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Consonants and vowels differentially contribute to lexical acquisition. From 8 months on, infants' preferential reliance on consonants has been shown to predict their lexical outcome. Here, the predictive value of German-learning infants' (n = 58, 29 girls, 29 boys) trajectories of consonant and vowel perception, indicated by the electrophysiological mismatch response, across 2, 6, and 10 months for later lexical acquisition was studied. The consonant-perception trajectory from 2 to 6 months (β = −2.95) and 6 to 10 months (β = −.91), but not the vowel-perception trajectory, significantly predicted receptive vocabulary at 12 months. These results reveal an earlier predictive value of consonant perception for word learning than previously found, and a particular role of the longitudinal maturation of this skill in lexical acquisition.

Adult second language (L2) learning is a challenging enterprise inducing neuroplastic changes in the human brain. However, it remains unclear how the structural language connectome and its subnetworks change during adult L2-learning. The current study investigated longitudinal changes in white matter (WM) language networks in each hemisphere, as well as their interconnection, in a large group of Arabic-speaking adults who learned German intensively for six months. We found a significant increase in WM-connectivity within bilateral temporal-parietal semantic and phonological subnetworks and right temporal-frontal pathways mainly in the second half of the learning period. At the same time, WM-connectivity between the two hemispheres decreased significantly. Crucially, these changes in WM-connectivity are correlated with L2 performance. The observed changes in subnetworks of the two hemispheres suggest a network reconfiguration due to lexical learning. The reduced interhemispheric connectivity may indicate a key role of the corpus callosum in L2-learning by reducing the inhibition of the language-dominant left hemisphere. Our study highlights the dynamic changes within and across hemispheres in adult language-related networks driven by L2 learning.SignificanceThe neuroplastic changes induced by learning a second language (L2) in adulthood open up new perspectives for understanding brain function. The current study shows structural changes in the language network of Arabic native speakers who learned German intensively in two phases of three months each. We found a marked change in the left-hemispheric lexical-semantic language system and the right fronto-temporal pathway, accompanied by a decrease in white matter connectivity in the corpus callosum during L2 learning, which occurred mainly in the second period of L2 acquisition. The reduced interhemispheric connectivity suggests that the inhibitory role of the corpus callosum, relevant for native language processing, is reduced by L2 learning. Our findings demonstrate a clear experience-dependent structural plasticity in the human brain during L2 learning.

Is the neuroanatomy of the language structural connectome modulated by the life-long experience of speaking a specific language? The current study compared the brain white matter connections of the language and speech production network in a large cohort of 94 native speakers of two very different languages: an Indo-European morphosyntactically complex language (German) and a Semitic root-based language (Arabic). Using high-resolution diffusion-weighted MRI and tractography-based network statistics of the language connectome, we demonstrated that German native speakers exhibited stronger connectivity in an intra-hemispheric frontal to parietal/temporal dorsal language network, known to be associated with complex syntax processing. In comparison, Arabic native speakers showed stronger connectivity in the connections between semantic language regions, including the left temporo-parietal network, and stronger inter-hemispheric connections via the posterior corpus callosum connecting bilateral superior temporal and inferior parietal regions. The current study suggests that the structural language connectome develops and is modulated by environmental factors such as the characteristic processing demands of the native language.

Human language is supported by a cortical network involving Broca’s area which comprises Brodmann Areas 44 and 45 (BA44, BA45). While cytoarchitectonic homolog areas have been identified in nonhuman primates, it remains unknown how these regions evolved to support human language. Here, we use histological data and advanced cortical registration methods to precisely compare the morphology of BA44 and 45 between humans and chimpanzees. We found a general expansion of Broca’s areas in humans, with the left BA44 enlarging the most, growing anteriorly into a region known to process syntax. Together with recent functional studies, our findings show that BA44 evolved from a purely action-related region to a more expanded region in humans, with a posterior portion supporting action and an anterior portion supporting syntactic processes. Furthermore, our findings provide a solution for the longstanding debate concerning the structural and functional evolution of Broca’s area and its role in action and language.

Grammar learning requires memory for dependencies between nonadjacent elements in speech. Immediate learning of nonadjacent dependencies has been observed in very young infants, but their memory of such dependencies has remained unexplored. Here we used event-related potentials to investigate whether 6- to 8-month-olds retain nonadjacent dependencies and if sleep after learning affects this memory. Infants were familiarised with two rule-based morphosyntactic dependencies, presented in sentences of an unknown language. Brain responses after a retention period reveal memory of the nonadjacent dependencies, independent of whether infants napped or stayed awake. Napping, however, altered a specific processing stage, suggesting that memory evolves during sleep. Infants with high left frontal spindle activity show an additional brain response indicating memory of individual speech phrases. Results imply that infants as young as 6 months are equipped with memory mechanisms relevant to grammar learning. They also suggest that during sleep, consolidation of highly specific information can co-occur with changes in the nature of generalised memory.

The capacity for language is a defining property of our species, yet despite decades of research evidence on its neural basis is still mixed and a generalized consensus is difficult to achieve. We suggest that this is partly caused by researchers defining “language” in different ways, with focus on a wide range of phenomena, properties, and levels of investigation. Accordingly, there is very little agreement amongst cognitive neuroscientists of language on the operationalization of fundamental concepts to be investigated in neuroscientific experiments. Here, we review chains of derivation in the cognitive neuroscience of language, focusing on how the hypothesis under consideration is defined by a combination of theoretical and methodological assumptions. We first attempt to disentangle the complex relationship between linguistics, psychology, and neuroscience in the field. Next, we focus on how conclusions that can be drawn from any experiment are inherently constrained by auxiliary assumptions, both theoretical and methodological, on which the validity of conclusions drawn rests. These issues are discussed in the context of classical experimental manipulations as well as study designs that employ novel approaches such as naturalistic stimuli and computational modelling. We conclude by proposing that a highly interdisciplinary field such as the cognitive neuroscience of language requires researchers to form explicit statements concerning the theoretical definitions, methodological choices, and other constraining factors involved in their work.

Humans are equipped with the remarkable ability to comprehend an infinite number of utterances. Relations between grammatical categories restrict the way words combine into phrases and sentences. How the brain recognizes different word combinations remains largely unknown, although this is a necessary condition for combinatorial unboundedness in language. Here, we used functional magnetic resonance imaging and multivariate pattern analysis to explore whether distinct neural populations of a known language network hub—Broca’s area—are specialized for recognizing distinct simple word combinations. The phrases consisted of a noun (flag) occurring either with a content word, an adjective (green flag), or with a function word, a determiner (that flag). The key result is that the distribution of neural populations classifying word combination in Broca’s area seems sensitive to neuroanatomical subdivisions within this area, irrespective of task. The information patterns for adjective + noun were localized in its anterior part (BA45) whereas those for determiner + noun were localized in its posterior part (BA44). Our findings provide preliminary answers to the fundamental question of how lexical and grammatical category information interact during simple word combination, with the observation that Broca’s area is sensitive to the recognition of categorical relationships during combinatory processing, based on different demands placed on syntactic and semantic information. This supports the hypothesis that the combinatorial power of language consists of some neural computation capturing phrasal differences when processing linguistic input.

Individual differences in the ability to deal with language have long been discussed. The neural basis of these, however, is yet unknown. Here we investigated the relationship between long-range white matter connectivity of the brain, as revealed by diffusion tractography, and the ability to process syntactically complex sentences in the participants’ native language as well as the improvement thereof by multi-day training. We identified specific network motifs that indeed related white matter tractography to individual language processing performance. First, for two such motifs, one in the left and one in the right hemisphere, their individual prevalence significantly predicted the individual language performance suggesting a predisposition for the individual ability to process syntactically complex sentences, which manifests itself in the white matter brain structure. Both motifs comprise a number of cortical regions, but seem to be dominated by areas known for the involvement in working memory rather than the classical language network itself. Second, we identified another left hemispheric network motif, whose change of prevalence over the training period significantly correlated with the individual change in performance, thus reflecting training induced white matter plasticity. This motif comprises diverse cortical areas including regions known for their involvement in language processing, working memory and motor functions. The present findings suggest that individual differences in language processing and learning can be explained, in part, by individual differences in the brain’s white matter structure. Brain structure may be a crucial factor to be considered when discussing variations in human cognitive performance, more generally.

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.

Goal-directed behavior crucially relies on our capacity to suppress impulses and predominant behavioral responses. This ability, called inhibitory control, emerges in early childhood with marked improvements between 3 and 4 years. Here, we ask which brain structures are related to the emergence of this critical ability. Using a multimodal approach, we relate the pronounced behavioral improvements in different facets of 3-and 4-year-olds’ (N = 37, 20 female) inhibitory control to structural indices of maturation in the developing brain assessed with MRI. Our results show that cortical and subcortical structure of core regions in the adult cognitive control network, including the PFC, thalamus, and the inferior parietal cortices, are associated with early inhibitory functioning in preschool children. Probabilistic tractography revealed an association of frontoparietal (i.e., the superior longitudinal fascicle) and thalamocortical connections with early inhibitory control. Notably, these associations to brain structure were distinct for different facets of early inhibitory control, often referred to as motivational (‘hot’) and cognitive (‘cold’) inhibitory control. Our findings thus reveal the structural brain networks and connectivity related to the emergence of this core faculty of human cognition.Significance StatementThe capacity to suppress impulses and behavioral responses is crucial for goal-directed behavior. This ability, called inhibitory control, develops between the ages of 3 and 4 years. The factors behind this developmental milestone have been debated intensely for decades, however, the brain structure that underlies the emergence of inhibitory control in early childhood is largely unknown. Here, we relate the pronounced behavioral improvements in inhibitory control between 3 and 4 years with structural brain markers of grey matter and white matter maturation. Using a multimodal approach that combines analyses of cortical surface structure, subcortical structures, and white matter connectivity, our results reveal the structural brain networks and connectivity related to this core faculty of human cognition.

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., We demonstrate brain plasiticity to language training with a longitudinal auditory experiment during which native German speakers were trained to understand syntactically complex sentences. Based on MEG recordings, we identified a gamma power decrease in the left BA 44 for the more complex syntactic structures but not for the simpler ones. Over the training days, a correlation was observed between performance and gamma‐ based connectivity for pairwise connections from and to BA 44.

Grammar is central to any natural language. In the past decades, the artificial grammar of the AnBn 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 AnBn 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., Hierarchical syntactic processing in language is distinguished from associative processing. A novel artificial grammar is designed to guarantee hierarchical syntactic structure construction on the basis of category relations. Brodmann area (BA) 44 and the posterior superior temporal gyrus (pSTG) are specifically involved in hierarchical syntactic structure build‐up. Hierarchical syntactic processing engages a distinctive syntactic network including BA 44 and the pSTG.

Language comprehension depends on tight functional interactions between distributed brain regions. While these interactions are established for semantic and syntactic processes, the functional network of speech intonation – the linguistic variation of pitch – has been scarcely defined. Particularly little is known about intonation in tonal languages, in which pitch not only serves intonation but also expresses meaning via lexical tones. The present study used psychophysiological interaction analyses of functional magnetic resonance imaging data to characterise the neural networks underlying intonation and tone processing in native Mandarin Chinese speakers. Participants categorised either intonation or tone of monosyllabic Mandarin words that gradually varied between statement and question and between Tone 2 and Tone 4. Intonation processing induced bilateral fronto‐temporal activity and increased functional connectivity between left inferior frontal gyrus and bilateral temporal regions, likely linking auditory perception and labelling of intonation categories in a phonological network. Tone processing induced bilateral temporal activity, associated with the auditory representation of tonal (phonemic) categories. Together, the present data demonstrate the breadth of the functional intonation network in a tonal language including higher‐level phonological processes in addition to auditory representations common to both intonation and tone., The present functional magnetic resonance imaging study investigated task‐specific functional connectivity in intonation and tone processing in Mandarin speakers.Intonation processing increased functional interactions between left inferior frontal and bilateral temporal regions and between lateral‐medial frontal regions, likely reflecting higher‐level phonological processes and verbal response preparation.Tone processing only induced bilateral temporal activity, likely associated with auditory representation of tone (phonemic) categories.

Significance The ability to reason about other people’s thoughts and beliefs characterizes the complex social interaction among humans. This ability, called Theory of Mind (ToM), has long been argued to develop around 4 y when children start explicitly reasoning about others' beliefs. However, when tested nonverbally, infants already show action expectations congruent with others’ beliefs before the age of 2 y. Do these behaviors reflect different systems for understanding others’ minds—an early and a later developing one—or when does ToM develop? We show that these abilities are supported by the maturation of independent brain networks, suggesting different systems for explicit verbal ToM and early nonverbal action expectations., Human social interaction crucially relies on the ability to infer what other people think. Referred to as Theory of Mind (ToM), this ability has long been argued to emerge around 4 y of age when children start passing traditional verbal ToM tasks. This developmental dogma has recently been questioned by nonverbal ToM tasks passed by infants younger than 2 y of age. How do young children solve these tests, and what is their relation to the later-developing verbal ToM reasoning? Are there two different systems for nonverbal and verbal ToM, and when is the developmental onset of mature adult ToM? To address these questions, we related markers of cortical brain structure (i.e., cortical thickness and surface area) of 3- and 4-y-old children to their performance in novel nonverbal and traditional verbal TM tasks. We showed that verbal ToM reasoning was supported by cortical surface area and thickness of the precuneus and temporoparietal junction, classically involved in ToM in adults. Nonverbal ToM reasoning, in contrast, was supported by the cortical structure of a distinct and independent neural network including the supramarginal gyrus also involved in emotional and visual perspective taking, action observation, and social attention or encoding biases. This neural dissociation suggests two systems for reasoning about others’ minds—mature verbal ToM that emerges around 4 y of age, whereas nonverbal ToM tasks rely on different earlier-developing possibly social-cognitive processes.

In recent years a growing number of studies on syntactic processing has employed basic two-word constructions (e.g., “the tree”) to characterize the fundamental aspects of linguistic composition. This large body of evidence allows, for the first time, to closely examine which cognitive processes and neural substrates support the combination of two syntactic units into a more complex one, mirroring the nature of combinatory operations described in theoretical linguistics. The present review comprehensively examines behavioural, neuroimaging and neurostimulation studies investigating basic syntactic composition, covering more than 40 years of psycho- and neuro-linguistic research. Across several paradigms, four key features of syntactic composition have emerged: (1) the rule-based and (2) automatic nature of the combinatorial process, (3) a central role of Broca’s area and the posterior temporal lobe in representing and combining syntactic features, and (4) the reliance on efficient bottom-up integration rather than top-down prediction.