The role of inferior frontal gyrus in processing Chinese classifiers

The Chinese classifier system classifies nouns and builds a relation between classifiers and their corresponding nouns. This functional magnetic resonance imaging (fMRI) study examined brain activation of Chinese classifiers during reading comprehension. Thirty-four participants read and performed semantic congruency judgments on congruent, inside-classifier (IC) violated, and outside-classifier (OC) violated sentences. The IC and OC violations were created by changing the correct classifier to an inappropriate classifier and a non-classifier, respectively. The comparison of the IC violation vs. the congruent condition produced greater activation in the mid-ventral region (BA 45) of the left inferior frontal gyrus (IFG), suggesting an increased demand on semantic processing. Contrasting different subtypes of IC violation produced greater activation in the right IFG (BAs 45 and 47), indicating that processing mass/count classifiers involves distinct brain activations. The OC violation produced greater activation in the left IFG (BAs 45 and 44), suggesting both semantic and syntactic processing. These results indicate that different parts of the IFG contribute to syntactic and semantic processing of classifier phrases in reading Chinese for comprehension.     

Processing lexical semantic and syntactic information in first and second language: fMRI evidence from German and Russian

We introduce two experiments that explored syntactic and semantic processing of spoken sentences by native and non-native speakers. In the first experiment, the neural substrates corresponding to detection of syntactic and semantic violations were determined in native speakers of two typologically different languages using functional magnetic resonance imaging (fMRI). The results show that the underlying neural response of participants to stimuli across different native languages is quite similar. In the second experiment, we investigated how non-native speakers of a language process the same stimuli presented in the first experiment. First, the results show a more similar pattern of increased activation between native and non-native speakers in response to semantic violations than to syntactic violations. Second, the non-native speakers were observed to employ specific portions of the frontotemporal language network differently from those employed by native speakers. These regions included the inferior frontal gyrus (IFG), superior temporal gyrus (STG), and subcortical structures of the basal ganglia.     

Repair,revision, and complexity in syntactic analysis: an electrophysiological differentiation

One of the core aspects of human sentence processing is the ability to detect errors and to recover from erroneous analysis through revision of ambiguous sentences and repair of ungrammatical sentences. In the present study, we used event-related potentials (ERPs) to help identify the nature of these processes by directly comparing ERPs to complex ambiguous sentence structures with and without grammatical violations, and to simpler unambiguous sentence structures with and without grammatical violations. In ambiguous sentences, preference of syntactic analysis was manipulated such that in one condition, the structures agreed with the preferred analysis, and in another condition, a nonpreferred but syntactically correct analysis (garden path) was imposed. Nonpreferred ambiguous structures require revision, whereas ungrammatical structures require repair. We found that distinct ERPs reflected different characteristics of syntactic processing. Specifically, our results are consistent with the idea that a positivity with a posterior distribution across the scalp (posterior P600) is an index of syntactic processing difficulty, including repair and revision, and that a frontally distributed positivity (frontal P600) is related to ambiguity resolution and/or to an increase in discourse level complexity.     

Left cytoarchitectonic BA 44 processes syntactic gender violations in determiner phrases

Recent neuroimaging studies make contradictory predictions about the involvement of left Brodmann's area (BA) 44 in processing local syntactic violations in determiner phrases (DPs). Some studies suggest a role for BA 44 in detecting local syntactic violations, whereas others attribute this function to the left premotor cortex. Therefore, the present event‐related functional magnetic resonance imaging (fMRI) study investigated whether left‐cytoarchitectonic BA 44 was activated when German DPs involving syntactic gender violations were compared with correct DPs (correct: ‘der Baum’—the[masculine] tree[masculine]; violated: ‘das Baum’—the[neuter] tree[masculine]). Grammaticality judgements were made for both visual and auditory DPs to be able to generalize the results across modalities. Grammaticality judgements involved, among others, left BA 44 and left BA 6 in the premotor cortex for visual and auditory stimuli. Most importantly, activation in left BA 44 was consistently higher for violated than for correct DPs. This finding was behaviourally corroborated by longer reaction times for violated versus correct DPs. Additional brain regions, showing the same effect, included left premotor cortex, supplementary motor area, right middle and superior frontal cortex, and left cerebellum. Based on earlier findings from the literature, the results indicate the involvement of left BA 44 in processing local syntactic violations when these include morphological features, whereas left premotor cortex seems crucial for the detection of local word category violations. Hum Brain Mapp, 2010. © 2010 Wiley‐Liss, Inc.     

Brain regions that process case: Evidence from basque

The aim of this event‐related fMRI study was to investigate the cortical networks involved in case processing, an operation that is crucial to language comprehension yet whose neural underpinnings are not well‐understood. What is the relationship of these networks to those that serve other aspects of syntactic and semantic processing? Participants read Basque sentences that contained case violations, number agreement violations or semantic anomalies, or that were both syntactically and semantically correct. Case violations elicited activity increases, compared to correct control sentences, in a set of parietal regions including the posterior cingulate, the precuneus, and the left and right inferior parietal lobules. Number agreement violations also elicited activity increases in left and right inferior parietal regions, and additional activations in the left and right middle frontal gyrus. Regions‐of‐interest analyses showed that almost all of the clusters that were responsive to case or number agreement violations did not differentiate between these two. In contrast, the left and right anterior inferior frontal gyrus and the dorsomedial prefrontal cortex were only sensitive to semantic violations. Our results suggest that whereas syntactic and semantic anomalies clearly recruit distinct neural circuits, case, and number violations recruit largely overlapping neural circuits and that the distinction between the two rests on the relative contributions of parietal and prefrontal regions, respectively. Furthermore, our results are consistent with recently reported contributions of bilateral parietal and dorsolateral brain regions to syntactic processing, pointing towards potential extensions of current neurocognitive theories of language. Hum Brain Mapp, 2012. © 2011 Wiley Periodicals, Inc.     

Spatial and temporal analysis of fMRI data on word and sentence reading

Written language comprehension at the word and the sentence level was analysed by the combination of spatial and temporal analysis of functional magnetic resonance imaging (fMRI). Spatial analysis was performed via general linear modelling (GLM). Concerning the temporal analysis, local differences in neurovascular coupling may confound a direct comparison of blood oxygenation level-dependent (BOLD) response estimates between regions. To avoid this problem, we parametrically varied linguistic task demands and compared only task-induced within-region BOLD response differences across areas. We reasoned that, in a hierarchical processing system, increasing task demands at lower processing levels induce delayed onset of higher-level processes in corresponding areas. The flow of activation is thus reflected in the size of task-induced delay increases. We estimated BOLD response delay and duration for each voxel and each participant by fitting a model function to the event-related average BOLD response. The GLM showed increasing activations with increasing linguistic demands dominantly in the left inferior frontal gyrus (IFG) and the left superior temporal gyrus (STG). The combination of spatial and temporal analysis allowed a functional differentiation of IFG subregions involved in written language comprehension. Ventral IFG region (BA 47) and STG subserve earlier processing stages than two dorsal IFG regions (BA 44 and 45). This is in accordance with the assumed early lexical semantic and late syntactic processing of these regions and illustrates the complementary information provided by spatial and temporal fMRI data analysis of the same data set.     

Localizing the distributed language network responsible for the N400 measured by MEG during auditory sentence processing

We studied auditory sentence comprehension using magnetoencephalography while subjects listened to sentences whose correctness they had to judge subsequently. The localization and the time course of brain electrical activity during processing of correct and semantically incorrect sentences were estimated by computing a brain surface current density within a cortical layer for both conditions. Finally, a region of interest (ROI) analysis was conducted to determine the time course of specific locations. A magnetic N400 was present in six spatially different ROIs. Semantic anomalies caused an exclusive involvement of the ventral portion of the left inferior frontal gyrus (BA 47) and left pars triangularis (BA 45). The anterior parts of the superior (BA 22) and inferior (BA 20/21) temporal gyri bilaterally were activated by both conditions. The activation for the correct condition, however, peaked earlier in both left temporal regions (approximately 32 ms). In general, activation due to semantic violations was more pronounced, started later, and lasted longer as compared to correct sentences. The findings reveal a clear left-hemispheric dominance during language processing indicated firstly by the mere number of activated regions (four in the left vs. two in the right hemisphere) and secondly by the observed specificity of the left inferior frontal ROIs to semantic violations. The temporal advantage observed for the correct condition in the left temporal regions is supporting the notion that the established context eases the processing of the final word. Semantically incorrect words that do not fit into the context result in longer integration times.     

Disentangling syntax and intelligibility in auditory language comprehension

Studies of the neural basis of spoken language comprehension typically focus on aspects of auditory processing by varying signal intelligibility, or on higher‐level aspects of language processing such as syntax. Most studies in either of these threads of language research report brain activation including peaks in the superior temporal gyrus (STG) and/or the superior temporal sulcus (STS), but it is not clear why these areas are recruited in functionally different studies. The current fMRI study aims to disentangle the functional neuroanatomy of intelligibility and syntax in an orthogonal design. The data substantiate functional dissociations between STS and STG in the left and right hemispheres: first, manipulations of speech intelligibility yield bilateral mid‐anterior STS peak activation, whereas syntactic phrase structure violations elicit strongly left‐lateralized mid STG and posterior STS activation. Second, ROI analyses indicate all interactions of speech intelligibility and syntactic correctness to be located in the left frontal and temporal cortex, while the observed right‐hemispheric activations reflect less specific responses to intelligibility and syntax. Our data demonstrate that the mid‐to‐anterior STS activation is associated with increasing speech intelligibility, while the mid‐to‐posterior STG/STS is more sensitive to syntactic information within the speech. Hum Brain Mapp, 2010. © 2009 Wiley‐Liss, Inc.     

Asyntactic comprehension, working memory, and acute ischemia in Broca's area versus angular gyrus

We evaluated sentence comprehension of variety of sentence constructions and components of short-term memory (STM) in 53 individuals with acute ischemic stroke, to test some current hypotheses about the role of Broca's area in these tasks. We found that some patients show structure-specific, task-independent deficits in sentence comprehension, with chance level of accuracy on passive reversible sentences, more impaired comprehension of object-cleft than subject-cleft sentences, and more impaired comprehension of reversible than irreversible sentences in both sentence-picture matching and enactment tasks. In a dichotomous analysis, this pattern of "asyntactic comprehension" was associated with dysfunctional tissue in left angular gyrus, rather than dysfunctional tissue in Broca's area as previously proposed. Tissue dysfunction in left Brodmann area (BA) 44, part of Broca's area, was associated with phonological STM impairment defined by forward digit span≤4. Verbal working memory (VWM) defined by backward digit span≤2 was associated with tissue dysfunction left premotor cortex (BA 6). In a continuous analysis, patients with acute ischemia in left BA 44 were impaired in phonological STM. Patients with ischemia in left BA 45 and BA 6 were impaired in passive, reversible sentences, STM, and VWM. Patients with ischemia in left BA 39 were impaired in passive reversible sentences, object-cleft sentences, STM, and VWM. Therefore, various components of working memory seem to depend on a network of brain regions that include left angular gyrus and posterior frontal cortex (BA 6, 44, 45); left BA 45 and angular gyrus (BA 39) may have additional roles in comprehension of syntax such as thematic role checking.     

ERP effects of listening to speech compared to reading: the P600/SPS to syntactic violations in spoken sentences and rapid serial visual presentation

In this study, event-related brain potential effects of speech processing are obtained and compared to similar effects in sentence reading. In two experiments sentences were presented that contained three different types of grammatical violations. In one experiment sentences were presented word by word at a rate of four words per second. The grammatical violations elicited a Syntactic Positive Shift (P600/SPS), 500 ms after the onset of the word that rendered the sentence ungrammatical. The P600/SPS consisted of two phases, an early phase with a relatively equal anterior-posterior distribution and a later phase with a strong posterior distribution. We interpret the first phase as an indication of structural integration complexity, and the second phase as an indication of failing parsing operations and/or an attempt at reanalysis. In the second experiment the same syntactic violations were presented in sentences spoken at a normal rate and with normal intonation. These violations elicited a P600/SPS with the same onset as was observed for the reading of these sentences. In addition two of the three violations showed a preceding frontal negativity, most clearly over the left hemisphere.     

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.     

fMRI evidence for dual routes to the mental lexicon in visual word recognition.

Event-related fMRI was used to investigate lexical decisions to words of high and low frequency of occurrence and to pseudowords. The results obtained strongly support dual-route models of visual word processing. By contrasting words with pseudowords, bilateral occipito-temporal brain areas and posterior left middle temporal gyrus (MTG) were identified as contributing to the successful mapping of orthographic percepts onto visual word form representations. Low-frequency words and pseudowords elicited greater activations than high-frequency words in the superior pars opercularis [Brodmann's area (BA) 44] of the left inferior frontal gyrus (IFG), in the anterior insula, and in the thalamus and caudate nucleus. As processing of these stimuli during lexical search is known to rely on phonological information, it is concluded that these brain regions are involved in grapheme-to-phoneme conversion. Activation in the pars triangularis (BA 45) of the left IFG was observed only for low-frequency words. It is proposed that this region is involved in processes of lexical selection.     

Ventral premotor cortex lesions disrupt learning of sequential grammatical structures

IntroductionRecent functional magnetic resonance imaging (fMRI) evidence shows differential involvement of the inferior frontal gyrus (IFG) and the ventral premotor cortex (PMv) in syntactic processing. Our main goal is to specify the precise role of the PMv in the processing of sequential structures and whether these processes are a necessary prerequisite for the successful acquisition of grammatical structure.MethodsWe tested patients with PMv lesions in an artificial grammar (AG) learning task, including correct sentences and sentences with violations of local (referring to adjacent elements within an AG string) and long-distance dependencies (incorporating recursive structures). In addition to performance measures event-related potentials to these violations were recorded.Results and conclusionsCompared to matched controls, patients displayed impaired acquisition of the AG. This impairment was more pronounced for local than for long-distance dependencies. This effect was paralleled by a selective reduction of the P600 component in response to violations of local dependencies. Most importantly, the P600 elicited by violations of long-distance dependencies was comparable between groups. Together, behavioral and ERP results indicate a PMv involvement in processing local sequential information.     

Reviewing the functional basis of the syntactic Merge mechanism for language: A coordinate-based activation likelihood estimation meta-analysis

The ability to create structures out of single words is a key aspect of human language. This combinatorial capacity relies on a low-level syntactic mechanism—Merge—assembling words into hierarchies. Neuroscience has explored Merge by comparing syntax to word-lists. Here, we first review potential issues with the word-lists materials. We then perform an activation likelihood estimation (ALE) on the reported foci, to reveal functional convergence for Merge at whole-brain level. Finally, we run probabilistic tractography on an independent population to observe how these convergent activations anatomically connect. Functionally, we found that when confounding activity was removed, consistency for Merge was only observable in the left pars opercularis (BA44) and in the inferior part of the posterior superior temporal sulcus/gyrus (pSTS/STG; BA22). Structurally, we could confirm that the two regions are connected through dorsal fiber bundles. We therefore suggest that the cortical implementation of linguistic Merge consists of a left fronto-temporal interaction between BA44 (syntactic processor) and pSTS/STG (integrative processor), which communicate to each other along dorsal white matter fascicles.     

Differential effects of syntactic and semantic processing on the subregions of Broca's area

This study attempts to specify the contribution of two subregions of Broca's area during syntactic and semantic processing of sentences by examining brain activation in a grammaticality judgment task. The processing of two types of ungrammatical sentences was examined. One type leaves the thematic interpretation generally unaffected, by violating the noun-verb agreement in number, while the other type introduces an extraneous verb, which cannot be incorporated into the developing thematic structure. Pars triangularis was more sensitive to the extra verb violation, whereas pars opercularis was more sensitive to the noun-verb agreement violation. The current study adds to the growing literature that suggests there are separable functional subregions of Broca's area, with pars triangularis more involved in thematic processing and pars opercularis more involved in syntactic processing. The posterior left temporal area was also involved in both types of processing.     

Determining hierarchical functional networks from auditory stimuli fMRI

We determined connectivity of the human brain using functional magnetic resonance imaging (fMRI) while subjects experienced auditory stimuli in a 2-by-2 factorial design. The two factors in this study were "speaker" (same or different speaker) and "sentence" (same or different sentences). Connectivity studies allow us to ask how spatially remote brain regions are neurophysiologically related given these stimuli. In the context of this study, we examined how the "speaker" effect and "sentence" effect influenced these relationships. We applied a Bayesian connectivity method that determines hierarchical functional networks of functionally connected brain regions. Hierarchy in these functional networks is determined by conditional probabilities of elevated activity. For example, a brain region that becomes active a superset of the time of another region is considered ascendant to that brain region in the hierarchical network. For each factor level, we found a baseline functional network connecting the primary auditory cortex (Brodmann's Area [BA] 41) with the BA 42 and BA 22 of the superior temporal gyrus (STG). We also found a baseline functional network that includes Wernicke's Area (BA 22 posterior), STG, and BA 44 for each factor level. However, we additionally observed a strong ascendant connection from BA 41 to the posterior cingulate (BA 30) and Broca's Area and a stronger connection from Wernicke's Area to STG and the posterior cingulate while passively listening to different sentences rather than the same sentence repeatedly. Finally, our results revealed no significant "speaker" effect or interaction between "speaker" and "sentence."     

Response of anterior temporal cortex to syntactic and prosodic manipulations during sentence processing

Previous research has implicated a portion of the anterior temporal cortex in sentence-level processing. This region activates more to sentences than to word-lists, sentences in an unfamiliar language, and environmental sound sequences. The current study sought to identify the relative contributions of syntactic and prosodic processing to anterior temporal activation. We presented auditory stimuli where the presence of prosodic and syntactic structure was independently manipulated during functional magnetic resonance imaging (fMRI). Three "structural" conditions included normal sentences, sentences with scrambled word order, and lists of content words. These three classes of stimuli were presented either with sentence prosody or with flat supra-lexical (list-like) prosody. Sentence stimuli activated a portion of the left anterior temporal cortex in the superior temporal sulcus (STS) and extending into the middle temporal gyrus, independent of prosody, and to a greater extent than any of the other conditions. An interaction between the structural conditions and prosodic conditions was seen in a more dorsal region of the anterior temporal lobe bilaterally along the superior temporal gyrus (STG). A post-hoc analysis revealed that this region responded either to syntactically structured stimuli or to nonstructured stimuli with sentence-like prosody. The results suggest a parcellation of anterior temporal cortex into 1) an STG region that is sensitive both to the presence of syntactic information and is modulated by prosodic manipulations (in nonsyntactic stimuli); and 2) a more inferior left STS/MTG region that is more selective for syntactic structure.     

Overt sentence production in event-related fMRI

The use of syntactic structures on a sentence level is a unique human ability. Functional imaging studies have usually investigated syntax comprehension. However, language production may be performed by different neuronal resources. We have investigated syntax generation on a sentence level with functional magnetic resonance imaging (fMRI). BOLD contrast was measured while subjects articulated utterances aloud. In the active condition 'sentence generation' (SG), subjects had to produce subject verb object (SVO) sentences (e.g. "The child throws the ball") according to syntactically incomplete stimuli (e.g. "throw ball child") presented visually. In the control condition 'word reading' (WR), subjects had to read identical stimuli without completing the syntactic structure, while in a second control condition 'sentence reading' (SR), subjects had to read complete sentences. The semantic meaning of all expressions was obvious despite the syntactically incomplete structure in conditions SG and WR. In both contrasts, SG minus WR and SG minus SR, activation was mainly present in the left inferior frontal (BA 44/45) and medial frontal (BA 6) gyri, the superior parietal lobule (BA 7) and the right insula (BA 13). A region of interest analysis revealed significantly stronger left-dominant activation in BA 45 compared to BA 44. Our data illustrates the crucial involvement of the left BA 45 in syntactic encoding and is in line with more recent imaging and brain lesion data on syntax processing on a sentence level, emphasizing the involvement of a distributed left and right hemispheric network in syntax generation.     

An fMRI examination of the effects of acoustic-phonetic and lexical competition on access to the lexical-semantic network

The current study explored how factors of acoustic-phonetic and lexical competition affect access to the lexical-semantic network during spoken word recognition. An auditory semantic priming lexical decision task was presented to subjects while in the MR scanner. Prime-target pairs consisted of prime words with the initial voiceless stop consonants /p/, /t/, and /k/ followed by word and nonword targets. To examine the neural consequences of lexical and sound structure competition, primes either had voiced minimal pair competitors or they did not, and they were either acoustically modified to be poorer exemplars of the voiceless phonetic category or not. Neural activation associated with semantic priming (Unrelated–Related conditions) revealed a bilateral fronto-temporo-parietal network. Within this network, clusters in the left insula/inferior frontal gyrus (IFG), left superior temporal gyrus (STG), and left posterior middle temporal gyrus (pMTG) showed sensitivity to lexical competition. The pMTG also demonstrated sensitivity to acoustic modification, and the insula/IFG showed an interaction between lexical competition and acoustic modification. These findings suggest the posterior lexical-semantic network is modulated by both acoustic-phonetic and lexical structure, and that the resolution of these two sources of competition recruits frontal structures.