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Contains fulltext : 115434.pdf (Publisher’s version ) (Closed access) Repetition suppression in fMRI studies is generally thought to underlie behavioural facilitation effects (i.e., priming) and it is often used to identify the neuronal representations associated with a stimulus. However, this pays little heed to the large number of repetition enhancement effects observed under similar conditions. In this review, we identify several cognitive variables biasing repetition effects in the BOLD response towards enhancement instead of suppression. These variables are stimulus recognition, learning, attention, expectation and explicit memory. We also evaluate which models can account for these repetition effects and come to the conclusion that there is no one single model that is able to embrace all repetition enhancement effects. Accumulation, novel network formation as well as predictive coding models can all explain subsets of repetition enhancement effects.

The human capacity to acquire language is an outstanding scientific challenge to understand. Somehow our language capacities arise from the way the human brain processes, develops and learns in interaction with its environment. To set the stage, we begin with a summary of what is known about the neural organization of language and what our artificial grammar learning (AGL) studies have revealed. We then review the Chomsky hierarchy in the context of the theory of computation and formal learning theory. Finally, we outline a neurobiological model of language acquisition and processing based on an adaptive, recurrent, spiking network architecture. This architecture implements an asynchronous, event-driven, parallel system for recursive processing. We conclude that the brain represents grammars (or more precisely, the parser/generator) in its connectivity, and its ability for syntax is based on neurobiological infrastructure for structured sequence processing. The acquisition of this ability is accounted for in an adaptive dynamical systems framework. Artificial language learning (ALL) paradigms might be used to study the acquisition process within such a framework, as well as the processing properties of the underlying neurobiological infrastructure. However, it is necessary to combine and constrain the interpretation of ALL results by theoretical models and empirical studies on natural language processing. Given that the faculty of language is captured by classical computational models to a significant extent, and that these can be embedded in dynamic network architectures, there is hope that significant progress can be made in understanding the neurobiology of the language faculty. Max Planck Institute for Psycholinguistics; Donders Institute for Brain, Cognition and Behaviour; Fundacao para a Ciencia e Tecnologia (IBB/CBME, LA, FEDER/POCI) [PTDC/PSI-PCO/110734/2009]; Vetenskapsradet info:eu-repo/semantics/publishedVersion

In this paper we examine the neurobiological correlates of syntax, the processing of structured sequences, by comparing FMRI results on artificial and natural language syntax. We discuss these and similar findings in the context of formal language and computability theory. We used a simple right-linear unification grammar in an implicit artificial grammar learning paradigm in 32 healthy Dutch university students (natural language FMRI data were already acquired for these participants). We predicted that artificial syntax processing would engage the left inferior frontal region (BA 44/45) and that this activation would overlap with syntax-related variability observed in the natural language experiment. The main findings of this study show that the left inferior frontal region centered on BA 44/45 is active during artificial syntax processing of well-formed (grammatical) sequence independent of local subsequence familiarity. The same region is engaged to a greater extent when a syntactic violation is present and structural unification becomes difficult or impossible. The effects related to artificial syntax in the left inferior frontal region (BA 44/45) were essentially identical when we masked these with activity related to natural syntax in the same subjects. Finally, the medial temporal lobe was deactivated during this operation, consistent with the view that implicit processing does not rely on declarative memory mechanisms that engage the medial temporal lobe. In the context of recent FMRI findings, we raise the question whether Broca's region (or subregions) is specifically related to syntactic movement operations or the processing of hierarchically nested non-adjacent dependencies in the discussion section. We conclude that this is not the case. Instead, we argue that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner, independent of whether there are any processing requirements related to syntactic movement or hierarchically nested structures. In addition, we argue that the Chomsky hierarchy is not directly relevant for neurobiological systems. (C) 2010 Elsevier Inc. All rights reserved. Max Planck Institute for Psycholinguistics; Donders Institute for Brain, Cognition and Behaviour; FCT; LA; FEDER/POCI; Vetenskapsradet [8276]; Hedlunds Stiftelse; Stockholm County Council (ALF, FoUU) info:eu-repo/semantics/publishedVersion

Item does not contain fulltext Research on pattern perception and rule learning, grounded in formal language theory (FLT) and using artificial grammar learning paradigms, has exploded in the last decade. This approach marries empirical research conducted by neuroscientists, psychologists and ethologists with the theory of computation and FLT, developed by mathematicians, linguists and computer scientists over the last century. Of particular current interest are comparative extensions of this work to non-human animals, and neuroscientific investigations using brain imaging techniques. We provide a short introduction to the history of these fields, and to some of the dominant hypotheses, to help contextualize these ongoing research programmes, and finally briefly introduce the papers in the current issue. 8 p.

A recent hypothesis in empirical brain research on language is that the fundamental difference between animal and human communication systems is captured by the distinction between finite-state and more complex phrase-structure grammars, such as context-free and context-sensitive grammars. However, the relevance of this distinction for the study of language as a neurobiological system has been questioned and it has been suggested that a more relevant and partly analogous distinction is that between non-adjacent and adjacent dependencies. Online memory resources are central to the processing of non-adjacent dependencies as information has to be maintained across intervening material. One proposal is that an external memory device in the form of a limited push-down stack is used to process non-adjacent dependencies. We tested this hypothesis in an artificial grammar learning paradigm where subjects acquired non-adjacent dependencies implicitly. Generally, we found no qualitative differences between the acquisition of non-adjacent dependencies and adjacent dependencies. This suggests that although the acquisition of non-adjacent dependencies requires more exposure to the acquisition material, it utilizes the same mechanisms used for acquiring adjacent dependencies. We challenge the push-down stack model further by testing its processing predictions for nested and crossed multiple non-adjacent dependencies. The push-down stack model is partly supported by the results, and we suggest that stack-like properties are some among many natural properties characterizing the underlying neurophysiological mechanisms that implement the online memory resources used in language and structured sequence processing. info:eu-repo/semantics/publishedVersion

Item does not contain fulltext We investigated how two distortions of the speech signal - added background noise and speech in an unfamiliar accent - affect comprehension of speech using functional Magnetic Resonance Imaging (fMRI). Listeners performed a speeded sentence verification task for speech in quiet in Standard Dutch, in Standard Dutch with added background noise and for speech in an unfamiliar accent of Dutch. The behavioural results showed slower responses for both types of distortion compared to clear speech, and no difference between the two distortions. The neuroimaging results showed that, compared to clear speech, processing noise resulted in more activity bilaterally in Inferior Frontal Gyrus, Frontal Operculum, while processing accented speech recruited an area in left Superior Temporal Gyrus/Sulcus. It is concluded that the neural bases for processing different distortions of the speech signal dissociate. It is suggested that current models of the cortical organisation of speech are updated to specifically associate bilateral inferior frontal areas with processing external distortions (e.g., background noise) and left temporal areas with speaker-related distortions (e.g., accents). Crown Copyright (C) 2011 Published by Elsevier Ltd. All rights reserved.

Contains fulltext : 108202.pdf (Publisher’s version ) (Closed access) Identifying genetic variants influencing human brain structures may reveal new biological mechanisms underlying cognition and neuropsychiatric illness. The volume of the hippocampus is a biomarker of incipient Alzheimer's disease and is reduced in schizophrenia, major depression and mesial temporal lobe epilepsy. Whereas many brain imaging phenotypes are highly heritable, identifying and replicating genetic influences has been difficult, as small effects and the high costs of magnetic resonance imaging (MRI) have led to underpowered studies. Here we report genome-wide association meta-analyses and replication for mean bilateral hippocampal, total brain and intracranial volumes from a large multinational consortium. The intergenic variant rs7294919 was associated with hippocampal volume (12q24.22; N = 21,151; P = 6.70 x 10(-16)) and the expression levels of the positional candidate gene TESC in brain tissue. Additionally, rs10784502, located within HMGA2, was associated with intracranial volume (12q14.3; N = 15,782; P = 1.12 x 10(-12)). We also identified a suggestive association with total brain volume at rs10494373 within DDR2 (1q23.3; N = 6,500; P = 5.81 x 10(-7)). 01 mei 2012

Contains fulltext : 115530.pdf (Publisher’s version ) (Closed access) Models of speaking distinguish producing meaning, words and syntax as three different linguistic components of speaking. Nevertheless, little is known about the brain's integrated neuronal infrastructure for speech production. We investigated semantic, lexical and syntactic aspects of speaking using fMRI. In a picture description task, we manipulated repetition of sentence meaning, words, and syntax separately. By investigating brain areas showing response adaptation to repetition of each of these sentence properties, we disentangle the neuronal infrastructure for these processes. We demonstrate that semantic, lexical and syntactic processes are carried out in partly overlapping and partly distinct brain networks and show that the classic left-hemispheric dominance for language is present for syntax but not semantics. (C) 2012 Elsevier Inc. All rights reserved.

Contains fulltext : 115492.pdf (Publisher’s version ) (Open Access) Synesthesia provides an elegant model to investigate neural mechanisms underlying individual differences in subjective experience in humans. In grapheme-color synesthesia, written letters induce color sensations, accompanied by activation of color area V4. Competing hypotheses suggest that enhanced V4 activity during synesthesia is either induced by direct bottom-up cross-activation from grapheme processing areas within the fusiform gyrus, or indirectly via higher-order parietal areas. Synesthetes differ in the way synesthetic color is perceived: "projector" synesthetes experience color externally colocalized with a presented grapheme, whereas "associators" report an internally evoked association. Using dynamic causal modeling for fMRI, we show that V4 cross-activation during synesthesia was induced via a bottom-up pathway (within fusiform gyrus) in projector synesthetes, but via a top-down pathway (via parietal lobe) in associators. These findings show how altered coupling within the same network of active regions leads to differences in subjective experience. Our findings reconcile the two most influential cross-activation accounts of synesthesia. 6 p.

Contains fulltext : 86126.pdf (Publisher’s version ) (Closed access) Human communication is often thought about in terms of transmitted messages in a conventional code like a language. But communication requires a specialized interactive intelligence. Senders have to be able to perform recipient design, while receivers need to be able to do intention recognition, knowing that recipient design has taken place. To study this interactive intelligence in the lab, we developed a new task that taps directly into the underlying abilities to communicate in the absence of a conventional code. We show that subjects are remarkably successful communicators under these conditions, especially when senders get feedback from receivers. Signaling is accomplished by the manner in which an instrumental action is performed, such that instrumentally dysfunctional components of an action are used to convey communicative intentions. The findings have important implications for the nature of the human communicative infrastructure, and the task opens up a line of experimentation on human communication.

Contains fulltext : 90335.pdf (Publisher’s version ) (Open Access) There is growing evidence suggesting that synchronization changes in the oscillatory neuronal dynamics in the EEG or MEG reflect the transient coupling and uncoupling of functional networks related to different aspects of language comprehension. In this work, we examine how sentence-level syntactic unification operations are reflected in the oscillatory dynamics of the MEG. Participants read sentences that were either correct, contained a word category violation, or were constituted of random word sequences devoid of syntactic structure. A time-frequency analysis of MEG power changes revealed three types of effects. The first type of effect was related to the detection of a (word category) violation in a syntactically structured sentence, and was found in the alpha and gamma frequency bands. A second type of effect was maximally sensitive to the syntactic manipulations: A linear increase in beta power across the sentence was present for correct sentences, was disrupted upon the occurrence of a word category violation, and was absent in syntactically unstructured random word sequences. We therefore relate this effect to syntactic unification operations. Thirdly, we observed a linear increase in theta power across the sentence for all syntactically structured sentences. The effects are tentatively related to the building of a working memory trace of the linguistic input. In conclusion, the data seem to suggest that syntactic unification is reflected by neuronal synchronization in the lower-beta frequency band.

We performed a resting-state functional connectivity study to investigate directly the functional correlations within the perisylvian language networks by seeding from 3 subregions of Broca's complex (pars opercularis, pars triangularis, and pars orbitalis) and their right hemisphere homologues. A clear topographical functional connectivity pattern in the left middle frontal, parietal, and temporal areas was revealed for the 3 left seeds. This is the first demonstration that a functional connectivity topology can be observed in the perisylvian language networks. The results support the assumption of the functional division for phonology, syntax, and semantics of Broca's complex as proposed by the memory, unification, and control (MUC) model and indicated a topographical functional organization in the perisylvian language networks, which suggests a possible division of labor for phonological, syntactic, and semantic function in the left frontal, parietal, and temporal areas.

Contains fulltext : 70720.pdf (Publisher’s version ) (Open Access) When do listeners take into account who the speaker is? We asked people to listen to utterances whose content sometimes did not match inferences based on the identity of the speaker (e.g., "If only I looked like Britney Spears" in a male voice, or "I have a large tattoo on my back" spoken with an upper-class accent). Event-related brain responses revealed that the speaker's identity is taken into account as early as 200-300 msec after the beginning of a spoken word, and is processed by the same early interpretation mechanism that constructs sentence meaning based on just the words. This finding is difficult to reconcile with standard "Gricean" models of sentence interpretation in which comprehenders initially compute a local, context-independent meaning for the sentence ("semantics") before working out what it really means given the wider communicative context and the particular speaker ("pragmatics"). Because the observed brain response hinges on voice-based and usually stereotype-dependent inferences about the speaker, it also shows that listeners rapidly classify speakers on the basis of their voices and bring the associated social stereotypes to bear on what is being said. According to our event-related potential results, language comprehension takes very rapid account of the social context, and the construction of meaning based on language alone cannot be separated from the social aspects of language use. The linguistic brain relates the message to the speaker immediately.

Contains fulltext : 49746.pdf (Publisher’s version ) (Closed access) An event-related brain potential experiment was carried out to investigate the temporal relationship between lexical selection and the semantic integration in auditory sentence processing. Participants were presented with spoken sentences that ended with a word that was either semantically congruent or anomalous. Information about the moment in which a sentence-final word could uniquely be identified, its isolation point (IP), was compared with the onset of the elicited N400 congruity effect, reflecting semantic integration processing. The results revealed that the onset of the N400 effect occurred prior to the IP of the sentence-final words. Moreover, the factor early or late IP did not affect the onset of the N400. These findings indicate that lexical selection and semantic integration are cascading processes, in that semantic integration processing can start before the acoustic information allows the selection of a unique candidate and seems to be attempted in parallel for multiple candidates that are still compatible with the bottom-up acoustic input.

Comprehension of pointing gestures is fundamental to human communication. However, the neural mechanisms that subserve the integration of pointing gestures and speech in visual contexts in comprehension are unclear. Here we present the results of an fMRI study in which participants watched images of an actor pointing at an object while they listened to her referential speech. The use of a mismatch paradigm revealed that the semantic unication of pointing gesture and speech in a triadic context recruits left inferior frontal gyrus. Complementing previous ndings, this suggests that left inferior frontal gyrus semantically integrates information across modalities and semiotic domains.

Contains fulltext : 150217.pdf (Publisher’s version ) (Open Access) Frequent and predictable words are generally pronounced with less effort and are therefore acoustically more reduced than less frequent or unpredictable words. Local predictability can be operationalised by Transitional Probability (TP), which indicates how likely a word is to occur given its immediate context. We investigated whether and how probabilistic reduction effects on word durations change with adult age when reading aloud content words embedded in sentences. The results showed equally large frequency effects on verb and noun durations for both younger (Mage = 20 years) and older (Mage = 68 years) adults. Backward TP also affected word duration for younger and older adults alike. ForwardTP, however, had no significant effect on word duration in either age group. Our results resemble earlier findings of more robust BackwardTP effects compared to ForwardTP effects. Furthermore, unlike often reported decline in predictive processing with aging, probabilistic reduction effects remain stable across adulthood.

Speakers can adapt their speech and co-speech gestures for addressees. Here, we investigate whether this ability is modulated by age. Younger and older adults participated in a comic narration task in which one participant (the speaker) narrated six short comic stories to another participant (the addressee). One half of each story was known to both participants, the other half only to the speaker. Younger but not older speakers used more words and gestures when narrating novel story content as opposed to known content. We discuss cognitive and pragmatic explanations of these findings and relate them to theories of gesture production.

Speakers adapt their speech and gestures in various ways for their audience. We investigated further whether they use ostensive signals (eye gaze, ostensive speech (e.g. like this, this) or a combination of both) in relation to their gestures when talking to different addressees, i.e., to another adult or a child in a multimodal demonstration task. While adults used more eye gaze towards their gestures with other adults than with children, they were more likely to use combined ostensive signals for children than for adults. Thus speakers mark the communicative relevance of their gestures with different types of ostensive signals and by taking different types of addressees into account.

Contains fulltext : 150206.pdf (Publisher’s version ) (Open Access) We investigated music and language processing in a group of early bilinguals who spoke a tone language and a non-tone language (Cantonese and Dutch). We assessed online speech-music processing interactions, that is, interactions that occur when speech and music are processed simultaneously in songs, with a speeded classification task. In this task, participants judged sung pseudowords either musically (based on the direction of the musical interval) or phonologically (based on the identity of the sung vowel). We also assessed longer-term effects of linguistic experience on musical ability, that is, the influence of extensive prior experience with language when processing music. These effects were assessed with a task in which participants had to learn to identify musical intervals and with four pitch-perception tasks. Our hypothesis was that due to their experience in two different languages using lexical versus intonational tone, the early Cantonese-Dutch bilinguals would outperform the Dutch control participants. In online processing, the Cantonese-Dutch bilinguals processed speech and music more holistically than controls. This effect seems to be driven by experience with a tone language, in which integration of segmental and pitch information is fundamental. Regarding longer-term effects of linguistic experience, we found no evidence for a bilingual advantage in either the music-interval learning task or the pitch-perception tasks. Together, these results suggest that being a Cantonese-Dutch bilingual does not have any measurable longer-term effects on pitch and music processing, but does have consequences for how speech and music are processed jointly. 20 p.

Contains fulltext : 145518.pdf (Publisher’s version ) (Closed access) Emotions are often expressed metaphorically, and both emotion and metaphor are ways through which abstract meaning can be grounded in language. Here we investigate specifically whether motion-related verbs when used metaphorically are differentially sensitive to a preceding emotional context, as compared to when they are used in a literal manner. Participants read stories that ended with ambiguous action/motion sentences (e.g., he got it), in which the action/motion could be interpreted metaphorically (he understood the idea) or literally (he caught the ball) depending on the preceding story. Orthogonal to the metaphorical manipulation, the stories were high or low in emotional content. The results showed that emotional context modulated the neural response in visual motion areas to the metaphorical interpretation of the sentences, but not to their literal interpretations. In addition, literal interpretations of the target sentences led to stronger activation in the visual motion areas as compared to metaphorical readings of the sentences. We interpret our results as suggesting that emotional context specifically modulates mental simulation during metaphor processing. 7 p.

In this study, we explore the possibility to predict the semantic category of words from brain signals in a free word generation task. Participants produced single words from different semantic categories in a modified semantic fluency task. A Bayesian logistic regression classifier was trained to predict the semantic category of words from single-trial MEG data. Significant classification accuracies were achieved using sensor-level MEG time series at the time interval of conceptual preparation. Semantic category prediction was also possible using source-reconstructed time series, based on minimum norm estimates of cortical activity. Brain regions that contributed most to classification on the source level were identified. These were the left inferior frontal gyrus, left middle frontal gyrus, and left posterior middle temporal gyrus. Additionally, the temporal dynamics of brain activity underlying the semantic preparation during word generation was explored. These results provide important insights about central aspects of language production.

Item does not contain fulltext We sketch four applications of Marr's levels-of-analysis methodology to the relations between logic and experimental data in the cognitive neuroscience of language and reasoning. The first part of the paper illustrates the explanatory power of computational level theories based on logic. We show that a Bayesian treatment of the suppression task in reasoning with conditionals is ruled out by EEG data, supporting instead an analysis based on defeasible logic. Further, we describe how results from an EEG study on temporal prepositions can be reanalyzed using formal semantics, addressing a potential confound. The second part of the article demonstrates the predictive power of logical theories drawing on EEG data on processing progressive constructions and on behavioral data on conditional reasoning in people with autism. Logical theories can constrain processing hypotheses all the way down to neurophysiology, and conversely neuroscience data can guide the selection of alternative computational level models of cognition. 13 p.

Contains fulltext : 127147.pdf (Publisher’s version ) (Open Access) A number of recent studies have hypothesized that monitoring in speech production may occur via domain-general mechanisms responsible for the detection of response conflict. Outside of language, two ERR components have consistently been elicited in conflict-inducing tasks (e.g., the flanker task): the stimulus-locked N2 on correct trials, and the response-locked error-related negativity (ERN). The present investigation used these electrophysiological markers to test whether a common response conflict monitor is responsible for monitoring in speech and non-speech tasks. Electroencephalography (EEG) was recorded while participants performed a tongue twister (TT) task and a manual version of the flanker task. In the TT task, people rapidly read sequences of four nonwords arranged in TT and non-TT patterns three times. In the flanker task, people responded with a left/right button press to a center-facing arrow, and conflict was manipulated by the congruency of the flanking arrows. Behavioral results showed typical effects of both tasks, with increased error rates and slower speech onset times forTT relative to non-TT trials and for incongruent relative to congruent flanker trials. In the flanker task, stimulus-locked EEG analyses replicated previous results, with a larger N2 for incongruent relative to congruent trials, and a response-locked ERN. In the TT task, stimulus-locked analyses revealed broad, frontally-distributed differences beginning around 50 ms and lasting until just before speech initiation, with TT trials more negative than non-TT trials; response-locked analyses revealed an ERN. Correlation across these measures showed some correlations within a task, but little evidence of systematic cross-task correlation. Although the present results do not speak against conflict signals from the production system serving as cues to self-monitoring, they are not consistent with signatures of response conflict being mediated by a single, domain-general conflict monitor. 16 p.

Item does not contain fulltext Language and action systems are highly interlinked. A critical piece of evidence is that speech and its accompanying gestures are tightly synchronized. Five experiments were conducted to test 2 hypotheses about the synchronization of speech and gesture. According to the interactive view, there is continuous information exchange between the gesture and speech systems, during both their planning and execution phases. According to the ballistic view, information exchange occurs only during the planning phases of gesture and speech, but the 2 systems become independent once their execution has been initiated. In all experiments, participants were required to point to and/or name a light that had just lit up. Virtual reality and motion tracking technologies were used to disrupt their gesture or speech execution. Participants delayed their speech onset when their gesture was disrupted. They did so even when their gesture was disrupted at its late phase and even when they received only the kinesthetic feedback of their gesture. Also, participants prolonged their gestures when their speech was disrupted. These findings support the interactive view and add new constraints on models of speech and gesture production. 16 p.

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Contains fulltext : 139798.pdf (Publisher’s version ) (Open Access) A fundamental property of language is that it can be used to refer to entities in the extra-linguistic physical context of a conversation in order to establish a joint focus of attention on a referent. Typological and psycholinguistic work across a wide range of languages has put forward at least two different theoretical views on demonstrative reference. Here we contrasted and tested these two accounts by investigating the electrophysiological brain activity underlying the construction of indexical meaning in comprehension. In two EEG experiments, participants watched pictures of a speaker who referred to one of two objects using speech and an index-finger pointing gesture. In contrast with separately collected native speakers' linguistic intuitions, N400 effects showed a preference for a proximal demonstrative when speaker and addressee were in a face-to-face orientation and all possible referents were located in the shared space between them, irrespective of the physical proximity of the referent to the speaker. These findings reject egocentric proximity-based accounts of demonstrative reference, support a sociocentric approach to deixis, suggest that interlocutors construe a shared space during conversation, and imply that the psychological proximity of a referent may be more important than its physical proximity. 21 p.

Item does not contain fulltext To investigate the neural underpinnings of word decoding, and how it changes as a function of repeated exposure, we trained Dutch participants repeatedly over the course of a month of training to articulate a set of novel disyllabic input strings written in Greek script to avoid the use of familiar orthographic representations. The syllables in the input were phonotactically legal combinations but non-existent in the Dutch language, allowing us to assess their role in novel word decoding. Not only trained disyllabic pseudowords were tested but also pseudowords with recombined patterns of syllables to uncover the emergence of syllabic representations. We showed that with extensive training, articulation became faster and more accurate for the trained pseudowords. On the neural level, the initial stage of decoding was reflected by increased activity in visual attention areas of occipito-temporal and occipito-parietal cortices, and in motor coordination areas of the precentral gyrus and the inferior frontal gyrus. After one month of training, memory representations for holistic information (whole word unit) were established in areas encompassing the angular gyrus, the precuneus and the middle temporal gyrus. Syllabic representations also emerged through repeated training of disyllabic pseudowords, such that reading recombined syllables of the trained pseudowords showed similar brain activation to trained pseudowords and were articulated faster than novel combinations of letter strings used in the trained pseudowords. 16 p.

Contains fulltext : 127930.pdf (Publisher’s version ) (Closed access) In this fMRI study we investigate the neural correlates of information structure integration during sentence comprehension in Dutch. We looked into how prosodic cues (pitch accents) that signal the information status of constituents to the listener (new information) are combined with other types of information during the unification process. The difficulty of unifying the prosodic cues into overall sentence meaning was manipulated by constructing sentences in which the pitch accent did (focus-accent agreement), and sentences in which the pitch accent did not (focus-accent disagreement) match the expectations for focus constituents of the sentence. In case of a mismatch, the load on unification processes increases. Our results show two anatomically distinct effects of focus-accent disagreement, one located in the posterior left inferior frontal gyrus (LIFG, BA6/44), and one in the more anterior-ventral LIFG (BA 47/45). Our results confirm that information structure is taken into account during unification, and imply an important role for the LIFG in unification processes, in line with previous fMRI studies.

Contains fulltext : 121905.pdf (Publisher’s version ) (Open Access) A neurobiological model of language is discussed that overcomes the shortcomings of the classical Wernicke-Lichtheim-Geschwind model. It is based on a subdivision of language processing into three components: Memory, Unification, and Control. The functional components as well as the neurobiological underpinnings of the model are discussed. In addition, the need for extension of the model beyond the classical core regions for language is shown. The attention network and the network for inferential processing are crucial to realize language comprehension beyond single word processing and beyond decoding propositional content. It is shown that this requires the dynamic interaction between multiple brain regions.

Contains fulltext : 132365.pdf (Publisher’s version ) (Open Access) Even though language allows us to say exactly what we mean, we often use language to say things indirectly, in a way that depends on the specific communicative context. For example, we can use an apparently straightforward sentence like "It is hard to give a good presentation" to convey deeper meanings, like "Your talk was a mess!" One of the big puzzles in language science is how listeners work out what speakers really mean, which is a skill absolutely central to communication. However, most neuroimaging studies of language comprehension have focused on the arguably much simpler, context-independent process of understanding direct utterances. To examine the neural systems involved in getting at contextually constrained indirect meaning, we used functional magnetic resonance imaging as people listened to indirect replies in spoken dialog. Relative to direct control utterances, indirect replies engaged dorsomedial prefrontal cortex, right temporo-parietal junction and insula, as well as bilateral inferior frontal gyrus and right medial temporal gyrus. This suggests that listeners take the speaker's perspective on both cognitive (theory of mind) and affective (empathy-like) levels. In line with classic pragmatic theories, our results also indicate that currently popular "simulationist" accounts of language comprehension fail to explain how listeners understand the speaker's intended message. 7 p.

Item does not contain fulltext Here we review evidence from cognitive neuroscience for a tight relation between language and action in the brain. We focus on two types of relation between language and action. First, we investigate whether the perception of speech and speech sounds leads to activation of parts of the cortical motor system also involved in speech production. Second, we evaluate whether understanding action-related language involves the activation of parts of the motor system. We conclude that whereas there is considerable evidence that understanding language can involve parts of our motor cortex, this relation is best thought of as inherently flexible. As we explain, the exact nature of the input as well as the intention with which language is perceived influences whether and how motor cortex plays a role in language processing.

Contains fulltext : 115430.pdf (Publisher’s version ) (Closed access) In spoken language, pitch accent can mark certain information as focus, whereby more attentional resources are allocated to the focused information. Using functional magnetic resonance imaging, this study examined whether pitch accent, used for marking focus, recruited general attention networks during sentence comprehension. In a language task, we independently manipulated the prosody and semantic/pragmatic congruence of sentences. We found that semantic/pragmatic processing affected bilateral inferior and middle frontal gyrus. The prosody manipulation showed bilateral involvement of the superior/inferior parietal cortex, superior and middle temporal cortex, as well as inferior, middle, and posterior parts of the frontal cortex. We compared these regions with attention networks localized in an auditory spatial attention task. Both tasks activated bilateral superior/inferior parietal cortex, superior temporal cortex, and left precentral cortex. Furthermore, an interaction between prosody and congruence was observed in bilateral inferior parietal regions: for incongruent sentences, but not for congruent ones, there was a larger activation if the incongruent word carried a pitch accent, than if it did not. The common activations between the language task and the spatial attention task demonstrate that pitch accent activates a domain general attention network, which is sensitive to semantic/pragmatic aspects of language. Therefore, attention and language comprehension are highly interactive. 13 p.

Contains fulltext : 116682-OA.pdf (Publisher’s version ) (Open Access) The posterior middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) are two critical nodes of the brain's language network. Previous neuroimaging evidence has supported a dissociation in language comprehension in which parts of the MTG are involved in the retrieval of lexical syntactic information and the IFG in unification operations that maintain, select, and integrate multiple sources of information over time. In the present investigation, we tested for causal evidence of this dissociation by modulating activity in IFG and MTG using an offline TMS procedure: continuous theta-burst stimulation. Lexical–syntactic retrieval was manipulated by using sentences with and without a temporarily word-class (noun/verb) ambiguity (e.g., run). In one group of participants, TMS was applied to the IFG and MTG, and in a control group, no TMS was applied. Eye movements were recorded and quantified at two critical sentence regions: a temporarily ambiguous region and a disambiguating region. Results show that stimulation of the IFG led to a modulation of the ambiguity effect (ambiguous–unambiguous) at the disambiguating sentence region in three measures: first fixation durations, total reading times, and regressive eye movements into the region. Both IFG and MTG stimulation modulated the ambiguity effect for total reading times in the temporarily ambiguous sentence region relative to the control group. The current results demonstrate that an offline repetitive TMS protocol can have influences at a different point in time during online processing and provide causal evidence for IFG involvement in unification operations during sentence comprehension. 14 p.

Contains fulltext : 115428.pdf (Publisher’s version ) (Closed access) Behavioral syntactic priming effects during sentence comprehension are typically observed only if both the syntactic structure and lexical head are repeated. In contrast, during production syntactic priming occurs with structure repetition alone, but the effect is boosted by repetition of the lexical head. We used fMRI to investigate the neuronal correlates of syntactic priming and lexical boost effects during sentence production and comprehension. The critical measure was the magnitude of fMRI adaptation to repetition of sentences in active or passive voice, with or without verb repetition. In conditions with repeated verbs, we observed adaptation to structure repetition in the left IFG and MTG, for active and passive voice. However, in the absence of repeated verbs, adaptation occurred only for passive sentences. None of the fMRI adaptation effects yielded differential effects for production versus comprehension, suggesting that sentence comprehension and production are subserved by the same neuronal infrastructure for syntactic processing.

Item does not contain fulltext Grapheme-color synesthetes perceive color when reading letters or digits. We investigated oscillatory brain signals of synesthetes vs. controls using magnetoencephalography. Brain oscillations specifically in the alpha band (∼10 Hz) have two interesting features: alpha has been linked to inhibitory processes and can act as a marker for attention. The possible role of reduced inhibition as an underlying cause of synesthesia, as well as the precise role of attention in synesthesia is widely discussed. To assess alpha power effects due to synesthesia, synesthetes as well as matched controls viewed synesthesia-inducing graphemes, colored control graphemes, and non-colored control graphemes while brain activity was recorded. Subjects had to report a color change at the end of each trial which allowed us to assess the strength of synesthesia in each synesthete. Since color (synesthetic or real) might allocate attention we also included an attentional cue in our paradigm which could direct covert attention. In controls the attentional cue always caused a lateralization of alpha power with a contralateral decrease and ipsilateral alpha increase over occipital sensors. In synesthetes, however, the influence of the cue was overruled by color: independent of the attentional cue, alpha power decreased contralateral to the color (synesthetic or real). This indicates that in synesthetes color guides attention. This was confirmed by reaction time effects due to color, i.e. faster RTs for the color side independent of the cue. Finally, the stronger the observed color dependent alpha lateralization, the stronger was the manifestation of synesthesia as measured by congruency effects of synesthetic colors on RTs. Behavioral and imaging results indicate that color induces a location-specific, automatic shift of attention towards color in synesthetes but not in controls. We hypothesize that this mechanism can facilitate coupling of grapheme and color during the development of synesthesia.

Around their first birthday infants begin to talk, yet they comprehend words long before. This study investigated the event-related potentials (ERP) responses of nine-month-olds on basic level picture-word pairings. After a familiarization phase of six picture-word pairings per semantic category, comprehension for novel exemplars was tested in a picture-word matching paradigm. ERPs time-locked to pictures elicited a modulation of the Negative Central (Nc) component, associated with visual attention and recognition. It was attenuated by category repetition as well as by the type-token ratio of picture context. ERPs time-locked to words in the training phase became more negative with repetition (N300-600), but there was no influence of picture type-token ratio, suggesting that infants have identified the concept of each picture before a word was presented. Results from the test phase provided clear support that infants integrated word meanings with (novel) picture context. Here, infants showed different ERP responses for words that did or did not align with the picture context: a phonological mismatch (N200) and a semantic mismatch (N400). Together, results were informative of visual categorization, word recognition and word-to-world-mappings, all three crucial processes for vocabulary construction. (C) 2012 Elsevier Ltd. All rights reserved.

We carried out the first study on the relationship between individual language aptitude and structural connectivity of language pathways in the adult brain. We measured four components of language aptitude (vocabulary learning, VocL; sound recognition, SndRec; sound-symbol correspondence, SndSym; and grammatical inferencing, GrInf) using the LLAMA language aptitude test (Meara, 2005). Spatial working memory (SWM), verbal working memory (VWM) and IQ were also measured as control factors. Diffusion Tensor Imaging (DTI) was employed to investigate the structural connectivity of language pathways in the perisylvian language network. Principal Component Analysis (PCA) on behavioural measures suggests that a general ability might be important to the first stages of L2 acquisition. It also suggested that VocL, SndSy and SWM are more closely related to general IQ than SndRec and VocL, and distinguished the tasks specifically designed to tap into L2 acquisition (VocL, SndRec,SndSym and GrInf) from more generic measures (IQ, SWM and VWM). Regression analysis suggested significant correlations between most of these behavioural measures and the structural connectivity of certain language pathways, i.e., VocL and BA47-Parietal pathway, SndSym and inter-hemispheric BA45 pathway, GrInf and BA45-Temporal pathway and BA6-Temporal pathway, IQ and BA44-Parietal pathway, BA47-Parietal pathway, BA47-Temporal pathway and inter-hemispheric BA45 pathway, SWM and inter-hemispheric BA6 pathway and BA47-Parietal pathway, and VWM and BA47-Temporal pathway. These results are discussed in relation to relevant findings in the literature.

Contains fulltext : 102636.pdf (Publisher’s version ) (Open Access) Information structure facilitates communication between interlocutors by highlighting relevant information. It has previously been shown that information structure modulates the depth of semantic processing. Here we used event-related potentials to investigate whether information structure can modulate the depth of syntactic processing. In question-answer pairs, subtle (number agreement) or salient (phrase structure) syntactic violations were placed either in focus or out of focus through information structure marking. P600 effects to these violations reflect the depth of syntactic processing. For subtle violations, a P600 effect was observed in the focus condition, but not in the non-focus condition. For salient violations, comparable P600 effects were found in both conditions. These results indicate that information structure can modulate the depth of syntactic processing, but that this effect depends on the salience of the information. When subtle violations are not in focus, they are processed less elaborately. We label this phenomenon the Chomsky illusion. 9 p.

Geen andere soort dan homo sapiens heeft in de loop van zijn evolutionaire geschiedenis een communicatiesysteem ontwikkeld waarin een eindig aantal symbolen samen met een reeks van regels voor het combineren daarvan een oneindig aantal uitdrukkingen mogelijk maakt. Dit natuurlijke taalsysteem stelt leden van onze soort in staat gedachten een uiterlijke vorm te geven en uit te wisselen met de sociale groep en, door de uitvinding van schriftsystemen, met de gehele samenleving. Spraak en taal zijn effectieve middelen voor het behoud van sociale cohesie in samenlevingen waarvan de groepsgrootte en de complexe sociale organisatie van dien aard is dat dit niet langer kan door middel van ‘vlooien’, de wijze waarop onze genetische buren, de primaten van de oude wereld, sociale cohesie bevorderen [1,2].

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Contains fulltext : 102326.pdf (Publisher’s version ) (Open Access) The genetic FOXP2-CNTNAP2 pathway has been shown to be involved in the language capacity. We investigated whether a common variant of CNTNAP2 (rs7794745) is relevant for syntactic and semantic processing in the general population by using a visual sentence processing paradigm while recording ERPs in 49 healthy adults. While both AA homozygotes and T-carriers showed a standard N400 effect to semantic anomalies, the response to subject-verb agreement violations differed across genotype groups. T-carriers displayed an anterior negativity preceding the P600 effect, whereas for the AA group only a P600 effect was observed. These results provide another piece of evidence that the neuronal architecture of the human faculty of language is shaped differently by effects that are genetically determined. 8 p.

Item does not contain fulltext Rhyme processing is reflected in the electrophysiological signals of the brain as a negative deflection for non-rhyming as compared to rhyming stimuli around 450 ms after stimulus onset. Studies have shown that this N450 component is not solely sensitive to rhyme but also responds to other types of phonological overlap. In the present study, we examined whether the N450 component can be used to gain insight into the global similarity effect, indicating that rhyme judgment skills decrease when participants are presented with word pairs that share a phonological overlap but do not rhyme (e.g., bell-ball). We presented 20 adults with auditory rhyming, globally similar overlapping and unrelated word pairs. In addition to measuring behavioral responses by means of a yes/no button press, we also took EEG measures. The behavioral data showed a clear global similarity effect; participants judged overlapping pairs more slowly than unrelated pairs. However, the neural outcomes did not provide evidence that the N450 effect responds differentially to globally similar and unrelated word pairs, suggesting that globally similar and dissimilar non-rhyming pairs are processed in a similar fashion at the stage of early lexical access. 8 p.

Contains fulltext : 102797-OA.pdf (Publisher’s version ) (Open Access) Research from the past decade has shown that understanding the meaning of words and utterances (i.e., abstracted symbols) engages the same systems we used to perceive and interact with the physical world in a content-specific manner. For example, understanding the word "grasp" elicits activation in the cortical motor network, that is, part of the neural substrate involved in planned and executing a grasping action. In the embodied literature, cortical motor activation during language comprehension is thought to reflect motor simulation underlying conceptual knowledge [note that outside the embodied framework, other explanations for the link between action and language are offered, e.g., Mahon, B. Z., & Caramazza, A. A critical look at the embodied cognition hypothesis and a new proposal for grouding conceptual content. Journal of Physiology, 102, 59-70, 2008; Hagoort, P. On Broca, brain, and binding: A new framework. Trends in Cognitive Sciences, 9, 416-423, 2005]. Previous research has supported the view that the coupling between language and action is flexible, and reading an action-related word form is not sufficient for cortical motor activation [Van Dam, W. O., van Dijk, M., Bekkering, H., & Rueschemeyer, S.-A. Flexibility in embodied lexical-semantic representations. Human Brain Mapping, doi: 10.1002/hbm.21365, 2011]. The current study goes one step further by addressing the necessity of action-related word forms for motor activation during language comprehension. Subjects listened to indirect requests (IRs) for action during an fMRI session. IRs for action are speech acts in which access to an action concept is required, although it is not explicitly encoded in the language. For example, the utterance "It is hot here!" in a room with a window is likely to be interpreted as a request to open the window. However, the same utterance in a desert will be interpreted as a statement. The results indicate (1) that comprehension of IR sentences activates cortical motor areas reliably more than comprehension of sentences devoid of any implicit motor information. This is true despite the fact that IR sentences contain no lexical reference to action. (2) Comprehension of IR sentences also reliably activates substantial portions of the theory of mind network, known to be involved in making inferences about mental states of others. The implications of these findings for embodied theories of language are discussed. 11 p.

Item does not contain fulltext The relationship between the evoked responses (ERPs/ERFs) and the event-related changes in EEG/MEG power that can be observed during sentence-level language comprehension is as yet unclear. This study addresses a possible relationship between MEG power changes and the N400m component of the event-related field. Whole-head MEG was recorded while subjects listened to spoken sentences with incongruent (IC) or congruent (C) sentence endings. A clear N400m was observed over the left hemisphere, and was larger for the IC sentences than for the C sentences. A timefrequency analysis of power revealed a decrease in alpha and beta power over the left hemisphere in roughly the same time range as the N400m for the IC relative to the C condition. A linear regression analysis revealed a positive linear relationship between N400m and beta power for the IC condition, not for the C condition. No such linear relation was found between N400m and alpha power for either condition. The sources of the beta decrease were estimated in the LIFG, a region known to be involved in semantic unification operations. One source of the N400m was estimated in the left superior temporal region, which has been related to lexical retrieval. We interpret our data within a framework in which beta oscillations are inversely related to the engagement of task-relevant brain networks. The source reconstructions of the beta power suppression and the N400m effect support the notion of a dynamic communication between the LIFG and the left superior temporal region during language comprehension. Hum Brain Mapp 33:2898-2912, 2012. (c) 2012 Wiley Periodicals, Inc. 15 p.

Item does not contain fulltext Infants’ ability to recognize words in continuous speech is vital for building a vocabulary. We here examined the amount and type of exposure needed for 10-month-olds to recognize words. Infants first heard a word, either embedded within an utterance or in isolation, then recognition was assessed by comparing event-related potentials to this word versus a word that they had not heard directly before. Although all 10-month-olds showed recognition responses to words first heard in isolation, not all infants showed such responses to words they had first heard within an utterance. Those that did succeed in the latter, harder, task, however, understood more words and utterances when re-tested at 12 months, and understood more words and produced more words at 24 months, compared with those who had shown no such recognition response at 10 months. The ability to rapidly recognize the words in continuous utterances is clearly linked to future language development. 11 p.

Contains fulltext : 102503.pdf (Publisher’s version ) (Open Access) When an adult claims he cannot sleep without his teddy bear, people tend to react surprised. Language interpretation is, thus, influenced by social context, such as who the speaker is. The present study reveals inter-individual differences in brain reactivity to social aspects of language. Whereas women showed brain reactivity when stereotype-based inferences about a speaker conflicted with the content of the message, men did not. This sex difference in social information processing can be explained by a specific cognitive trait, one's ability to empathize. Individuals who empathize to a greater degree revealed larger N400 effects (as well as a larger increase in gamma-band power) to socially relevant information. These results indicate that individuals with high-empathizing skills are able to rapidly integrate information about the speaker with the content of the message, as they make use of voice-based inferences about the speaker to process language in a top-down manner. Alternatively, individuals with lower empathizing skills did not use information about social stereotypes in implicit sentence comprehension, but rather took a more bottom-up approach to the processing of these social pragmatic sentences. 11 p.

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Contains fulltext : 102872.pdf (Publisher’s version ) (Closed access) During speaking and listening syntactic processing is a crucial step. It involves specifying syntactic relations between words in a sentence. If the production and comprehension modality share the neuronal substrate for syntactic processing then processing syntax in one modality should lead to adaptation effects in the other modality. In the present functional magnetic resonance imaging experiment, participants either overtly produced or heard descriptions of pictures. We looked for brain regions showing adaptation effects to the repetition of syntactic structures. In order to ensure that not just the same brain regions but also the same neuronal populations within these regions are involved in syntactic processing in speaking and listening, we compared syntactic adaptation effects within processing modalities (syntactic production-to-production and comprehension-to-comprehension priming) with syntactic adaptation effects between processing modalities (syntactic comprehension-to-production and production-to-comprehension priming). We found syntactic adaptation effects in left inferior frontal gyrus (Brodmann's area [BA] 45), left middle temporal gyrus (BA 21), and bilateral supplementary motor area (BA 6) which were equally strong within and between processing modalities. Thus, syntactic repetition facilitates syntactic processing in the brain within and across processing modalities to the same extent. We conclude that that the same neurobiological system seems to subserve syntactic processing in speaking and listening. 9 p.

Item does not contain fulltext Semantic unification, the process by which small blocks of semantic information are combined into a coherent utterance, has been studied with various types of tasks. However, whether the brain activations reported in these studies are attributed to semantic unification per se or to other task-induced concomitant processes still remains unclear. The neural basis for semantic unification in sentence comprehension was examined using event-related potentials (ERP) and functional Magnetic Resonance Imaging (fMRI). The semantic unification load was manipulated by varying the goodness of fit between a critical word and its preceding context (in high cloze, low cloze and violation sentences). The sentences were presented in a serial visual presentation mode. The participants were asked to perform one of three tasks: semantic congruency judgment (SEM), silent reading for comprehension (READ), or font size judgment (FONT), in separate sessions. The ERP results showed a similar N400 amplitude modulation by the semantic unification load across all of the three tasks. The brain activations associated with the semantic unification load were found in the anterior left inferior frontal gyrus (aLIFG) in the FONT task and in a widespread set of regions in the other two tasks. These results suggest that the aLIFG activation reflects a semantic unification, which is different from other brain activations that may reflect task-specific strategic processing. 8 p.

Item does not contain fulltext A repetitionsuppression functional magnetic resonance imaging paradigm was used to explore the neuroanatomical substrates of processing two types of acoustic variationspeaker and accentduring spoken sentence comprehension. Recordings were made for two speakers and two accents: Standard Dutch and a novel accent of Dutch. Each speaker produced sentences in both accents. Participants listened to two sentences presented in quick succession while their haemodynamic responses were recorded in an MR scanner. The first sentence was spoken in Standard Dutch; the second was spoken by the same or a different speaker and produced in Standard Dutch or in the artificial accent. This design made it possible to identify neural responses to a switch in speaker and accent independently. A switch in accent was associated with activations in predominantly left-lateralized areas including posterior temporal regions, including superior temporal gyrus, planum temporale (PT), and supramarginal gyrus, as well as in frontal regions, including left pars opercularis of the inferior frontal gyrus (IFG). A switch in speaker recruited a predominantly right-lateralized network, including middle frontal gyrus and prenuneus. It is concluded that posterior temporal areas, including PT, and frontal areas, including IFG, are involved in processing accent variation in spoken sentence comprehension. Hum Brain Mapp, 2012. (C) 2010 Wiley Periodicals, Inc.