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In everyday life we need to continuously regulate our emotional responses according to their social context. Strategies of emotion regulation allow individuals to control time, intensity, nature and expression of emotional responses to environmental stimuli. The left inferior frontal gyrus (LIFG) is involved in the cognitive control of the selection of semantic content. We hypothesized that it might also be involved in the regulation of emotional feelings and expressions. We applied continuous theta burst stimulation (cTBS) over LIFG or a control site before a newly-developed ecological regulation task that required participants to produce storytelling of pictures with negative or neutral valence to either a peer (unregulated condition) or a child (regulated condition). Linguistic, expressive, and physiological responses were analyzed in order to assess the effects of LIFG-cTBS on emotion regulation. Results showed that the emotion regulation context modulated the emotional content of narrative productions, but not the physiologic orienting response or the early expressive behavior to negative stimuli. Furthermore, LIFG-cTBS disrupted the text-level structuring of negative picture storytelling and the early cardiac and muscular response to negative pictures; however, it did not affect the contextual emotional regulation of storytelling. These results may suggest that LIFG is involved in the initial detection of the affective arousal of emotional stimuli., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Semantic priming is a crucial phenomenon to study the organization of semantic memory. A novel type of priming effect, integrative priming, has been identified behaviorally, whereby a prime word facilitates recognition of a target word when the 2 concepts can be combined to form a unitary representation. We used both functional and anatomical imaging approaches to investigate the neural substrates supporting such integrative priming, and compare them with those in semantic priming. Similar behavioral priming effects for both semantic (Bread-Cake) and integrative conditions (Cherry-Cake) were observed when compared with an unrelated condition. However, a clearly dissociated brain response was observed between these 2 types of priming. The semantic-priming effect was localized to the posterior superior temporal and middle temporal gyrus. In contrast, the integrative-priming effect localized to the left anterior inferior frontal gyrus and left anterior temporal cortices. Furthermore, fiber tractography showed that the integrative-priming regions were connected via uncinate fasciculus fiber bundle forming an integrative circuit, whereas the semantic-priming regions connected to the posterior frontal cortex via separated pathways. The results point to dissociable neural pathways underlying the 2 distinct types of priming, illuminating the neural circuitry organization of semantic representation and integration., (© The Author 2015. Published by Oxford University Press. All rights reserved. For Permissions, please e-mail: journals.permissions@oup.com.)

Background: A hallmark symptom after psychological trauma is the presence of intrusive memories. It is unclear why only some moments of trauma become intrusive, and how these memories involuntarily return to mind. Understanding the neural mechanisms involved in the encoding and involuntary recall of intrusive memories may elucidate these questions., Method: Participants (n = 35) underwent functional magnetic resonance imaging (fMRI) while being exposed to traumatic film footage. After film viewing, participants indicated within the scanner, while undergoing fMRI, if they experienced an intrusive memory of the film. Further intrusive memories in daily life were recorded for 7 days. After 7 days, participants completed a recognition memory test. Intrusive memory encoding was captured by comparing activity at the time of viewing 'Intrusive scenes' (scenes recalled involuntarily), 'Control scenes' (scenes never recalled involuntarily) and 'Potential scenes' (scenes recalled involuntarily by others but not that individual). Signal change associated with intrusive memory involuntary recall was modelled using finite impulse response basis functions., Results: We found a widespread pattern of increased activation for Intrusive v. both Potential and Control scenes at encoding. The left inferior frontal gyrus and middle temporal gyrus showed increased activity in Intrusive scenes compared with Potential scenes, but not in Intrusive scenes compared with Control scenes. This pattern of activation persisted when taking recognition memory performance into account. Intrusive memory involuntary recall was characterized by activity in frontal regions, notably the left inferior frontal gyrus., Conclusions: The left inferior frontal gyrus may be implicated in both the encoding and involuntary recall of intrusive memories.

One of the most replicated findings in neurolinguistic literature on syntax is the increase of hemodynamic activity in the left inferior frontal gyrus (LIFG) in response to object relative (OR) clauses compared to subject relative clauses. However, behavioral studies have shown that ORs are primarily only costly when similarity-based interference is involved and recently, Leiken and Pylkkänen (2014) showed with magnetoencephalography (MEG) that an LIFG increase at an OR gap is also dependent on such interference. However, since ORs always involve a cue indicating an upcoming dependency formation, OR dependencies could be processed already prior to the gap-site and thus show no sheer dependency effects at the gap itself. To investigate the role of gap predictability in LIFG dependency effects, this MEG study compared ORs to verb phrase ellipsis (VPE), which was used as an example of a non-predictable dependency. Additionally, we explored LIFG sensitivity to filler-gap order by including right node raising structures, in which the order of filler and gap is reverse to that of ORs and VPE. Half of the stimuli invoked similarity-based interference and half did not. Our results demonstrate that LIFG effects of dependency can be elicited regardless of whether the dependency is predictable, the stimulus materials evoke similarity-based interference, or the filler precedes the gap. Thus, contrary to our own prior data, the current findings suggest a highly general role for the LIFG in dependency interpretation that is not limited to environments involving similarity-based interference. Additionally, the millisecond time-resolution of MEG allowed for a detailed characterization of the temporal profiles of LIFG dependency effects across our three constructions, revealing that the timing of these effects is somewhat construction-specific.

Prior research has shown that patients with damage to the left inferior frontal gyrus (LIFG) can have difficulties in executive control as well as understanding syntactic garden-paths, that is, sentences with a temporary syntactic ambiguity that resolve towards a less preferred interpretation. The present study tested two LIFG patients on object/subject garden path and matched syntactically unambiguous sentences. Besides syntactic ambiguity, support for the preferred but ultimately inappropriate interpretation was manipulated via verb bias, using verbs that were neutral between alternative analyses and verbs that were biased towards the context-inappropriate option. The LIFG patients, a non-LIFG patient, and healthy controls were tested on a sentence interpretation task (Experiment 1) and grammaticality judgement (Experiment 2). In contrast to the non-LIFG patient and controls, the LIFG patients showed impaired thematic role assignment across garden-path as well as unambiguous sentences, which tended to be worse with biased verbs. The results argue for a role of executive control in overcoming verb bias across diverse sentence processing situations, including, but not limited to, garden-path revision.

Tics in Gilles de la Tourette syndrome (GTS) resemble fragments of normal motor behaviour but appear in an intrusive, repetitive and context-inappropriate manner. Although tics can be voluntarily inhibited on demand, the neural correlates of this process remain unclear. 14 GTS adults without relevant comorbidities participated in this study. First, tic severity and voluntary tic inhibitory capacity were evaluated outside the scanner. Second, patients were examined with resting state functional magnetic resonance imaging (RS-fMRI) in two states, free ticcing and voluntary tic inhibition. Local synchronization of spontaneous fMRI-signal was analysed with regional homogeneity (ReHo) and differences between both states (free ticcing

Social information influences decision-making through an integration of information derived from individual experience with that derived from observing the actions of others. This raises the question as to which extent one should utilize social information. One strategy is to make use of uncertainty estimates, leading to a copy-when-uncertain strategy that weights information from individual and social sources based on their respective reliabilities. Here, we investigate this integration process by extending models of Bayes optimal integration of sensory information to a social decision context. We then use a key parameter of our behavioral model in conjunction with functional magnetic resonance imaging to identify the neural substrate that is specifically linked to the fidelity of this integration process. We show that individuals behave near Bayes optimal when integrating two distinct sources of social information but systematically deviate from Bayes optimal choice when integrating individual with social information. This systematic behavioral deviation from optimality is linked to activity of left inferior frontal gyrus. Thus, an ability to optimally exploit social information depends on processes that overcome an egocentric bias, and this regulatory role involves the left inferior prefrontal cortex. The findings provide a mechanistic explanation for observations wherein individuals neglect the benefits from exploiting social information., (© The Author (2013). Published by Oxford University Press.)

Although research on language production has developed detailed maps of the brain basis of single word production in both time and space, little is known about the spatiotemporal dynamics of the processes that combine individual words into larger representations during production. Studying composition in production is challenging due to difficulties both in controlling produced utterances and in measuring the associated brain responses. Here, we circumvent both problems using a minimal composition paradigm combined with the high temporal resolution of magnetoencephalography (MEG). With MEG, we measured the planning stages of simple adjective-noun phrases ('red tree'), matched list controls ('red, blue'), and individual nouns ('tree') and adjectives ('red'), with results indicating combinatorial processing in the ventro-medial prefrontal cortex (vmPFC) and left anterior temporal lobe (LATL), two regions previously implicated for the comprehension of similar phrases. These effects began relatively quickly (∼180 ms) after the presentation of a production prompt, suggesting that combination commences with initial lexical access. Further, while in comprehension, vmPFC effects have followed LATL effects, in this production paradigm vmPFC effects occurred mostly in parallel with LATL effects, suggesting that a late process in comprehension is an early process in production. Thus, our results provide a novel neural bridge between psycholinguistic models of comprehension and production that posit functionally similar combinatorial mechanisms operating in reversed order., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Accounts of the role of Broca's area in sentence comprehension range from specific syntactic operations to domain-general processes. The present study was designed to tease apart these two general accounts by measuring the BOLD response to two syntactically distinct long-distance dependencies that invoke abstractly similar predictive processes: backward anaphora and filler-gap dependencies. Previous research has observed distance effects in Broca's area for filler-gap dependencies, but not canonical anaphora, which has been interpreted in support of a syntactic movement account. However, filler-gap dependencies engage predictive mechanisms, resulting in active search for the gap, while canonical anaphora do not. Backward anaphora correct for this asymmetry as they engage a predictive mechanism that parallels the active search in filler-gap dependencies. The results revealed a distance effect in the pars triangularis of Broca's area for the backward anaphora condition, supporting a prediction-based role for this region rather than one for a particular syntactic operation., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Genetic studies found the A allele of the single nucleotide polymorphism rs1006737 in the CACNA1C gene, which encodes for the alpha 1C subunit of the voltage-dependent, L-type calcium ion channel Cav1.2, to be overrepresented in patients with major depressive disorder (MDD). Altered prefrontal brain functioning and impaired semantic verbal fluency (SVF) are robust findings in these patients. A recent functional magnetic resonance imaging (fMRI) study found the A allele to be associated with poorer performance and increased left inferior frontal gyrus (IFG) activation during SVF tasks in healthy subjects. In the present study, we investigated the effects of rs1006737 on neural processing during SVF in MDD. In response to semantic category cues, 40 patients with MDD and 40 matched controls overtly generated words while brain activity was measured with fMRI. As revealed by whole brain analyses, genotype significantly affected brain activity in patients. Compared to patients with GG genotype, patients with A allele demonstrated increased task-related activation in the left middle/inferior frontal gyrus and the bilateral cerebellum. Patients with A allele also showed enhanced functional coupling between left middle/inferior and right superior/middle frontal gyri. No differential effects of genotype on SVF performance or brain activation were found between diagnostic groups. The current data provide further evidence for an impact of rs1006737 on the left IFG and demonstrate that genetic variation in CACNA1C modulates neural responses in patients with MDD. The observed functional alterations in prefrontal and cerebellar areas might represent a mechanism by which rs1006737 influences susceptibility to MDD., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

In this fMRI study we investigate the neural correlates of information structure integration during sentence comprehension in Dutch. We looked into how prosodic cues (pitch accents) that signal the information status of constituents to the listener (new information) are combined with other types of information during the unification process. The difficulty of unifying the prosodic cues into overall sentence meaning was manipulated by constructing sentences in which the pitch accent did (focus-accent agreement), and sentences in which the pitch accent did not (focus-accent disagreement) match the expectations for focus constituents of the sentence. In case of a mismatch, the load on unification processes increases. Our results show two anatomically distinct effects of focus-accent disagreement, one located in the posterior left inferior frontal gyrus (LIFG, BA6/44), and one in the more anterior-ventral LIFG (BA 47/45). Our results confirm that information structure is taken into account during unification, and imply an important role for the LIFG in unification processes, in line with previous fMRI studies., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Verbal insight problem solving means to break mental sets, to select the novel semantic information and to form novel, task-related associations. Although previous studies have identified the brain regions associated with these key processes, the interaction among these regions during insight is still unclear. In the present study, we explored the functional connectivity between the key regions during solving Chinese 'chengyu' riddles by using event-related functional magnetic resonance imaging. Results showed that both insight and noninsight solutions activated the bilateral inferior frontal gyri, middle temporal gyri and hippocampi, and these regions constituted a frontal to temporal to hippocampal neural pathway. Compared with noninsight solution, insight solution had a stronger functional connectivity between the inferior frontal gyrus and middle temporal gyrus in the right hemisphere. Our study reveals the neural pathway of information processing during verbal insight problem solving, and supports the right-hemisphere advantage theory of insight., (Copyright © 2013 IBRO. Published by Elsevier Ltd. All rights reserved.)

Objective: Contrasts of verbal fluency and automatic speech provide an opportunity to evaluate the neural underpinnings of generativity and flexibility in autism spectrum disorders (ASD)., Method: We used functional magnetic resonance imaging (fMRI) to contrast brain activity in high functioning ASD (n=17, mean verbal IQ=117) and neurotypical (NT; n=20, mean verbal IQ=112) adolescent and young adult males (12-23years). Participants responded to three word generation conditions: automatic speech (reciting months), category fluency, and letter fluency., Results: Our paradigm closely mirrored behavioral fluency tasks by requiring overt, free recall word generation while controlling for differences in verbal output between the groups and systematically increasing the task demand. The ASD group showed reduced neural response compared to the NT participants during fluency tasks in multiple regions of left anterior and posterior cortices, and sub-cortical structures. Six of these regions fell in cortico-striatal circuits previously linked to repetitive behaviors (Langen, Durston, Kas, van Engeland, & Staal, 2011), and activity in two of them (putamen and thalamus) was negatively correlated with autism repetitive behavior symptoms in the ASD group. In addition, response in left inferior frontal gyrus was differentially modulated in the ASD, relative to the NT, group as a function of task demand., Conclusions: These data indicate a specific, atypical brain response in ASD to demanding generativity tasks that may have relevance to repetitive behavior symptoms in ASD as well as to difficulties generating original verbal responses., (Published by Elsevier Inc.)

According to the cue-based parsing approach (Lewis, Vasishth, & Van Dyke, 2006), sentence comprehension difficulty derives from interference from material that partially matches syntactic and semantic retrieval cues. In a 2 (low vs. high semantic interference)×2 (low vs. high syntactic interference) fMRI study, greater activation was observed in left BA44/45 for high versus low syntactic interference conditions following sentences and in left BA45/47 for high versus low semantic interference conditions following comprehension questions. A conjunction analysis showed BA45 associated with both types of interference, while BA47 was associated with only semantic interference. Greater activation was also observed in the left STG in the high interference conditions. Importantly, the results for the LIFG could not be attributed to greater working memory capacity demands for high interference conditions. The results favor a fractionation of the LIFG wherein BA45 is associated with post-retrieval selection and BA47 with controlled retrieval of semantic information., (Copyright © 2013 Elsevier Inc. All rights reserved.)

The proposal of a general conflict resolution mechanism in lIFG was investigated further in relation to language errors. In an fMRI study participants read sentences containing syntactic and plausibility violations. Furthermore, they were presented with sentences that were difficult to comprehend, due to degradation of the bottom-up signal (i.e., the visual form) of the language. We were interested whether comprehension difficulties caused by degradation would activate cognitive control mechanisms in the same manner as other language violations. To localize cognitive control processes participants performed a Stroop task. Both the violations and the visual degradation condition elicited co-localized lIFG activation with the Stroop conflict. These results indicate that lIFG implements control adjustments to resolve situations in which extra attention is needed more generally. Next to biasing attention to resolve representational conflicts arising from different types of errors, lIFG may also adjust control to compensate for a temporary lack of bottom-up information., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Although it is widely accepted that nouns and verbs are functionally independent linguistic entities, it is less clear whether their processing recruits different brain areas. This issue is particularly relevant for those theories of lexical semantics (and, more in general, of cognition) that suggest the embodiment of abstract concepts, i.e., based strongly on perceptual and motoric representations. This paper presents a formal meta-analysis of the neuroimaging evidence on noun and verb processing in order to address this dichotomy more effectively at the anatomical level. We used a hierarchical clustering algorithm that grouped fMRI/PET activation peaks solely on the basis of spatial proximity. Cluster specificity for grammatical class was then tested on the basis of the noun-verb distribution of the activation peaks included in each cluster. Thirty-two clusters were identified: three were associated with nouns across different tasks (in the right inferior temporal gyrus, the left angular gyrus, and the left inferior parietal gyrus); one with verbs across different tasks (in the posterior part of the right middle temporal gyrus); and three showed verb specificity in some tasks and noun specificity in others (in the left and right inferior frontal gyrus and the left insula). These results do not support the popular tenets that verb processing is predominantly based in the left frontal cortex and noun processing relies specifically on temporal regions; nor do they support the idea that verb lexical-semantic representations are heavily based on embodied motoric information. Our findings suggest instead that the cerebral circuits deputed to noun and verb processing lie in close spatial proximity in a wide network including frontal, parietal, and temporal regions. The data also indicate a predominant-but not exclusive-left lateralization of the network.

Inner speech involvement in self-reflection was examined by reviewing 130 studies assessing brain activation during self-referential processing in key self-domains: agency, self-recognition, emotions, personality traits, autobiographical memory, and miscellaneous (e.g., prospection, judgments). The left inferior frontal gyrus (LIFG) has been shown to be reliably recruited during inner speech production. The percentage of studies reporting LIFG activity for each self-dimension was calculated. Fifty five percent of all studies reviewed indicated LIFG (and presumably inner speech) activity during self-reflection tasks; on average LIFG activation is observed 16% of the time during completion of non-self tasks (e.g., attention, perception). The highest LIFG activation rate was observed during retrieval of autobiographical information. The LIFG was significantly more recruited during conceptual tasks (e.g., prospection, traits) than during perceptual tasks (agency and self-recognition). This constitutes additional evidence supporting the idea of a participation of inner speech in self-related thinking.

During times of emotional stress, individuals often engage in emotion regulation to reduce the experiential and physiological impact of negative emotions. Interestingly, emotion regulation strategies also influence memory encoding of the event. Cognitive reappraisal is associated with enhanced memory while expressive suppression is associated with impaired explicit memory of the emotional event. However, the mechanism by which these emotion regulation strategies affect memory is unclear. We used event-related fMRI to investigate the neural mechanisms that give rise to memory formation during emotion regulation. Twenty-five participants viewed negative pictures while alternately engaging in cognitive reappraisal, expressive suppression, or passive viewing. As part of the subsequent memory design, participants returned to the laboratory two weeks later for a surprise memory test. Behavioral results showed a reduction in negative affect and a retention advantage for reappraised stimuli relative to the other conditions. Imaging results showed that successful encoding during reappraisal was uniquely associated with greater co-activation of the left inferior frontal gyrus, amygdala, and hippocampus, suggesting a possible role for elaborative encoding of negative memories. This study provides neurobehavioral evidence that engaging in cognitive reappraisal is advantageous to both affective and mnemonic processes.