Left Dorsal Inferior Frontal Gyrus Research Articles

Phonological and Semantic Specialization in 9- to 10-Year-Old Children During Auditory Word Processing.

One of the core features of brain maturation is functional specialization. Previous research has found that 7- to 8-year-old children start to specialize in both the temporal and frontal lobes. However, as children continue to develop their phonological and semantic skills rapidly until approximately 10 years old, it remained unclear whether any changes in specialization later in childhood would be detected. Thus, the goal of the current study was to examine phonological and semantic specialization in 9- to 10-year-old children during auditory word processing. Sixty-one children were included in the analysis. They were asked to perform a sound judgment task and a meaning judgment task, each with both hard and easy conditions to examine parametric effects. Consistent with previous results from 7- to 8-year-old children, direct task comparisons revealed language specialization in both the temporal and frontal lobes in 9- to 10-year-old children. Specifically, the left dorsal inferior frontal gyrus showed greater activation for the sound than the meaning task whereas the left middle temporal gyrus showed greater activation for the meaning than the sound task. Interestingly, in contrast to the previously reported finding that 7- to 8-year-old children primarily engage a general control region during the harder condition for both tasks, we showed that 9- to 10-year-old children recruited language-specific regions to process the more difficult task conditions. Specifically, the left superior temporal gyrus showed greater activation for the phonological parametric manipulation whereas the left ventral inferior frontal gyrus showed greater activation for the semantic parametric manipulation.

Distinct neural correlates of poor decoding and poor comprehension in children with reading disability.

Reading disability (RD) can manifest itself as a word decoding problem or a reading comprehension problem. In the current study, we identified 3 subtypes of RD: poor decoders (PD), poor comprehenders (PC), and poor-in-both (PB). We found that PD had greater deficits in meta-linguistic skills such as phonological awareness, orthographic skills, and morphological skills than PC, whereas PC had greater deficits in listening comprehension than PD. In the brain, we also found different patterns of deficits during an auditory rhyming judgment task using functional magnetic resonance imaging. PD showed less activation than PC and age controls in the left dorsal inferior frontal gyrus (IFG) and pre-supplementary motor area (SMA), brain activation of which was correlated with phonological awareness and working memory. In contrast, PC showed less activation in the left fusiform gyrus than PD and age controls, which was correlated with reading comprehension fluency and morphological skill. Last, PB showed both PD's and PC's deficits, as well as additional deficits in the bilateral lingual gyri. Our findings contribute to revealing different neural signatures of poor decoding and poor comprehension, which are distinct disorders but co-occur very often. These findings implicate possibility and necessity of precise diagnosis and individualized intervention.

A hierarchical deficit model of reading disability: Evidence from dynamic causal modelling analysis

Deficits have been documented in visuo-orthographic processing as well as phonological retrieval/manipulation during visual word reading in individuals with reading disability (RD); however, the relationship between these deficits remains unclear. Previously, we found that during word reading, visuo-orthographic deficit appears to be a neural signature of RD, but deficits in phonological retrieval/manipulation appears to be a consequence of being RD (Cao et al., 2020). Therefore, in the current study, we directly tested the hypothesis that during visual word reading, deficit in phonological retrieval/manipulation may result from weakened input from visuo-orthographic regions, and that this relationship tends to be universal across languages. We conducted a dynamic causal modelling analysis of fMRI data from Chinese-English bilingual children (9–11 years, N = 78) with or without RD during a visual word rhyming judgment task. We found a weaker connection from the left inferior temporal gyrus (ITG) to the left dorsal inferior frontal gyrus (dIFG) in children with RD and reading controls than the connection found in age controls for both Chinese and English. This finding suggests that the phonological deficit at the dIFG may result from weak input from the visuo-orthographic region and this connection appears to be responsive to reading level rather than RD, because the reading-control children were similar to children with RD. We also found that the left ITG was selectively connected with language-specific regions (i.e., the left inferior parietal lobe (IPL) for Chinese and the left ventral inferior frontal gyrus (vIFG) for English) depending on the language being processed; however, this language selectivity was reduced in children with RD, suggesting that decreased language specialization is associated with RD. Using a double control design, our study suggests that during reading, the visuo-orthographic deficit of RD constrains the development of the connection from orthography to phonology and to other language-specific processing due to distorted quantity and quality of reading.

Reading Disability in Chinese Children Learning English as an L2

To examine whether there are common or specific deficits of reading disability (RD) in first (L1) and second languages (L2), Chinese children (9-11years, N=76) with or without RD who learn English as an L2 were studied during a visual word rhyming judgment task. Evidence was found for common deficits in L1 and L2 in visuo-orthographic processes in left inferior temporal gyrus and left precuneus, as well as in phonological processes in left dorsal inferior frontal gyrus as children with RD showed less activation than controls in both languages. Furthermore, the visuo-orthographic deficit appears to be a RD effect, whereas the phonological deficit appears to be a reading/performance effect. Some weak evidence for language specific effects was also found.

Dynamic Functional Connectivity Within the Fronto-Limbic Network Induced by Intermittent Theta-Burst Stimulation: A Pilot Study.

PurposeThe utility of transcranial magnetic stimulation (TMS) has been growing rapidly in both neurocognitive studies and clinical applications in decades. However, it remains unclear how the responses of the stimulated site and the site-related functional network to the external TMS manipulation dynamically change over time.MethodsA multi-session combining TMS-fMRI experiment was conducted to explore the spatiotemporal effects of TMS within the fronto-limbic network. Ten healthy volunteers were modulated by intermittent theta-burst stimulation (iTBS) at a precise site within the left dorsolateral prefrontal cortex (DLPFC, MNI coordinate [-44 36 20]), navigated by individual structural MRI image. Three-session resting-state fMRI images were acquired before iTBS (TP1), immediately after iTBS (TP2), and 15 min after iTBS (TP3) for each participant. Seventy-four regions of interests (ROIs) within the fronto-limbic network were chosen including the bilateral superior frontal gyrus (SFG), middle frontal gyrus (MidFG), inferior frontal gyrus (IFG), orbital gyrus (OrG), cingulate gyrus (CG), and subcortical nuclei (hippocampus and amygdala). Regional fractional amplitude of low-frequency fluctuation (fALFF) and ROI-to-ROI functional connectivity (FC) were compared among TP1, TP2, and TP3.ResultsThe immediate iTBS effect was observed at the stimulated site. FC between the left dorsolateral SFG and left dorsal IFG and between the left rostral IFG and right MidFG increased at TP2 as compared to at TP1 (all FDR-p < 0.05), while FC within the left OrG decreased. The relatively long-term iTBS effect transmitted with decreased FC between the left IFG and right amygdala, increased FC between the left MidFG and left OrG, and decreased FC between bilateral IFG and OrG at TP3 than at TP1 (all FDR-p < 0.05). Meanwhile, mean fALFF values over the left SFG, MidFG, ventral CG, and IFG were significantly increased at TP3 as compared to those at TP2 (all p < 0.05 with Bonferroni correction).ConclusionBy combining TMS and fMRI, it becomes possible to track the spatiotemporal dynamics of TMS after-effects within the fronto-limbic network. Our findings suggested that the iTBS effect dynamically changed over time from the local neural activation at the stimulated site to its connected remote regions within the fronto-limbic network.

Hemodynamics of speech production: An fNIRS investigation of children who stutter

Stuttering affects nearly 1% of the population worldwide and often has life-altering negative consequences, including poorer mental health and emotional well-being, and reduced educational and employment achievements. Over two decades of neuroimaging research reveals clear anatomical and physiological differences in the speech neural networks of adults who stutter. However, there have been few neurophysiological investigations of speech production in children who stutter. Using functional near-infrared spectroscopy (fNIRS), we examined hemodynamic responses over neural regions integral to fluent speech production including inferior frontal gyrus, premotor cortex, and superior temporal gyrus during a picture description task. Thirty-two children (16 stuttering and 16 controls) aged 7–11 years participated in the study. We found distinctly different speech-related hemodynamic responses in the group of children who stutter compared to the control group. Whereas controls showed significant activation over left dorsal inferior frontal gyrus and left premotor cortex, children who stutter exhibited deactivation over these left hemisphere regions. This investigation of neural activation during natural, connected speech production in children who stutter demonstrates that in childhood stuttering, atypical functional organization for speech production is present and suggests promise for the use of fNIRS during natural speech production in future research with typical and atypical child populations.

Activation changes of the left inferior frontal gyrus for the factors of construction and scrambling in a sentence.

Our previous functional magnetic resonance imaging (fMRI) studies have indicated that the left dorsal inferior frontal gyrus (L. dF3op/F3t) and left lateral premotor cortex (L. LPMC) are crucial regions for syntactic processing among the syntax-related networks. In the present study, we further examined how activations in these regions were modified by the factors of construction and scrambling (object-initial type). Using various sentence types, we clarified three major points. First, we found that the main effects of construction and scrambling consistently activated the L. dF3op/F3t and L. LPMC. Secondly, the main effects of scrambling clearly localized activation in the L. dF3op/F3t and L. LPMC, indicating the more narrowed down processing of syntax. Thirdly, step-wise percent signal changes were observed in the L. dF3op/F3t, demonstrating synergistic effects of construction and scrambling. These results demonstrate the abstract and intensive nature of syntactic processing carried out by these regions, i.e., the grammar center.

Shared orthographic neuronal representations for spelling and reading

A central question in the study of the neural basis of written language is whether reading and spelling utilize shared orthographic representations. While recent studies employing fMRI to test this question report that the left inferior frontal gyrus (IFG) and ventral occipitotemporal cortex (vOTC) are active during both spelling and reading in the same subjects (Purcell et al., 2011a; Rapp and Lipka, 2011), the spatial resolution of fMRI limits the interpretation of these findings. Specifically, it is unknown if the neurons which encode orthography for reading are also involved in spelling of the same words. Here we address this question by employing an event-related functional magnetic resonance imaging-adaptation (fMRI-A) paradigm designed to examine shared orthographic representations across spelling and reading. First, we identified areas that independently showed adaptation to reading, and adaptation to spelling. Then we identified spatial convergence for these two separate maps via a conjunction analysis. Consistent with previous studies (Purcell et al., 2011a; Rapp and Lipka, 2011), this analysis revealed the left dorsal IFG, vOTC and supplementary motor area. To further validate these observations, we then interrogated these regions using an across-task adaptation technique, and found adaptation across reading and spelling in the left dorsal IFG (BA 44/9). Our final analysis focused specifically on the Visual Word Form Area (VWFA) in the vOTC, whose variability in location among subjects requires the use of subject-specific identification mechanisms (Glezer and Riesenhuber, 2013). Using a functional localizer for reading, we defined the VWFA in each subject, and found adaptation effects for both within the spelling and reading conditions, respectively, as well as across spelling and reading. Because none of these effects were observed during a phonological/semantic control condition, we conclude that the left dorsal IFG and VWFA are involved in accessing the same orthography-specific representations for spelling and reading.

Neural signatures of phonological deficits in Chinese developmental dyslexia

There has been debate on whether phonological deficits explain reading difficulty in Chinese, since Chinese is a logographic language which does not employ grapheme-phoneme-correspondence rules and remote memorization seems to be the main method to acquire reading. In the current study, we present neuroimaging evidence that the phonological deficit is also a signature of Chinese dyslexia. Specifically, we found that Chinese children with dyslexia (DD) showed reduced brain activation in the left dorsal inferior frontal gyrus (dIFG) when compared to both age-matched controls (AC) and reading-matched controls (RC) during an auditory rhyming judgment task. This suggests that the phonological processing deficit in this region may be a signature of dyslexia in Chinese, rather than a difference due to task performance or reading ability, which was matched on DD and RC. At exactly the same region of the left dIFG, we found a positive correlation between brain activation and reading skill in DD, suggesting that the phonological deficit is associated with the severity of dyslexia. We also found increased brain activation in the right precentral gyrus in DD than both AC and RC, suggesting a compensation of reliance on articulation. Functional connectivity analyses revealed that DD had a weaker connection between the left superior temporal gyrus (STG) and fusiform gyrus (FG) than the two control groups, suggesting that the reduced connection between phonology and orthography is another neural signature of dyslexia. In contrast, DD showed greater connectivity between the left dIFG and the left inferior parietal lobule (IPL) than both control groups, suggesting a reduced segregation between the language network and default mode network in dyslexic children. We also found that connectivity between the left STG and the left dIFG was sensitive to task performance and/or reading skill rather than being dyslexic or not, because AC was greater than both RC and DD, while the connectivity between the left middle occipital gyrus (MOG) and left STG was sensitive to age, because both AC and DD were greater than RC. In summary, our study provides the very first neurological evidence of phonological deficits in Chinese developmental dyslexia and we successfully distinguished variations of brain activity/functional connectivity due to age, performance, and dyslexia by comparing AC, RC, and DD.

Sentence processing and verbal working memory in a white-matter-disconnection patient

The Arcuate Fasciculus/Superior Longitudinal Fasciculus (AF/SLF) is the white-matter bundle that connects posterior superior temporal and inferior frontal cortex. Its causal functional role in sentence processing and verbal working memory is currently under debate. While impairments of sentence processing and verbal working memory often co-occur in patients suffering from AF/SLF damage, it is unclear whether these impairments result from shared white-matter damage to the verbal-working-memory network. The present study sought to specify the behavioral consequences of focal AF/SLF damage for sentence processing and verbal working memory, which were assessed in a single patient suffering from a cleft-like lesion spanning the deep left superior temporal gyrus, sparing most surrounding gray matter. While tractography suggests that the ventral fronto-temporal white-matter bundle is intact in this patient, the AF/SLF was not visible to tractography. In line with the hypothesis that the AF/SLF is causally involved in sentence processing, the patient׳s performance was selectively impaired on sentences that jointly involve both complex word orders and long word-storage intervals. However, the patient was unimpaired on sentences that only involved long word-storage intervals without involving complex word orders. On the contrary, the patient performed generally worse than a control group across standard verbal-working-memory tests. We conclude that the AF/SLF not only plays a causal role in sentence processing, linking regions of the left dorsal inferior frontal gyrus to the temporo-parietal region, but moreover plays a crucial role in verbal working memory, linking regions of the left ventral inferior frontal gyrus to the left temporo-parietal region. Together, the specific sentence-processing impairment and the more general verbal-working-memory impairment may imply that the AF/SLF subserves both sentence processing and verbal working memory, possibly pointing to the AF and SLF respectively supporting each.

The neural substrates associated with attentional resources and difficulty of concurrent processing of the two verbal tasks

The kana pick-out test has been widely used in Japan to evaluate the ability to divide attention in both adult and pediatric patients. However, the neural substrates underlying the ability to divide attention using the kana pick-out test, which requires participants to pick out individual letters (vowels) in a story while also reading for comprehension, thus requiring simultaneous allocation of attention to both activities, are still unclear. Moreover, outside of the clinical area, neuroimaging studies focused on the mechanisms of divided attention during complex story comprehension are rare. Thus, the purpose of the present study, to clarify the neural substrates of kana pick-out test, improves our current understanding of the basic neural mechanisms of dual task performance in verbal memory function. We compared patterns of activation in the brain obtained during performance of the individual tasks of vowel identification and story comprehension, to levels of activation when participants performed the two tasks simultaneously during the kana pick-out test. We found that activations of the left dorsal inferior frontal gyrus and superior parietal lobule increase in functional connectivity to a greater extent during the dual task condition compared to the two single task conditions. In contrast, activations of the left fusiform gyrus and middle temporal gyrus, which are significantly involved in picking out letters and complex sentences during story comprehension, respectively, were reduced in the dual task condition compared to during the two single task conditions. These results suggest that increased activations of the dorsal inferior frontal gyrus and superior parietal lobule during dual task performance may be associated with the capacity for attentional resources, and reduced activations of the left fusiform gyrus and middle temporal gyrus may reflect the difficulty of concurrent processing of the two tasks. In addition, the increase in synchronization between the left dorsal inferior frontal gyrus and superior parietal lobule in the dual task condition may induce effective communication between these brain regions and contribute to more attentional processing than in the single task condition, due to greater and more complex demands on voluntary attentional resources.

Higher or lower? The functional anatomy of perceived allocentric social hierarchies

The perception and judgement of social hierarchies forms an integral part of social cognition. Hierarchical judgements can be either self-referential or allocentric (pertaining to two or more external agents). In psychiatric conditions such as dissocial personality disorder and schizophrenia, the impact of hierarchies may be problematic. We sought to elucidate the brain regions involved in judging allocentric social hierarchies. Twenty-two healthy male subjects underwent three fMRI scans. During scanning, subjects answered questions concerning visually-presented target pairs of human individual's relative superiority within a specific social hierarchy or their perceived degree of social alliance (i.e., whether they were "friends or enemies"). Subjects also made judgements relating to target pairs' age, gender and fame to control for confounding factors and performed a baseline numerical task. Response times increased in line with hypothesized ascending executive load. Both social hierarchy and social alliance judgements activated left ventrolateral prefrontal cortex (VLPFC), left dorsal inferior frontal gyrus (IFG) and bilateral fusiform gyri. In addition, social alliance judgements activated right dorsal IFG and medial prefrontal cortex. When compared directly with social alliance, social hierarchy judgements activated left orbitofrontal cortex. Detecting the presence of social hierarchies and judging other's relative standing within them implicates the cognitive executive, in particular the VLPFC. Our finding informs accounts of 'normal' social cognition but our method also provides a means of probing the dissocial brain in personality disorder and schizophrenia where executive function may be dysfunctional.

Modality- and Task-specific Brain Regions Involved in Chinese Lexical Processing

fMRI was used to examine lexical processing in native adult Chinese speakers. A 2 task (semantics and phonology) x 2 modality (visual and auditory) within-subject design was adopted. The semantic task involved a meaning association judgment and the phonological task involved a rhyming judgment to two sequentially presented words. The overall effect across tasks and modalities was used to identify seven ROIs, including the left fusiform gyrus (FG), the left superior temporal gyrus (STG), the left ventral inferior frontal gyrus (VIFG), the left middle temporal gyrus (MTG), the left dorsal inferior frontal gyrus (DIFG), the left inferior parietal lobule (IPL), and the left middle frontal gyrus (MFG). ROI analyses revealed two modality-specific areas, FG for visual and STG for auditory, and three task-specific areas, IPL and DIFG for phonology and VIFG for semantics. Greater DIFG activation was associated with conflicting tonal information between words for the auditory rhyming task, suggesting this region's role in strategic phonological processing, and greater VIFG activation was correlated with lower association between words for both the auditory and the visual meaning task, suggesting this region's role in retrieval and selection of semantic representations. The modality- and task-specific effects in Chinese revealed by this study are similar to those found in alphabetical languages. Unlike English, we found that MFG was both modality- and task-specific, suggesting that MFG may be responsible for the visuospatial analysis of Chinese characters and orthography-to-phonology integration at a syllabic level.

Neural correlates of morphosyntactic and verb-argument structure processing: An EfMRI study

In the current study, we investigated the processing of ungrammatical sentences containing morphosyntactic and verb-argument structure violations in an fMRI paradigm. In the morphosyntactic condition, participants listened to German perfect tense sentences with morphosyntactic violations which were neither related to finiteness nor to agreement but which were based on a syntactic feature mismatch between two verbal elements. When compared to correct sentences, morphosyntactically ungrammatical sentences elicited an increase in brain activity in the left middle to posterior superior temporal gyrus (STG). In the verb-argument structure condition, sentences were either correct or contained an intransitive verb with an unlicensed direct object. Ungrammatical sentences of this type elicited brain activations in the left inferior frontal gyrus (IFG) (BA 44). Thus, we found evidence for different brain activity patterns as a function of violation type. The left posterior STG, an area known to support lexical–syntactic integration was strongly implicated in morphosyntactic processing whereas the left dorsal IFG (BA 44) was seen to be involved in the processing of verb-argument structure. Our results suggest that lexical, syntactic and semantic features of verbal stimuli interact in a complex fashion during language comprehension.

Developmental changes in brain regions involved in phonological and orthographic processing during spoken language processing

Developmental differences in brain activation of 9- to 15-year-old children were examined during an auditory rhyme decision task to spoken words using functional magnetic resonance imaging (fMRI). As a group, children showed activation in the left superior/middle temporal gyri (BA 22, 21), right middle temporal gyrus (BA 21), dorsal (BA 45, pars opercularis) and ventral (BA 46, pars triangularis) aspects of the left inferior frontal gyrus, and left fusiform gyrus (BA 37). There was a developmental increase in activation in the left middle temporal gyrus (BA 22) across all lexical conditions, suggesting that automatic semantic processing increases with age regardless of task demands. Activation in the left dorsal inferior frontal gyrus also showed developmental increases for the conflicting (e.g. PINT–MINT) compared to the non-conflicting (e.g. PRESS–LIST) non-rhyming conditions, indicating that this area becomes increasingly involved in strategic phonological processing in the face of conflicting orthographic and phonological representations. Left inferior temporal/fusiform gyrus (BA 37) activation was also greater for the conflicting (e.g. PINT–MINT) condition, and a developmental increase was found in the positive relationship between individuals' reaction time and activation in the left lingual/fusiform gyrus (BA 18) in this condition, indicating an age-related increase in the association between longer reaction times and greater visual-orthographic processing in this conflicting condition. These results suggest that orthographic processing is automatically engaged by children in a task that does not require access to orthographic information for correct performance, especially when orthographic and phonological representations conflict, and especially for longer response latencies in older children.

Fractionation of the Component Processes Underlying Successful Episodic Encoding: A Combined fMRI and Divided-attention Study

Considerable evidence suggests that attentional resources are necessary for the encoding of episodic memories, but the nature of the relationship between attention and neural correlates of encoding is unclear. Here we address this question using functional magnetic resonance imaging and a divided-attention paradigm in which competition for different types of attentional resources was manipulated. Fifteen volunteers were scanned while making animacy judgments to visually presented words and concurrently performing one of three tasks on auditorily presented words: male/female voice discrimination (control task), 1-back voice comparison (1-back task), or indoor/outdoor judgment (semantic task). The 1-back and semantic tasks were designed to compete for task-generic and task-specific attentional resources, respectively. Using the "remember/know" procedure, memory for the study words was assessed after 15 min. In the control condition, subsequent memory effects associated with later recollection were identified in the left dorsal inferior frontal gyrus and in the left hippocampus. These effects were differentially attenuated in the two more difficult divided-attention conditions. The effects of divided attention seem, therefore, to reflect impairments due to limitations at both task-generic and task-specific levels. Additionally, each of the two more difficult divided-attention conditions was associated with subsequent memory effects in regions distinct from those showing effects in the control condition. These findings suggest the engagement of alternative encoding processes to those engaged in the control task. The overall pattern of findings suggests that divided attention can impact later memory in different ways, and accordingly, that different attentional resources, including task-generic and task-specific resources, make distinct contributions to successful episodic encoding.

Encoding and the Durability of Episodic Memory: A Functional Magnetic Resonance Imaging Study

Memories vary in their durability even when encoding conditions apparently remain constant. We investigated whether, under these circumstances, memory durability is nonetheless associated with variation in the neural activity elicited during encoding. Event-related functional magnetic resonance imaging data were acquired while volunteers semantically classified visually presented words. Using the "remember/know" procedure, memory for one-half of the words was tested after 30 min and for the remaining half after 48 h. In several regions, including left hippocampus and left dorsal inferior frontal gyrus (IFG), activity at encoding differed depending on whether items were later recollected regardless of study-test delay. Delay-selective effects were also evident, however. Recollection after 48 h was associated with enhanced activity in bilateral ventral IFG, whereas recollection after 30 min was associated with greater fusiform activity. Thus, there is a relationship between the neural activity elicited by an event as it is encoded and the durability of the resulting memory representation.