

Neural correlates of syntactic transformations

Many agrammatic aphasics have a specific syntactic comprehension deficit involving processing syntactic transformations. It has been proposed that this deficit is due to a dysfunction of Broca's area, an area that is thought to be critical for comprehension of complex transformed sentences. The goal of this study was to investigate the role of Broca's area in processing canonical and non-canonical sentences in healthy subjects. The sentences were presented auditorily and were controlled for task difficulty. Subjects were asked to judge the grammaticality of the sentences while their brain activity was monitored using event-related functional magnetic resonance imaging. Processing both kinds of sentences resulted in activation of language-related brain regions. Comparison of non-canonical and canonical sentences showed greater activation in bilateral temporal regions; a greater activation of Broca's area in processing antecedent-gap relations was not found. Moreover, the posterior part of Broca's area was conjointly activated by both sentence conditions. Broca's area is thus involved in general syntactic processing as required by grammaticality judgments and does not seem to have a specific role in processing syntactic transformations.     

Phonological processing during language production: fMRI evidence for a shared production-comprehension network

Studies of phonological processes during language comprehension consistently report activation of the superior portion of Broca's area. In the domain of language production, however, there is no unequivocal evidence for the contribution of Broca's area to phonological processing. The present event-related fMRI study investigated the existence of a common neural network for phonological decisions in comprehension and production by using production tasks most comparable to those previously used in comprehension. Subjects performed two decision tasks on the initial phoneme of German picture names (/b/ or not? Vowel or not?). A semantic decision task served as a baseline for both phonological tasks. The contrasts between each phonological task and the semantic task were calculated, and a conjunction analysis was performed. There was significant activation in the superior portion of Broca's area (Brodmann's area (BA) 44) in the conjunction analysis, also present in each single contrast. In addition, further left frontal (BA 45/46) and temporal (posterior superior temporal gyrus) areas known to support phonological processing in both production and comprehension were activated. The results suggest the existence of a shared fronto-temporal neural network engaged in the processing of phonological information in both perception and production.     

Relative shift in activity from medial to lateral frontal cortex during internally versus externally guided word generation

Goldberg (1985) hypothesized that as language output changes from internally to externally guided production, activity shifts from supplementary motor area (SMA) to lateral premotor areas, including Broca's area. To test this hypothesis, 15 right-handed native English speakers performed three word generation tasks varying in the amount of internal guidance and a repetition task during functional magnetic resonance imaging (fMRI). Volumes of significant activity for each task versus a resting state were derived using voxel-by-voxel repeated-measures t tests (p <.001) across subjects. Changes in the size of activity volumes for left medial frontal regions (SMA and pre-SMA/BA 32) versus left lateral frontal regions (Broca's area, inferior frontal sulcus) were assessed as internal guidance of word generation decreased and external guidance increased. Comparing SMA to Broca's area, Goldberg's hypothesis was not verified. However, pre-SMA/BA 32 activity volumes decreased significantly and inferior frontal sulcus activity volumes increased significantly as word generation tasks moved from internally to externally guided.     

Sentence translation in proficient bilinguals: a direct electrostimulation brain mapping

Direct cortical electrostimulation was used to study cortical areas hypothetically involved in translation in bilinguals during brain tumour resections, with a view to sparing these functional areas. A series of seven proficient bilingual patients was studied: two left-handed and five right-handed individuals with no pre-existing language deficit. Hemispheric cortex (on the side contralateral to the patient's hand-dominance) was directly stimulated whilst the patient performed naming and reading tasks in both languages and a translation task (of a written text from their second 'learned' language to their first or 'native' language). Of the 147 different cortical sites studied, 26 'language functional sites' were detected, where electrostimulation affected reading and/or naming in the patient's native and/or second learned language. Of these, 8 sites (in 4 patients) were "task-specific" and "language-specific" i.e., affecting only naming or reading in only one of the patient's languages. Of the 26 "language sites", only 3 produced any interferences in translation. All of these were located in frontal regions. Electrostimulation at these sites caused the patient to stop translating abruptly, but no language switching or other translation-related phenomenon was observed. No site was found that was involved only in translation and not other language tasks. Overall, in contrast to other language tasks, cortical structures of the convexity were rarely involved in translation. We suggest that translation interference could be more readily detected by subcortical stimulations. This spatial dissociation within the brain of translation function versus other language functions could explain the cases of dissociated language impairments observed in some bilingual patients with brain lesions. On a practical level, because the cortical sites found by translation tasks are few and related with other cortical language sites, we think that translation tasks provide little additional helpful information for cortical brain mapping in bilingual neurosurgical patients.     

Detection of the brain response during a cognitive task using perfusion-based event-related functional MRI

Event-related (ER) fMRI has evoked great interest due to the ability to depict the dynamic features of human brain function during various cognitive tasks. Thus far, all cognitive ER-fMRI studies have been based on blood oxygenation level-dependent (BOLD) contrast techniques. Compared with BOLD-based fMRI techniques, perfusion-based fMRI is able to localize the region of neuronal activity more accurately. This report demonstrates, for the first time, the detection of the brain response to a cognitive task using high temporal resolution perfusion-based ER-fMRI. An English verb generation task was used in this study. Results show that perfusion-based ER-fMRI accurately depicts the activation in Broca's area. Average changes in regional relative cerebral blood flow reached a maximum value of 30.7% at approximately 6.5 s after the start of stimulation and returned to 10% of the maximum value at approximately 12.8 s. Our results show that perfusion-based ER-fMRI is a useful tool for cognitive neuroscience studies, providing comparable temporal resolution and better localization of brain function than BOLD ER-fMRI.     

Auditory stimulus repetition effects on cortical hemoglobin oxygenation: a near-infrared spectroscopy investigation

The cortical response to repeated sensory stimuli plateaus (or declines) as repetition frequencies increase beyond 2-8 Hz. This study examined the underlying changes in cortical oxygenated and deoxygenated hemoglobin associated with this phenomenon using near-infrared spectroscopy. The optical signal was measured from 11 healthy volunteers listening to noise-burst trains presented at 2, 10, and 35 Hz. In a bilateral region consistent with the posterior superior temporal gyrus there was an inverse relationship between deoxyhemoglobin concentration change and stimulus frequency: greatest at 2 Hz, intermediate at 10 Hz, and smallest at 35 Hz. These findings provide preliminary support for a relationship between the perceptual characteristics of auditory stimuli and modulation of cortical oxygenation as measured via an emerging neuromonitoring technique.     

The cortical organization of audio-visual sentence comprehension: an fMRI study at 4 Tesla

Neuroimaging studies of written and spoken sentence processing report greater left hemisphere than right hemisphere activation. However, a large majority of our experience with language is face-to-face interaction, which is much richer in information. The current study examines the neural organization of audio-visual (AV) sentence processing using functional magnetic resonance imaging (fMRI) at 4 Tesla. Participants viewed the face and upper body of a speaker via a video screen while listening to her produce, in alternating blocks, English sentences and sentences composed of pronounceable non-words. Audio-visual sentence processing was associated with activation in the left hemisphere in Broca's area, dorsolateral prefrontal cortex, the superior precentral sulcus, anterior and middle portions of the lateral sulcus, middle superior portions of the temporal sulcus, supramarginal gyrus and angular gyrus. Further, AV sentence processing elicited activation in the right anterior and middle lateral sulcus. Between-hemisphere analyses revealed a left hemisphere dominant pattern of activation. The findings support the hypothesis that the left hemisphere may be biased to process language independently of the modality through which it is perceived. These results are discussed in the context of previous neuroimaging results using American Sign Language (ASL).     

Quantitative Comparison of Language Deficits Produced by Extraoperative Electrical Stimulation of Broca's, Wernicke's, and Basal Temporal Language Areas

Summary: Subdural electrodes were implanted over the language-dominant left lateral convexity in 45 patients undergoing evaluation for epilepsy surgery. In 29 patients, additional electrodes were placed over the left basal temporal cortex. We identified language areas by using intermittent electrical stimuli applied while patients read aloud. Aphasia was classified as expressive or receptive based on additional testing performed when electrical stimulation elicited reading arrest in the absence of direct excitatory or inhibitory motor effects. Using correlated logistic regression, we noted no statistically significant differences among Broca's, and Wernicke's areas and the basal temporal language area (BTLA) regarding the frequency with which electrical stimulation interfered with language. Speech production deficits, however, occurred significantly more frequently in Broca's than in Wernicke's area (p = 0.012). In contrast, language comprehension deficits occurred with equal frequency when Broca's and Wernicke's areas were stimulated. These results suggest that both Broca's and Wernicke's areas play important roles in language comprehension and that the primarily expressive aphasia of patients with lesions of Broca's area results mainly from the predominant participation of Broca's area in language production.     

Functional assessment of Broca's area using near infrared spectroscopy in humans

We used near-infrared spectroscopy (NIRS) to compare functional hemoglobin concentration changes (delta[oxy-Hb] and delta[deoxy-Hb]) over human language and motor cortices. Eight subjects performed finger opposition, tongue movement, and covert visual object naming in an interleaved block paradigm design. NIRS revealed paradigm specific patterns of delta[oxy-Hb] and delta[deoxy-Hb] providing cortical localization of each function. During each task, significant response overlap was observed when comparing the [oxy-Hb] signals, whereas delta[deoxy-Hb] seemed more localized. Furthermore, by applying magnitude and time to significance measures to the delta[deoxy-Hb] response profile, Broca's area was easily distinguished from neighboring tongue (and hand) motor representation. Delta[oxy-Hb] did not provide this level of specificity. These findings suggest delta[deoxy-Hb] as the preferential NIRS parameter to map language cortices.     

Listening to action-related sentences activates fronto-parietal motor circuits

Observing actions made by others activates the cortical circuits responsible for the planning and execution of those same actions. This observation-execution matching system (mirror-neuron system) is thought to play an important role in the understanding of actions made by others. In an fMRI experiment, we tested whether this system also becomes active during the processing of action-related sentences. Participants listened to sentences describing actions performed with the mouth, the hand, or the leg. Abstract sentences of comparable syntactic structure were used as control stimuli. The results showed that listening to action-related sentences activates a left fronto-parieto-temporal network that includes the pars opercularis of the inferior frontal gyrus (Broca's area), those sectors of the premotor cortex where the actions described are motorically coded, as well as the inferior parietal lobule, the intraparietal sulcus, and the posterior middle temporal gyrus. These data provide the first direct evidence that listening to sentences that describe actions engages the visuomotor circuits which subserve action execution and observation.     

Temporal sorting of neural components underlying phonological processing

Event-related haemodynamic responses (EHRs) were recorded in subjects performing phonological tasks to test whether distinguishable temporal involvement of corresponding neural components would show through. A sequence of activation leading from primary auditory cortices to premotor regions emerged in the fast repetition and the phoneme monitoring tasks used. EHRs peaked significantly earlier in Wernicke's area (phonological decoding) than in Broca's area, the left supramarginal gyrus and the precentral gyrus (phonological rehearsal). Moreover, the sensitivity of within cluster temporal gradients to the nature of the tasks indicated either sensory to association cortex synchronization for fast repetition or delayed analysis for phoneme monitoring. These results are consistent with previous findings on working memory and show that fMRI permits temporal tracking of cognitive activations.     

fMRI功能导航下切除汉语运动性语言区附近病变

目的探讨功能磁共振(functional magnetic resonance imaging,fMRI)导航在汉语运动性语言区附近病变手术中的应用价值。方法对43例大脑皮层运动性语言区附近病变患者,通过组块设计的汉语朗读任务,利用血氧水平依赖性功能磁共振成像(BOLD-fMRI)获得运动性语言区激活,利用磁共振弥散张量成像技术(diffusion tensori maging,DTI)获得白质纤维束走行,将结构影像、fMRI功能影像与DTI的各项异性系数(FA)图像同时传入导航系统,术中定位fMRI确定的语言区,在唤醒状态利用皮层电刺激技术进行语言区验证,避开运动性语言区在显微镜下切除病变。结果本组患者29例术前语言功能正常,14例有不同程度语言功能障碍。38例获得了有效的fM-RI语言区激活,进行了36例核心激活脑区与皮层电刺激比较:25例为重叠关系,11例为邻近关系。手术全切除17例,次全切除14例,大部份切除12例。术后语言功能8例较术前好转,31例无变化,4例出现短暂性运动性失语。结论汉语朗读任务所获得的fMRI运动性语言区激活具有良好的敏感性与准确性;fMRI导航手术可以提高汉语运动性语言区附近病变切除程度,减少术后运动性失语的发生。