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Imaging and neurocognitive studies have searched for the brain areas involved in speech perception, specifically when speech is accompanied by noise, attempting to identify the underlying neural mechanism(s). Transcranial direct current stimulation (tDCS), a noninvasive, painless cortical neuromodulation technique, has been used to either excite or inhibit brain activity in order to better understand the neural mechanism underlying speech perception in noise. In the present study, anodal (excitatory) and cathodal (inhibitory) stimulations were performed on 48 participants, either over the left Inferior Frontal Gyrus (IFG), which includes Broca's area (n = 10 anodal, and n = 10 cathodal) or over the left Superior Temporal Gyrus (STG), which includes Wernicke's area (n = 13 anodal, n = 15 cathodal). Speech perception was measured using a sentence recognition task accompanied by white noise with a signal-to-noise ratio of -10 dB. Speech perception performance was measured four times: at baseline, after each of the two sessions of stimulation (one active and one sham session, the order of which was randomized between participants), and at a two-week follow-up session. Groups receiving anodal and cathodal stimulation over the left IFG did not show an effect of stimulation type. For groups receiving left STG stimulation, anodal stimulation resulted in higher scores, regardless of whether it was given before or after sham stimulation. However, cathodal stimulation showed an effect only when active stimulation was applied following sham stimulation. These results showed that tDCS had a direct effect on improving speech perception only over left STG. Furthermore, while anodal stimulation was effective in whatever order it was given, cathodal stimulation was effective only following sham stimulation, thereby allowing some amount of training. These findings carry both theoretical and clinical implications for the relationship between the DMN's left IFG and left STG areas during speech perception accompanied by background noise., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Language is bounded to the left hemisphere in the adult brain and the functional lateralization can already be observed early during development. Here we investigate whether this is paralleled by a lateralization of the white matter structural language network. We analyze the strength and microstructural properties of language-related fiber tracts connecting temporal and frontal cortices with a separation of two dorsal tracts, one targeting the posterior Broca's area (BA44) and one targeting the precentral gyrus (BA6). In a large sample of young children (3-6 years), we demonstrate that, in contrast to the BA6-targeting tract, the microstructural asymmetry of the BA44-targeting fiber tract significantly correlates locally with different aspects of development. While the asymmetry in its anterior segment reflects age, the asymmetry in its posterior segment is associated with the children's language skills. These findings demonstrate a fine-grained structure-to-function mapping in the lateralized network and go beyond our current view of language-related human brain maturation., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Language and complex tool use are often cited as behaviors unique to humans and may be evolutionarily linked owing to the underlying cognitive processes they have in common. We executed a quantitative activation likelihood estimation (ALE) meta-analysis (GingerALE 2.3) on published, whole-brain neuroimaging studies to identify areas associated with syntactic processing and/or tool use in humans. Significant clusters related to syntactic processing were identified in areas known to be related to language production and comprehension, including bilateral Broca's area in the inferior frontal gyrus. Tool use activation clusters were all in the left hemisphere and included the primary motor cortex and premotor cortex, in addition to other areas involved with sensorimotor transformation. Activation shared by syntactic processing and tool use was only significant at one cluster, located in the pars opercularis of the left inferior frontal gyrus. This minimal overlap between syntactic processing and tool use activation from our meta-analysis of neuroimaging studies indicates that there is not a widespread common neural network between the two. Broca's area may serve as an important hub that was initially recruited in early human evolution in the context of simple tool use, but was eventually co-opted for linguistic purposes, including the sequential and hierarchical ordering processes that characterize syntax. In the future, meta-analyses of additional components of language may allow for a more comprehensive examination of the functional networks that underlie the coevolution of human language and complex tool use., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Broca's area in the left hemisphere of the human neocortex has been suggested as a major hub for acquisition, storage, and access of linguistic information, abstract words in particular. Direct causal evidence for the latter, however, is still scarce; filling this gap was the goal of the present study. Using transcranial direct current stimulation (tDCS) of Broca's region, we aimed to delineate the involvement of this area in abstract and concrete word acquisition. The experiment used a between-subject design and involved 15 min of anodal or cathodal tDCS over Broca's area, or a sham/placebo control condition. The stimulation procedure was followed by a contextual learning session, in which participants were exposed to new concrete and abstract words embedded into short five-sentence texts. Finally, a set of behavioural assessment tasks was run to assess the learning outcomes immediately after the training (Day 1) and with a 24-hour delay (Day 2). The results showed that participants recognised novel abstract words more accurately after both anodal and cathodal tDCS in comparison with the sham condition on Day 1, which was also accompanied by longer recognition times (presumably due to deeper lexico-semantic processing), supporting the role of Broca's region in acquisition of abstract semantics. They were also more successful when recalling concrete words after cathodal tDCS, which indicates a degree of Broca's area involvement in forming memory circuits for concrete words as well. A decrease in the accuracy of recall of word forms and their meanings, as well as in recognition, was observed for all stimulation groups and both types of semantics on Day 2. The results suggest that both anodal and cathodal tDCS of Broca's area improves immediate contextual learning of novel vocabulary, predominantly affecting abstract semantics., (Copyright © 2022 Elsevier Inc. All rights reserved.)

An accumulating body of evidence suggests that transcranial direct current stimulation (tDCS) can be used to affect language processing, including word acquisition. There has been, however, no comprehensive study of effects of tDCS of the core language areas in relation to the main word-learning mechanisms. Two principal strategies have been posited as important for natural word acquisition: explicit encoding (EE) which relies on direct instructions and repetition of material, and fast mapping (FM) which operates implicitly, via context-based inference or deduction. We used anodal and cathodal tDCS of Broca's and Wernicke's areas to assess effects of stimulation site and polarity on novel word acquisition in both EE and FM regimes. 160 participants, divided into five groups, received 15 min of cathodal or anodal tDCS over one of the two areas or a sham (placebo) stimulation before learning eight novel words, presented ten times each in a short naturalistic audio-visual word-picture association session, fully counterbalanced across different learning regimes. The outcome of novel word acquisition was measured immediately after the training using a free recall task. The results showed elevated accuracy in all real stimulation groups in comparison with sham stimulation; however, this effect only reached full significance after anodal tDCS of Broca's area. Comparisons between the two learning modes indicated that Broca's anodal tDCS significantly improved both implicit and explicit acquisition of novel vocabulary in comparison with sham tDCS, without, however, any significant differences between EE and FM regimes as such. The results indicate involvement of the left inferior-frontal neocortex in the learning of novel vocabulary and suggest a possibility to promote different types of word acquisition using anodal tDCS of this area., (Copyright © 2022. Published by Elsevier Ltd.)

Manual gestures and speech recruit a common neural network, involving Broca's area in the left hemisphere. Such speech-gesture integration gave rise to theories on the critical role of manual gesturing in the origin of language. Within this evolutionary framework, research on gestural communication in our closer primate relatives has received renewed attention for investigating its potential language-like features. Here, using in vivo anatomical MRI in 50 baboons, we found that communicative gesturing is related to Broca homologue's marker in monkeys, namely the ventral portion of the Inferior Arcuate sulcus ( IA sulcus ). In fact, both direction and degree of gestural communication's handedness - but not handedness for object manipulation are associated and correlated with contralateral depth asymmetry at this exact IA sulcus portion. In other words, baboons that prefer to communicate with their right hand have a deeper left-than-right IA sulcus , than those preferring to communicate with their left hand and vice versa. Interestingly, in contrast to handedness for object manipulation, gestural communication's lateralisation is not associated to the Central sulcus depth asymmetry, suggesting a double dissociation of handedness' types between manipulative action and gestural communication. It is thus not excluded that this specific gestural lateralisation signature within the baboons' frontal cortex might reflect a phylogenetical continuity with language-related Broca lateralisation in humans., Competing Interests: YB, NC, KM, DM, MR, BN, JA, OC, AM No competing interests declared, (© 2022, Becker et al.)

During conversation, people take turns speaking by rapidly responding to their partners while simultaneously avoiding interruption 1,2 . Such interactions display a remarkable degree of coordination, as gaps between turns are typically about 200 milliseconds 3 -approximately the duration of an eyeblink 4 . These latencies are considerably shorter than those observed in simple word-production tasks, which indicates that speakers often plan their responses while listening to their partners 2 . Although a distributed network of brain regions has been implicated in speech planning 5-9 , the neural dynamics underlying the specific preparatory processes that enable rapid turn-taking are poorly understood. Here we use intracranial electrocorticography to precisely measure neural activity as participants perform interactive tasks, and we observe a functionally and anatomically distinct class of planning-related cortical dynamics. We localize these responses to a frontotemporal circuit centred on the language-critical caudal inferior frontal cortex 10 (Broca's region) and the caudal middle frontal gyrus-a region not normally implicated in speech planning 11-13 . Using a series of motor tasks, we then show that this planning network is more active when preparing speech as opposed to non-linguistic actions. Finally, we delineate planning-related circuitry during natural conversation that is nearly identical to the network mapped with our interactive tasks, and we find this circuit to be most active before participant speech during unconstrained turn-taking. Therefore, we have identified a speech planning network that is central to natural language generation during social interaction., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

To reveal the connectional specialization of the Broca's area (or its homologue), voxel-wise inter-species and individual differences, and inter-hemispheric asymmetry were respectively inspected in humans and macaques at both whole-brain connectivity and single tract levels. It was discovered that the developed connectivity blueprint approach is able to localize connectionally comparable voxels between the two species in Broca's area, whereas the quantitative differences between blueprints of locationally or connectionally corresponding voxels enable us to generate inter-hemispheric, inter-subject, and inter-species connectional variabilities, respectively. More importantly, the inter-species and inter-subject variabilities exhibited positive correlation in both two primates, and relatively higher variabilities were detected in the anatomically defined pars triangularis. By contrast, negative relationship was identified between the inter-species variability and hemispheric asymmetry in human brain. In particular, relatively higher asymmetry was revealed in the anatomically defined pars opercularis. Therefore, our novel findings demonstrated that pars triangularis, as compared to pars opercularis, might be a more active area during primate evolution, in which the brain connectivity and possible functions of pars triangularis show relatively higher degree in species specialization, yet lower in hemispheric specialization. Meanwhile, brain connectivity and possible functions of pars opercularis manifested an opposite pattern. At the tract level, functional roles related to the ventral stream in speech comprehension were relatively conservative and bilaterally organized, while those related to the dorsal stream in speech production show relatively higher species and hemispheric specializations., (Copyright © 2021. Published by Elsevier Inc.)

Analyzing the influence of emotion on false memory through electroencephalogram is helpful to further explore the cognition function of brain. In this study, material emotion and participant emotion were introduced into Deese-Roediger-McDermott (DRM) paradigm experiment at the same time, which made the experiment process closer to real life. Different music was used to induce the emotion of 28 participants, and they were divided into positive group and negative group. Then, we analyzed the difference between the two groups from the behavior data, source location and cortex functional network of event related potential. The results of difference analysis show that the false memory rate of positive group (85 ± 8.6%) is significantly higher than that of negative group (72 ± 9.7%), and the activation degree and voxel number of the positive group are significantly higher than those of the negative group in the brain regions related to semantic coding (BA24 and BA45), and the compactness, the speed of information transmission and the small-world property of the brain network in positive group are significantly higher than those in negative group. The above results show that in the positive group, more brain resources are used for semantic association when they judge keywords, which confuse the keywords and learning words, and result in more false memories., (Copyright © 2021 Elsevier B.V. All rights reserved.)

The adult human brain remains plastic even after puberty. However, whether first language (L1) training in adults can alter the language network is yet largely unknown. Thus, we conducted a longitudinal training experiment on syntactically complex German sentence comprehension. Sentence complexity was varied by the depth of the center embedded relative clauses (i.e., single or double embedded). Comprehension was tested after each sentence with a question on the thematic role assignment. Thirty adult, native German speakers were recruited for 4 days of training. Magnetoencephalography (MEG) data were recorded and subjected to spectral power analysis covering the classical frequency bands (i.e., theta, alpha, beta, low gamma, and gamma). Normalized spectral power, time-locked to the final closure of the relative clause, was subjected to a two-factor analysis ("sentence complexity" and "training days"). Results showed that for the more complex sentences, the interaction of sentence complexity and training days was observed in Brodmann area 44 (BA 44) as a decrease of gamma power with training. Moreover, in the gamma band (55-95 Hz) functional connectivity between BA 44 and other brain regions such as the inferior frontal sulcus and the inferior parietal cortex were correlated with behavioral performance increase due to training. These results show that even for native speakers, complex L1 sentence training improves language performance and alters neural activities of the left hemispheric language network. Training strengthens the use of the dorsal processing stream with working-memory-related brain regions for syntactically complex sentences, thereby demonstrating the brain's functional plasticity for L1 training., (© 2021 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Grammar is central to any natural language. In the past decades, the artificial grammar of the A n B n type in which a pair of associated elements can be nested in the other pair was considered as a desirable model to mimic human language syntax without semantic interference. However, such a grammar relies on mere associating mechanisms, thus insufficient to reflect the hierarchical nature of human syntax. Here, we test how the brain imposes syntactic hierarchies according to the category relations on linearized sequences by designing a novel artificial "Hierarchical syntactic structure-building Grammar" (HG), and compare this to the A n B n grammar as a "Nested associating Grammar" (NG) based on multilevel associations. Thirty-six healthy German native speakers were randomly assigned to one of the two grammars. Both groups performed a grammaticality judgment task on auditorily presented word sequences generated by the corresponding grammar in the scanner after a successful explicit behavioral learning session. Compared to the NG group, we found that the HG group showed a (a) significantly higher involvement of Brodmann area (BA) 44 in Broca's area and the posterior superior temporal gyrus (pSTG); and (b) qualitatively distinct connectivity between the two regions. Thus, the present study demonstrates that the build-up process of syntactic hierarchies on the basis of category relations critically relies on a distinctive left-hemispheric syntactic network involving BA 44 and pSTG. This indicates that our novel artificial grammar can constitute a suitable experimental tool to investigate syntax-specific processes in the human brain., (© 2021 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Humans are capable of acquiring multiple types of information presented in the same information stream. It has been suggested that at least two parallel learning processes are important during learning of sequential patterns-statistical learning and rule-based learning. Yet, the neurophysiological underpinnings of these parallel learning processes are not fully understood. To differentiate between the simultaneous mechanisms at the single trial level, we apply a temporal EEG signal decomposition approach together with sLORETA source localization method to delineate whether distinct statistical and rule-based learning codes can be distinguished in EEG data and can be related to distinct functional neuroanatomical structures. We demonstrate that concomitant but distinct aspects of information coded in the N2 time window play a role in these mechanisms: mismatch detection and response control underlie statistical learning and rule-based learning, respectively, albeit with different levels of time-sensitivity. Moreover, the effects of the two learning mechanisms in the different temporally decomposed clusters of neural activity also differed from each other in neural sources. Importantly, the right inferior frontal cortex (BA44) was specifically implicated in visuomotor statistical learning, confirming its role in the acquisition of transitional probabilities. In contrast, visuomotor rule-based learning was associated with the prefrontal gyrus (BA6). The results show how simultaneous learning mechanisms operate at the neurophysiological level and are orchestrated by distinct prefrontal cortical areas. The current findings deepen our understanding on the mechanisms of how humans are capable of learning multiple types of information from the same stimulus stream in a parallel fashion., (© 2021 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

Speaking in sentences requires selection from contextually determined lexical representations. Although posterior temporal cortex (PTC) and Broca's areas play important roles in storage and selection, respectively, of lexical representations, there has been no direct evidence for physiological interactions between these areas on time scales typical of lexical selection. Using intracranial recordings of cortical population activity indexed by high-gamma power (70-150 Hz) modulations, we studied the causal dynamics of cortical language networks while epilepsy surgery patients performed a sentence completion task in which the number of potential lexical responses was systematically varied. Prior to completion of sentences with more response possibilities, Broca's area was not only more active, but also exhibited more local network interactions with and greater top-down influences on PTC, consistent with activation of, and competition between, more lexical representations. These findings provide the most direct experimental support yet for network dynamics playing a role in lexical selection among competing alternatives during speech production., (© The Author(s) 2020. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Sign language (SL) conveys linguistic information using gestures instead of sounds. Here, we apply a meta-analytic estimation approach to neuroimaging studies (N = 23; subjects = 316) and ask whether SL comprehension in deaf signers relies on the same primarily left-hemispheric cortical network implicated in spoken and written language (SWL) comprehension in hearing speakers. We show that: (a) SL recruits bilateral fronto-temporo-occipital regions with strong left-lateralization in the posterior inferior frontal gyrus known as Broca's area, mirroring functional asymmetries observed for SWL. (b) Within this SL network, Broca's area constitutes a hub which attributes abstract linguistic information to gestures. (c) SL-specific voxels in Broca's area are also crucially involved in SWL, as confirmed by meta-analytic connectivity modeling using an independent large-scale neuroimaging database. This strongly suggests that the human brain evolved a lateralized language network with a supramodal hub in Broca's area which computes linguistic information independent of speech., (© 2020 The Authors. Human Brain Mapping published by Wiley Periodicals LLC.)

The ventrolateral frontal lobe (Broca's area) of the human brain is crucial in speech production. In macaques, neurons in the ventrolateral prefrontal cortex, the suggested monkey homologue of Broca's area, signal the volitional initiation of vocalizations. We explored whether this brain area became specialized for vocal initiation during primate evolution and trained macaques to alternate between a vocal and manual action in response to arbitrary cues. During task performance, single neurons recorded from the ventrolateral prefrontal cortex and the rostroventral premotor cortex of the inferior frontal cortex predominantly signaled the impending vocal or, to a lesser extent, manual action, but not both. Neuronal activity was specific for volitional action plans and differed during spontaneous movement preparations. This implies that the primate inferior frontal cortex controls the initiation of volitional utterances via a dedicated network of vocal selective neurons that might have been exploited during the evolution of Broca's area., Competing Interests: NG, AN No competing interests declared, (© 2021, Gavrilov and Nieder.)

Representations of sensory working memory can be found across the entire neocortex. But how are verbal working memory (VWM) contents retained in the human brain? Here we used fMRI and multi-voxel pattern analyses to study Chinese native speakers (15 males, 13 females) memorizing Chinese characters. Chinese characters are uniquely suitable to study VWM because verbal encoding is encouraged by their complex visual appearance and monosyllabic pronunciation. We found that activity patterns in Broca's area and left premotor cortex carried information about the memorized characters. These language-related areas carried (1) significantly more information about cued characters than those not cued for memorization, (2) significantly more information on the left than the right hemisphere and (3) significantly more information about Chinese symbols than complex visual patterns which are hard to verbalize. In contrast, early visual cortex carries a comparable amount of information about cued and uncued stimuli and is thus unlikely to be involved in memory retention. This study provides evidence for verbal working memory maintenance in a distributed network of language-related brain regions, consistent with distributed accounts of WM. The results also suggest that Broca's area and left premotor cortex form the articulatory network which serves articulatory rehearsal in the retention of verbal working memory contents., Competing Interests: Declaration of Competing Interest None., (Copyright © 2020. Published by Elsevier Inc.)

We have long known that language is lateralized to the left hemisphere (LH) in most neurologically healthy adults. In contrast, findings on lateralization of function during development are more complex. As in adults, anatomical, electrophysiological, and neuroimaging studies in infants and children indicate LH lateralization for language. However, in very young children, lesions to either hemisphere are equally likely to result in language deficits, suggesting that language is distributed symmetrically early in life. We address this apparent contradiction by examining patterns of functional MRI (fMRI) language activation in children (ages 4 through 13) and adults (ages 18 through 29). In contrast to previous studies, we focus not on lateralization per se but rather on patterns of left-hemisphere (LH) and right-hemisphere (RH) activation across individual participants over age. Our analyses show significant activation not only in the LH language network but also in their RH homologs in all of the youngest children (ages 4 through 6). The proportion of participants showing significant RH activation decreases over age, with over 60% of adults lacking any significant RH activation. A whole-brain correlation analysis revealed an age-related decrease in language activation only in the RH homolog of Broca's area. This correlation was independent of task difficulty. We conclude that, while language is left-lateralized throughout life, the RH contribution to language processing is also strong early in life and decreases through childhood. Importantly, this early RH language activation may represent a developmental mechanism for recovery following early LH injury., Competing Interests: The authors declare no competing interest.

When looking at a speaking person, the analysis of facial kinematics contributes to language discrimination and to the decoding of the time flow of visual speech. To disentangle these two factors, we investigated behavioural and fMRI responses to familiar and unfamiliar languages when observing speech gestures with natural or reversed kinematics. Twenty Italian volunteers viewed silent video-clips of speech shown as recorded (Forward, biological motion) or reversed in time (Backward, non-biological motion), in Italian (familiar language) or Arabic (non-familiar language). fMRI revealed that language (Italian/Arabic) and time-rendering (Forward/Backward) modulated distinct areas in the ventral occipito-temporal cortex, suggesting that visual speech analysis begins in this region, earlier than previously thought. Left premotor ventral (superior subdivision) and dorsal areas were preferentially activated with the familiar language independently of time-rendering, challenging the view that the role of these regions in speech processing is purely articulatory. The left premotor ventral region in the frontal operculum, thought to include part of the Broca's area, responded to the natural familiar language, consistent with the hypothesis of motor simulation of speech gestures., Competing Interests: The authors have declared that no competing interests exist.

Objective: The aim of our study was to determine any association between preservation of long latency response evoked by electrophysiological mapping of the caudal part of the pars opercularis (inferior frontal gyrus Broca area) and postoperative speech function after tumour removal in patients under general anesthesia., Patients and Methods: Twelve native Turkish-speaking patients with tumors in the dominant left frontal lobe, near the Broca area, were included in a single-center prospective cohort study. Hooked-wire electrodes were placed in both cricothyroid muscles (CTHY) before anesthesia and a monopolar electrode was used to stimulate the caudal portion of the pars opercularis of the inferior frontal gyrus before and after tumor removal. A long latency response (LLR) elicited at the contralateral (CTHY) muscle was interpreted as a positive stimulation of the Broca area. Patients received one pre-op and two post-op cognitive assessments. The Montreal Cognitive Assessment (MoCA) was used to assess global cognition and a "Cookie Theft" picture description task from the Boston Diagnostic Aphasia Examination test was used in assessing the language functions., Results: Electrical stimulation elicited a long latency response (LLR) in 9 (75 %) out of the 12 patients. The mean latency of the LLR was 50 ± 11 ms. Four (33.3 %) of the 12 patients showed temporary impairment in fluent speech postoperatively and all had full recovery during the 3-month follow-up period., Conclusions: The use of electrophysiological mapping methods by using EMG recording from laryngeal muscles may help to identify the opercular part of the Broca area under general anesthesia in order to preserve fluent speech functions., Competing Interests: Declaration of Competing Interest None of these authors has any conflict of interest to disclose., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Action is a cover term used to refer to a large set of motor processes differing in domain specificities (e.g. execution or observation). Here we review neuroimaging evidence on action processing (N = 416; Subjects = 5912) using quantitative Activation Likelihood Estimation (ALE) and Meta-Analytic Connectivity Modeling (MACM) approaches to delineate the functional specificities of six domains: (1) Action Execution, (2) Action Imitation, (3) Motor Imagery, (4) Action Observation, (5) Motor Learning, (6) Motor Preparation. Our results show distinct functional patterns for the different domains with convergence in posterior BA44 (pBA44) for execution, imitation and imagery processing. The functional connectivity network seeding in the motor-based localized cluster of pBA44 differs from the connectivity network seeding in the (language-related) anterior BA44. The two networks implement distinct cognitive functions. We propose that the motor-related network encompassing pBA44 is recruited when processing movements requiring a mental representation of the action itself., (Copyright © 2019. Published by Elsevier Inc.)

High-level cognitive capacities that serve communication, reasoning, and calculation are essential for finding our way in the world. But whether and to what extent these complex behaviors share the same neuronal substrate are still unresolved questions. The present study separated the aspects of logic from language and numerosity-mental faculties whose distinctness has been debated for centuries-and identified a new cytoarchitectonic area as correlate for an operation involving logical negation. A novel experimental paradigm that was implemented here in an RT/fMRI study showed a single cluster of activity that pertains to logical negation. It was distinct from clusters that were activated by numerical comparison and from the traditional language regions. The localization of this cluster was described by a newly identified cytoarchitectonic area in the left anterior insula, ventro-medial to Broca's region. We provide evidence for the congruence between the histologically and functionally defined regions on multiple measures. Its position in the left anterior insula suggests that it functions as a mediator between language and reasoning areas.

Current models of word-production in Broca's area (i.e. left ventro-lateral prefrontal cortex, VLPFC) posit that sequential and staggered semantic, lexical, phonological and articulatory processes precede articulation. Using millisecond-resolution intra-cranial recordings, we evaluated spatiotemporal dynamics and high frequency functional interconnectivity between left VLPFC regions during single-word production. Through the systematic variation of retrieval, selection, and phonological loads, we identified specific activation profiles and functional coupling patterns between these regions that fit within current psycholinguistic theories of word production. However, network interactions underpinning these processes activate in parallel (not sequentially), while the processes themselves are indexed by specific changes in network state. We found evidence that suggests that pars orbitalis is coupled with pars triangularis during lexical retrieval, while lexical selection is terminated via coupled activity with M1 at articulation onset. Taken together, this work reveals that speech production relies on very specific inter-regional couplings in rapid sequence in the language dominant hemisphere., Competing Interests: The authors have declared that no competing interests exist.

The superior temporal sulcus (STS) is an important region for speech comprehension. The greater language network is known to exhibit asymmetries in both structure and function, and consistent with that theory are reports of STS structural asymmetry in MRI-based, morphological measures such as mean thickness and sulcal depth. However, it is not known how these individual STS structural asymmetries relate to each other, or how they interact with the broader language asymmetry that manifests in other brain regions. In this study, we assess the interrelations of STS asymmetries in the human brain in vivo, using four independent datasets to validate our findings. For morphological measurements, we identify STS laterality effects consistent between our datasets and with the literature: leftward for surface area, and rightward for sulcal depth and mean thickness. We then add two more measurements of STS asymmetry: in T1, a quantitative index of the tissue's underlying biophysical properties; and in the projections to the STS from the arcuate fasciculus, a left-lateralized white-matter bundle that connects temporal regions (including STS) with frontal regions (including Broca's area). For these two new measurements, we identify no effect for T1 and a leftward effect for arcuate projections. We then test for correlations between these STS asymmetries, and find associations mainly between measurements of the same type (e.g., two morphological measurements). Finally, we ask if STS asymmetry is preferentially related to Broca asymmetry, as these are both important language regions and connected via the arcuate fasciculus. Using a linear model with cross-validation, we find that random regions are as successful as Broca's area in predicting STS, and no indication of a hypothesized leftward asymmetry. We conclude that although these different STS asymmetries are robust across datasets, they are not trivially related to each other, suggesting different biological or imaging sources for different aspects of STS lateralities.

The combination of transcranial magnetic stimulation and electroencephalography can be applied to probe effective connectivity. Neurons are excited by magnetic pulses, which produce transcranial magnetic stimulation-evoked potentials that can be monitored with electroencephalography. Effective connectivity refers to causal connections in the brain; it describes how different brain areas communicate with each other. Broca's area is crucial for all phases of speech processing and is located in the frontotemporal region of the cortex. Only a few studies have investigated this region using transcranial magnetic stimulation-electroencephalography because of the large cranial muscles that are located over these areas, resulting in large artifacts covering the transcranial magnetic stimulation-evoked potentials. However, it is shown that this obstacle can be overcome with new artifact-removal tools. We used minimum-norm estimation to locate the sources of the neuronal signals in electroencephalography data after stimulating the right-hemispheric homologue of Broca's area in three right-handed subjects; it was shown that the spreading of brain activity might be different for different individuals and that the brain activity spread fast to the contralateral hemisphere.

Transcranial direct current stimulation (tDCS) effects in cognition are inconsistent across studies. This study aimed to discuss why typical models might be insufficient to explain these effects, and to investigate a brain state factor, task load, with behavioral experiments on phonological processing. The motor theory of speech perception states that motor codes for articulation take part in speech perception, a view sharpened by neuroimaging findings, which show that the motor role in phonological processing is weighted by the nature of the tasks. Three groups of 20 participants, each under a different tDCS condition (anodal, cathodal, or sham), performed a categorical perception (CP), a lexical decision (LD), and a word naming (WN) task while stimulated on the pars opercularis of the left inferior frontal gyrus, a language area typically involved with the motor role. These tasks were assumed to be subserved by a network of nodes which included the target, believed to be increasingly relevant for performance from speech perception to speech production. A-tDCS facilitation and C-tDCS downregulation should directly increase with the relevance of the target for the task. Downregulation of a low relevance node could result in facilitation by compensation from other nodes. Overall, our brain stimulation findings support the neuroimaging literature in that motor participation in phonological processing depends on task nature and show that tDCS effects are modulated by task load relative to the target. Outcomes such as the improved performance following cathodal tDCS in CP and WN suggest that compensatory mechanisms may take place when the tasks involve more complex neuronal networks., (© 2019 Wiley Periodicals, Inc.)

Each brain hemisphere is dominant for certain functions such as speech. The determination of speech laterality prior to surgery is of paramount importance for accurate risk prediction. In this study, we aimed to determine speech laterality via EEG signals by using noninvasive machine learning techniques. The retrospective study included 67 subjects aged 18-65 years who had no chronic diseases and were diagnosed as healthy based on EEG examination. The subjects comprised 35 right-hand dominant (speech center located in the left hemisphere) and 32 left-hand dominant individuals (speech center located in the right hemisphere). A spectrogram was created for each of the 18 EEG channels by using various Convolutional Neural Networks (CNN) architectures including VGG16, VGG19, ResNet, MobileNet, NasNet, and DenseNet. These architectures were used to extract features from the spectrograms. The extracted features were classified using Support Vector Machines (SVM) and the classification performances of the CNN models were evaluated using Area Under the Curve (AUC). Of all the CNN models used in the study, VGG16 had a higher AUC value (0.83 ± 0.05) in the determination of speech laterality compared to all other models. The present study is a pioneer investigation into the determination of speech laterality via EEG signals with machine learning techniques, which, to our knowledge, has never been reported in the literature. Moreover, the classification results obtained in the study are promising and lead the way for subsequent studies though not practically feasible., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The localization of articulate language (speech) to the posterior third of the third left frontal convolution-Broca's area-did not occur to Broca as he reported the case of his first aphasic patient in 1861. Initially Broca localized articulate language to both frontal lobes, a position that he maintained for 4 years after publishing his first case. In the interval, the Academy of Medicine in Paris had received a copy of a paper authored in 1836 by Marc Dax, in which Dax claimed that the ability to speak resides within the left hemisphere alone. The Academy of Medicine convened in the spring of 1865 to adjudicate the issue. All of the distinguished speakers argued against Dax's contention by citing the prevailing paradigm, that bilaterally symmetrical organs, such as the eyes and ears, and the hemispheres of the brain, must perform the same function. The lone dissenting voice was that of Jules Baillarger, the discoverer of the laminar organization of the cerebral cortex, whose argument in favor of what he called "Dax's law" was so lucid that it carried the day. During his address to the Academy, Baillarger not only supported left-hemisphere dominance for speech, but for the first time described two forms of aphasia, fluent and nonfluent, now referred to as Wernicke's and Broca's aphasias, respectively, as well as the ability of aphasics to speak during emotional outbursts, to which we now refer as Baillarger-Jackson aphasia. It was 9 days after Baillarger's address that Broca, for the first time, unequivocally localized speech to the left frontal lobe.This paper is based on the author's reading of Dax's and Broca's original texts and of the texts read before the Academy of Medicine meeting held at the National Library of France between April 4, 1865, and June 13, 1865. From these primary sources it is concluded that the Academy of Medicine's debate was the last serious challenge to left-hemisphere dominance for speech and to the localization of articulate language to the left frontal lobe-and that Jules Baillarger played a pivotal role in what was a defining moment in neurobiology.

The sodium amytal test, or Wada test, named after Juhn Wada, has remained a pillar of presurgical planning and is used to identify the laterality of the dominant language and memory areas in the brain. What is perhaps less well known is that the original intent of the test was to abort seizure activity from an affected hemisphere and also to protect that hemisphere from the effects of electroconvulsive treatment. Some 80 years after Paul Broca described the frontal operculum as an essential area of expressive language and well before the age of MRI, Wada used the test to determine language dominance. The test was later adopted to study hemispheric memory dominance but was met with less consistent success because of the vascular anatomy of the mesial temporal structures. With the advent of functional MRI, the use of the Wada test has narrowed to application in select patients. The concept of selectively inhibiting part of the brain to determine its function, however, remains crucial to understanding brain function. In this review, the authors discuss the rise and fall of the Wada test, an important historical example of the innovation of clinicians in neuroscience.

Listening in a noisy environment is challenging for individuals with normal hearing and can be a significant burden for those with hearing impairment. The extent to which this burden is alleviated by a hearing device is a major, unresolved issue for rehabilitation. Here, we found adult users of cochlear implants (CIs) self-reported listening effort during a speech-in-noise task that was positively related to alpha oscillatory activity in the left inferior frontal cortex, canonical Broca's area, and inversely related to speech envelope coherence in the 2-5 Hz range originating in the superior-temporal plane encompassing auditory cortex. Left frontal cortex coherence in the 2-5 Hz range also predicted speech-in-noise identification. These data demonstrate that neural oscillations predict both speech perception ability in noise and listening effort.

Language is a highly important yet poorly understood function of the human brain. While studies of brain activation patterns during language comprehension are abundant, what is often critically missing is causal evidence of brain areas' involvement in a particular linguistic function, not least due to the unique human nature of this ability and a shortage of neurophysiological tools to study causal relationships in the human brain noninvasively. Recent years have seen a rapid rise in the use of transcranial direct current stimulation (tDCS) of the human brain, an easy, inexpensive and safe noninvasive technique that can modulate the state of the stimulated brain area (putatively by shifting excitation/inhibition thresholds), enabling a study of its particular contribution to specific functions. While mostly focusing on motor control, the use of tDCS is becoming more widespread in both basic and clinical research on higher cognitive functions, language included, but the procedures for its application remain variable. Here, we describe the use of tDCS in a psycholinguistic word-learning experiment. We present the techniques and procedures for application of cathodal and anodal stimulation of core language areas of Broca and Wernicke in the left hemisphere of the human brain, describe the procedures of creating balanced sets of psycholinguistic stimuli, a controlled yet naturalistic learning regime, and a comprehensive set of techniques to assess the learning outcomes and tDCS effects. As an example of tDCS application, we show that cathodal stimulation of Wernicke's area prior to a learning session can impact word learning efficiency. This impact is both present immediately after learning and, importantly, preserved over longer time after the physical effects of stimulation wear off, suggesting that tDCS can have long-term influence on linguistic storage and representations in the human brain.

Functional imaging data suggest different regions for semantic, syntactic, and phonological processing in an anterior-to-posterior direction along the inferior frontal gyrus. Language mapping by use of neuro-navigated transcranial magnetic stimulation (TMS) is frequently applied in clinical research to identify language-related cortical regions. Recently, we proposed a high spatial resolution approach for more detailed language mapping of cortical sub-areas such as Broca's region. Here, we employed a phonological picture-word interference paradigm in healthy subjects to reveal functional specialization in Broca's region for phonological processing. The behavioral phonological priming effect is characterized by accelerated naming responses to target pictures accompanied by phonologically related auditory distractor words. We hypothesized that the inhibitory effects of TMS on language processing would reduce phonological priming only at stimulation sites involved in phonological processing. In active as compared to sham TMS, we found reduced phonological facilitation specifically at sites overlapping with the probabilistic cytoarchitectonic area 44. Our findings complemented functional imaging data by revealing structure-function relationship in Broca's region. The introduction of a reaction time based interference paradigm into TMS language mapping increases the objectivity of the method and allows to explore functional specificity with high temporal resolution. Findings may help to interpret results in clinical applications.

The overall goal of this study is to investigate the role of parietal cortex in the control of walking in persons with Multiple Sclerosis (pwMS). We examined within-brain connectivity and cortico-muscular connectivity as pwMS and healthy control (HC) participants walked on an instrumented treadmill. Cortical activity was collected using EEG, muscle activity was collected using wireless EMG modules, and gait data were obtained by using the instrumented treadmill. Results show significant activation of sensorimotor and posterior parietal cortex during walking in both groups. Connectivity between parietal (posterior cingulate cortex PCC) and premotor regions (pars opercularis), and between PCC and contralateral muscles were higher in the healthy control group. Higher connectivity correlated with higher walking speed.

To evaluate transcranial direct current stimulation-induced changes in resting state quantitative EEG and cognitive-verbal performance of second language learners, 16 healthy individuals were randomly recruited to sham and real transcranial direct current stimulation groups receiving eight sessions of second language instruction accompanied by a 2 mA transcranial direct current stimulation over Broca's area with the cathode placed over the left arm. Quantitative EEG was recorded during the resting state after the stimulation session and second language instruction. Reduced theta activity at Fp1, F7, F3, and T5 caused by the stimulus current was reported. Multisession stimulation resulted in a significant increase in current density for beta power (25 Hz) in the language network. Cognitive-verbal pre-post stimulation performances suggest that anodal vs. sham transcranial direct current stimulation significantly improved the subjects test score on digit span, a cognitive-verbal ability. It is concluded that transcranial direct current stimulation of Broca's area increase cognitive-verbal performance by modulating brain electrical activity in language-related regions., Competing Interests: The authors declare no conflicts of interest statement., (©2019 Kheradmand Saadi et al. Published by IMR press. All rights reserved.)

Background: Repetitive transcranial magnetic stimulation (rTMS) has been used experimentally to facilitate naming abilities in individuals with chronic post-stroke aphasia. However, little is known about how rTMS confers clinical improvement, hampering its therapeutic value. The present study investigated the characteristics of naming failure that improve following administration of continuous theta burst stimulation (cTBS)-an inhibitory form of rTMS-to the right pars triangularis (rPTr) in persons with chronic aphasia., Methods: Eleven participants with chronic aphasia following left hemisphere stroke named pictures prior to and immediately following cTBS of the rPTr and a control site (vertex) in separate sessions. Prior to stimulation, we obtained two baseline measurements of picture naming ability to determine the extent and type (i.e., phonological vs. semantic) of naming impairment. Items presented for naming during stimulation were those that were named incorrectly in one or both of the baseline sessions (i.e., inconsistent vs. wrong items, respectively). Analyses assessed whether cTBS effects differed depending on the severity and/or type of naming impairment., Results: Relative to vertex, cTBS of the rPTr improved naming of inconsistent, but not wrong, items for individuals with more severe baseline naming impairment. Critically, baseline phonological but not semantic naming impairment severity marginally correlated with improved accuracy overall, and significantly correlated with decreased phonological errors following rPTr stimulation., Conclusion: CTBS of the rPTr enhances naming by facilitating phonological access during word retrieval, indicating that individuals whose naming impairment is localized to this stage of processing may be most likely to benefit from this rTMS approach., (Copyright © 2019 Elsevier Inc. All rights reserved.)

The basic steps in building up language involve binding words of different categories into a hierarchical structure. To what extent these steps are universal or differ across languages is an open issue. Here we examine the neural dynamics of phrase structure building in Chinese-a language that in contrast to other languages heavily depends on contextual semantic information. We used functional magnetic resonance imaging and dynamic causal modeling to identify the relevant brain regions and their dynamic relations. Language stimuli consisted of syntax-driving determiners, semantics-embedded classifiers, and nonverbal symbols making up for two-component sequences manipulated by the factors structure (phrase/list) and number of words (2-word/1-word). Processing phrases compared with word lists elicited greater activation in the anterior part of Broca's area, Brodmann area (BA) 45, and the left posterior superior/middle temporal gyri (pSTG/pMTG), while processing two words against one word led to stronger involvement of the left BA 45, BA 44, and insula. Differential network modulations emerging from subparts of Broca's area revealed that phrasal construction in particular highly modulated the direct connection from BA 44 to left pMTG, suggesting BA 44's primary role in phrase structure building. Conversely, the involvement of BA 45 rather appears sensitive to the reliance on lexico-semantic information in Chinese. Against the background of previous findings from other languages, the present results indicate that phrase structure building has a universal neural basis within the left fronto-temporal network. Most importantly, they provide the first evidence demonstrating that the structure-building network may be modulated by language-specific characteristics., (© 2018 Wiley Periodicals, Inc.)

Purpose: The present study aimed to explore the short-term effect of anodal transcranial direct current stimulation (tDCS) on tongue twister production., Method: Thirty healthy native Cantonese adult speakers were randomly assigned to the anodal tDCS group or the sham tDCS group. Anodal tDCS of 2 mA was applied over the Broca's area of the brain. The stimulation lasted for 20 min for the anodal tDCS group and 30 s for the sham tDCS group. The participants were instructed to produce a list of tongue twisters before, immediately after and 4 h after tDCS., Result: Speech rate and response accuracy measured immediately after stimulation were significantly faster and higher, respectively, than before stimulation. Although there was no change in speech rate measured at 4 h after stimulation, response accuracy at that time point was significantly lower than that measured immediately after stimulation. However, there were no significant differences between the anodal tDCS and sham tDCS groups in either speech rate or response accuracy., Conclusion: The findings revealed that a single session of anodal tDCS over the Broca's area did not significantly improve speech production during tongue twister production.

Musical improvisation is one of the most complex forms of creative behavior, which offers a realistic task paradigm for the investigation of real-time creativity where revision is not possible. Despite some previous studies on musical improvisation and brain activity, what and how brain areas are involved during musical improvisation are not clearly understood. In this article, we designed a new functional magnetic resonance imaging (fMRI) study, in which, while being in the MRI scanner, advanced jazz improvisers performed improvisatory vocalization and imagery as main tasks and performed a prelearned melody as a control task. We incorporated a musical imagery task to avoid possible confounds of mixed motor and perceptual variables in previous studies. We found that musical improvisation compared with prelearned melody is characterized by higher node activity in the Broca's area, dorsolateral prefrontal cortex, lateral premotor cortex, supplementary motor area and cerebellum, and lower functional connectivity in number and strength among these regions. We discuss various explanations for the divergent activation and connectivity results. These results point to the notion that a human creative behavior performed under real-time constraints is an internally directed behavior controlled primarily by a smaller brain network in the frontal cortex.

The ability to speak is a unique human capacity, but where is it located in our brains? This question is closely connected to the pioneering work of Pierre Paul Broca in the 1860s. Based on post-mortem observations of aphasic patients' brains, Broca located language production in the 3rd convolution of the left frontal lobe and thus reinitiated the localizationist view of brain functions. However, contemporary neuroscience has partially rejected this view in favor of a network-based perspective. This leads to the question, whether Broca's findings are still relevant today. In this mini-review, we discuss current and historical implications of Broca's work by focusing on his original contribution and contrasting it with contemporary knowledge. Borrowing from Broca's famous quote, our review shows that humans indeed "speak with the left hemisphere"- but Broca's area is not the sole "seat of articulatory language".

The ability to use word category information (WCI) for syntactic structure building has been hypothesized to be the essence of human language faculty. The neural substrate of the ability of using the WCI for the complex syntactic hierarchical structure processing, however, is yet unknown. Therefore, we directly conducted an fMRI experiment by using a pseudo-Chinese artificial language with syntactic structures containing a center-embedded relative clause. Thirty non-Chinese native (Korean) speakers were randomly divided into two groups: one acquired WCI and WCI-based syntactic rules (the WCI group) before the scanning session, and the other did not (the non-WCI group). Both groups were required to judge the grammaticality of the testing sentences, with critical long-distance dependencies between two elements (the main verb and the relativizer). Behaviorally, the WCI group's accuracy was significantly higher and its reaction time was shorter. The scanning results showed that the left superior temporal gyrus (STG) and Broca's area were more strongly activated for the WCI group, and the dynamic causal modeling analyses revealed a distinct effective connectivity pattern for this group. Therefore, the present research, for the first time, reveals that the activation of and the functional connectivity between Broca's area and the left STG play a critical role in the ability of the rule-based use of the WCI which is crucial for complex hierarchical structure building, and might be substantially corresponding to the "labeling competence" within the linguistic framework., (© 2018 Wiley Periodicals, Inc.)

The distinction between nouns and verbs is a language universal. Yet, functional neuroimaging studies comparing noun and verb processing have yielded inconsistent findings, ranging from a complete frontal(verb)-temporal(noun) dichotomy to a complete overlap in activation patterns. The current study addressed the debate about neural distinctions between nouns and verbs by conducting an activation likelihood estimation (ALE) meta-analysis of probabilistic cytoarchitectonic maps. Two levels of analysis were conducted: simple effects (Verbs vs. Baseline, Nouns vs. Baseline), and direct comparisons (Verbs vs. Nouns, Nouns vs. Verbs). Nouns were uniquely associated with a left medial temporal cluster (BA37). Activation foci for verbs included extensive inferior frontal (BA44-47) and mid-temporal (BA22, 21) regions in the left hemisphere. These findings confirm that the two grammatical classes have distinct neural architecture in supra-modal brain regions. Further, nouns and verbs overlapped in a small left lateral inferior temporal activation cluster (BA37), which is a region for modality-independent, grammatical class-independent lexical representations. These findings are most consistent with the view that as one acquires language, linguistic representations for a lexical category shift from the modality specific cortices which represent prototypical members of that category (e.g., motion for verbs) to abstract amodal representations in close proximity to modality specific cortices., (© 2018 Wiley Periodicals, Inc.)

Recent studies have shown that the left inferior frontal gyrus (IFG) plays a key role in language learning. Facilitatory stimulation over this region by means of anodal transcranial direct current stimulation (tDCS) can modulate linguistic abilities in healthy individuals and improve language performance in patients with post-stroke aphasia. Neuroimaging studies in healthy participants have suggested that anodal tDCS decreases task-related activity at the stimulated site when applied during different language tasks, and changes resting-state connectivity in a larger network of areas associated with language processing. However, to date, the neural correlates of the potential beneficial effects of tDCS on verb learning remain unclear. The current study investigated how anodal tDCS during verb learning modulates task-related activity and effective connectivity in the healthy language network. To this end, we combined a verb learning paradigm during functional neuroimaging with simultaneous tDCS over the left IFG in healthy human volunteers. We found that, relative to sham stimulation, anodal tDCS significantly decreased task-related activity at the stimulated left IFG and in the right homologue. Effective connectivity analysis showed that anodal tDCS significantly decreased task-related functional coupling between the left IFG and the right insula. Importantly, the individual decrease in connectivity was significantly correlated with the individual behavioural improvement during anodal tDCS. These results demonstrate, for the first time, that the behavioural improvements induced by anodal tDCS might be related to an overall decrease in processing effort both with respect to task-related activity and effective connectivity within a large language network., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Patient studies and brain stimulation evidence suggest that language processing can be enhanced by altering the interhemispheric balance: namely, preferentially enhancing left-hemisphere activity while suppressing right-hemisphere activity. To our knowledge, no study has yet compared the effects of such bilateral brain stimulation to both logically necessary control conditions (separate left- and right-hemisphere stimulation). This study did so in a between-group sham-controlled design, applying transcranial direct current stimulation over Broca's area and/or its homologue in 72 healthy participants. The effects were measured not only in a single-word-level task but also in a sentence-level task, rarely tested previously. We did not find either any significant overall effects of stimulation or greater stimulation effects in the bilateral compared to control groups. This null result, obtained in a large sample, contributes to the debate on whether tDCS can modulate language processing in healthy individuals., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Communication is an inferential process. In particular, language comprehension constantly requires top-down efforts, as often multiple interpretations are compatible with a given sentence. To assess top-down processing in the language domain, our experiment employed ambiguous sentences that allow for multiple interpretations (e.g., The client sued the murderer with the corrupt lawyer., where the corrupt lawyer could either belong to The client or the murderer). Interpretation thus depended on whether participants chunk the words of the sentence into short or long syntactic phrases. In principle, bottom-up acoustic information (i.e., the presence or absence of an intonational phrase boundary at the offset of the murderer) indicates one of the two possible interpretations. Yet, acoustic information often indicates interpretations that require words to be chunked into overly long phrases that would overburden working memory. Processing is biased against these demands, reflected in a top-down preference to chunk words into short rather than long phrases. It is often proposed, but also hotly debated, that the ability to chunk words into short phrases is subserved by the left inferior frontal gyrus (IFG). Here, we employed focal repetitive transcranial magnetic stimulation to perturb the left IFG, which resulted in a further decrease of the aptitude to tolerate long phrases, indicating the inability of the left IFG to assist the chunking of words into phrases. In contrast, the processing of auditory information was not affected. Our findings support a causal top-down role of the left inferior frontal gyrus in the chunking of words into phrases., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Listeners are constantly trying to predict what the speaker will say next. We concurrently measured the electrophysiological and haemodynamic correlates of syntactic pre-activation, investigating when and where the brain processes speech melody cues to upcoming word order structure. Pre-activation of syntactic structure was reflected in a left-lateralised pre-activation negativity (PrAN), which was subserved by Broca's area in the left inferior frontal gyrus, as well as the contiguous left anterior insula., (Copyright © 2018 The Authors. Published by Elsevier B.V. All rights reserved.)

It is clear that the left inferior frontal gyrus (LIFG) contributes in some fashion to sentence processing. While neuroimaging and neuropsychological evidence support a domain-general working memory function, recent neuroimaging data show that particular subregions of the LIFG, particularly the pars triangularis (pTri), show selective activation for sentences relative to verbal working memory and cognitive control tasks. These data suggest a language-specific function rather than a domain-general one. To resolve this apparent conflict, I propose separating claims of domain-generality and specificity independently for computations and representations-a given brain region may respond to a specific representation while performing a general computation over that representation, one shared with other systems. I hypothesize that the pTri underlies a language-specific working memory system, comprised of general memory retrieval/attention operations specialized for syntactic representations. There is a parallelism of top-down retrieval function among the phonological and semantic levels, localized to the pars opercularis and pars orbitalis, respectively. I further explore the idea of how such a system emerges in the human brain through the framework of neuronal retuning: the "borrowing" of domain-general mechanisms for language, either in evolution or development. The empirical data appear to tentatively support a developmental account of language-specificity in the pTri, possibly through connections to the posterior superior temporal sulcus (pSTS), a region that is both anatomically distinct for humans and functionally essential for language. Evidence of representational response specificity obtained from neuroimaging studies is useful in understanding how cognition is implemented in the brain. However, understanding the shared computations across domains and neural systems is necessary for a fuller understanding of this problem, providing potential answers to questions of how specialized systems, such as language, are implemented in the brain.

The Persian language can be considered to have a relatively more complex and combinatorial morpho-syntax than languages like Chinese and English. For example, the Persian verbal system is largely constituted of light verb constructions, in which light verbs are combined with specific items coming from other grammatical classes to generate entirely new verbal entities. This study was designed to examine the mediating effect of language-inherent properties related to morpho-syntax on activation of the left inferior frontal gyrus (LIFG), a brain area involved in morpho-syntactic processing. To this end, 20 late Persian-English bilinguals were required to covertly generate verbs and nouns from object and action pictures, within a cued grammatical context. Consistent with predictions, the results of an ROI analysis revealed an interaction between task and language in BA 44 of the LIFG and its right homologue, with greater activation of this region during the production of Persian compared to English verbs. In contrast, there was greater activation of the BA 44 during the production of English compared to Persian nouns, consistent with the more effortful processing of their less proficient second language (English). The findings suggest that language-specific properties such as morpho-syntactic complexity can modulate the recruitment of Broca's area, over and above the more well-documented effects of language proficiency., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Objective: In this study, electroencephalography data of imagined words were classified using four different feature extraction approaches. Eight subjects were recruited for the recording of imagination with five different words, namely; 'go', 'back', 'left', 'right', and 'stop'.

Language acquisition theories classically distinguish passive language understanding from active language production. However, recent findings show that brain areas such as Broca's region are shared in language understanding and production. Furthermore, these areas are also implicated in understanding and producing goal-oriented actions. These observations question the passive view of language development. In this work, we propose a cognitive developmental model of symbol acquisition, coherent with an active view of language learning. For that purpose, we introduce the concept of social babbling. In this view, symbols are learned in the same way as goal-oriented actions in the context of specific caregiver-infant interactions. We show that this model allows a virtual agent to learn both symbolic words and gestures to refer to objects while interacting with a caregiver. We validate our model by reproducing results from studies on the influence of parental responsiveness on infants language acquisition., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The ability to process structured sequences is a central feature of natural language but also characterizes many other domains of human cognition. In this fMRI study, we measured brain metabolic response in musicians as they generated structured and non-structured sequences in language and music. We employed a univariate and multivariate cross-classification approach to provide evidence that a common neural code underlies the production of structured sequences across the two domains. Crucially, the common substrate includes Broca's area, a region well known for processing structured sequences in language. These findings have several implications. First, they directly support the hypothesis that language and music share syntactic integration mechanisms. Second, they show that Broca's area is capable of operating supramodally across these two domains. Finally, these results dismiss the recent hypothesis that domain general processes of neighboring neural substrates explain the previously observed "overlap" between neuroimaging activations across the two domains., (Copyright © 2018 Elsevier Inc. All rights reserved.)

People suffering from neuromuscular disorders such as locked-in syndrome (LIS) are left in a paralyzed state with preserved awareness and cognition. In this study, it was hypothesized that changes in local hemodynamic activity, due to the activation of Broca's area during overt/covert speech, can be harnessed to create an intuitive Brain Computer Interface based on Near-Infrared Spectroscopy (NIRS). A 12-channel square template was used to cover inferior frontal gyrus and changes in hemoglobin concentration corresponding to six aloud (overtly) and six silently (covertly) spoken words were collected from eight healthy participants. An unsupervised feature extraction algorithm was implemented with an optimized support vector machine for classification. For all participants, when considering overt and covert classes regardless of words, classification accuracy of 92.88 ± 18.49% was achieved with oxy-hemoglobin (O2Hb) and 95.14 ± 5.39% with deoxy-hemoglobin (HHb) as a chromophore. For a six-active-class problem of overtly spoken words, 88.19 ± 7.12% accuracy was achieved for O2Hb and 78.82 ± 15.76% for HHb. Similarly, for a six-active-class classification of covertly spoken words, 79.17 ± 14.30% accuracy was achieved with O2Hb and 86.81 ± 9.90% with HHb as an absorber. These results indicate that a control paradigm based on covert speech can be reliably implemented into future Brain⁻Computer Interfaces (BCIs) based on NIRS.