Lateralization of dorsal fiber tract targeting Broca's area concurs with language skills during development.

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.

Coevolution of language and tools in the human brain: An ALE meta-analysis of neural activation during syntactic processing and tool use.

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.

Broca's area involvement in abstract and concrete word acquisition: tDCS evidence.

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.

Broca's cerebral asymmetry reflects gestural communication's lateralisation in monkeys (Papio anubis).

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.

A speech planning network for interactive language use.

During conversation, people take turns speaking by rapidly responding to their partners while simultaneously avoiding interruption1,2. Such interactions display a remarkable degree of coordination, as gaps between turns are typically about 200 milliseconds3-approximately the duration of an eyeblink4. 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 partners2. Although a distributed network of brain regions has been implicated in speech planning5-9, 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 cortex10 (Broca's region) and the caudal middle frontal gyrus-a region not normally implicated in speech planning11-13. 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.

Anodal tDCS over Broca's area improves fast mapping and explicit encoding of novel vocabulary.

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.

The influence of positive emotion and negative emotion on false memory based on EEG signal analysis.

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.

Species and individual differences and connectional asymmetry of Broca's area in humans and macaques.

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.

Functional brain plasticity during L1 training on complex sentences: Changes in gamma-band oscillatory activity.

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.

Neurophysiological and functional neuroanatomical coding of statistical and deterministic rule information during sequence learning.

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.

Hierarchical syntactic processing is beyond mere associating: Functional magnetic resonance imaging evidence from a novel artificial grammar.

Grammar is central to any natural language. In the past decades, the artificial grammar of the An Bn 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 An Bn 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.

Distinct neural networks for the volitional control of vocal and manual actions in the monkey homologue of Broca's area.

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.

Functional neuroanatomy of language without speech: An ALE meta-analysis of sign language.

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.

The Dynamics of Language Network Interactions in Lexical Selection: An Intracranial EEG Study.

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.

Decoding verbal working memory representations of Chinese characters from Broca's area.

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.

The neural basis of language development: Changes in lateralization over age.

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.

Sensitivity of occipito-temporal cortex, premotor and Broca's areas to visible speech gestures in a familiar language.

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.

Intraoperative motor speech mapping under general anesthesia using long-latency response from laryngeal muscles.

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.

The topographical organization of motor processing: An ALE meta-analysis on six action domains and the relevance of Broca's region.

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.

Logical negation mapped onto the brain.

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.