Publications

Abstract

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 is involved 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 a 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.

Abstract

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. Attentional networks as well as networks 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.

Abstract

This chapter sketches in very general terms the cognitive architecture of both language comprehension and production, as well as the neurobiological infrastructure that makes the human brain ready for language. Focus is on spoken language, since that compares most directly to processing music. It is worth bearing in mind that humans can also interface with language as a cognitive system using sign and text (visual) as well as Braille (tactile); that is to say, the system can connect with input/output processes in any sensory modality. Language processing consists of a complex and nested set of subroutines to get from sound to meaning (in comprehension) or meaning to sound (in production), with remarkable speed and accuracy. The fi rst section outlines a selection of the major constituent operations, from fractionating the input into manageable units to combining and unifying information in the construction of meaning. The next section addresses the neurobiological infrastructure hypothesized to form the basis for language processing. Principal insights are summarized by building on the notion of “brain networks” for speech–sound processing, syntactic processing, and the construction of meaning, bearing in mind that such a neat three-way subdivision overlooks important overlap and shared mechanisms in the neural architecture subserving language processing. Finally, in keeping with the spirit of the volume, some possible relations are highlighted between language and music that arise from the infrastructure developed here. Our characterization of language and its neurobiological foundations is necessarily selective and brief. Our aim is to identify for the reader critical questions that require an answer to have a plausible cognitive neuroscience of language processing.

Abstract

First paragraph:
MacDonald (2013) proposes that distributional properties of language and processing biases in language comprehension can to a large extent be attributed to consequences of the language production process. In essence, the account is derived from the principle of least effort that was formulated by Zipf, among others (Zipf, 1949; Levelt, 2013). However, in Zipf's view the outcome of the least effort principle was a compromise between least effort for the speaker and least effort for the listener, whereas MacDonald puts most of the burden on the production process.

Abstract

In human face-to-face communication, language comprehension is a multi-modal, situated activity. However, little is known about how we combine information from these different modalities, and how perceived communicative intentions, often signaled through visual signals, such as eye
gaze, may influence this processing. We address this question by simulating a triadic communication context in which a
speaker alternated her gaze between two different recipients. Participants thus viewed speech-only or speech+gesture
object-related utterances when being addressed (direct gaze) or unaddressed (averted gaze). Two object images followed
each message and participants’ task was to choose the object that matched the message. Unaddressed recipients responded significantly slower than addressees for speech-only
utterances. However, perceiving the same speech accompanied by gestures sped them up to a level identical to
that of addressees. That is, when speech processing suffers due to not being addressed, gesture processing remains intact and enhances the comprehension of a speaker’s message

Abstract

The ability to extract word forms from continuous speech is a prerequisite for constructing a vocabulary and emerges in the first year of life. Electrophysiological (ERP) studies of speech segmentation by 9- to 12-month-old listeners in several languages have found a left-localized negativity linked to word onset as a marker of word detection. We report an ERP study showing significant evidence of speech segmentation in Dutch-learning 7-month-olds. In contrast to the left-localized negative effect reported with older infants, the observed overall mean effect had a positive polarity. Inspection of individual results revealed two participant sub-groups: a majority showing a positive-going response, and a minority showing the left negativity observed in older age groups. We retested participants at age three, on vocabulary comprehension and word and sentence production. On every test, children who at 7 months had shown the negativity associated with segmentation of words from speech outperformed those who had produced positive-going brain responses to the same input. The earlier that infants show the left-localized brain responses typically indicating detection of words in speech, the better their early childhood language skills.

Abstract

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.

Abstract

Many authors have recently highlighted the importance of prediction for language comprehension. Pickering & Garrod (P&G) are the first to propose a central role for prediction in language production. This is an intriguing idea, but it is not clear what it means for speakers to predict their own utterances, and how prediction during production can be empirically distinguished from production proper.

Abstract

In everyday communication, people not only use speech but
also hand gestures to convey information. One intriguing
question in gesture research has been why gestures take the
specific form they do. Previous research has identified the
speaker-gesturer’s communicative intent as one factor
shaping the form of iconic gestures. Here we investigate
whether communicative intent also shapes the form of
pointing gestures. In an experimental setting, twenty-four
participants produced pointing gestures identifying a referent
for an addressee. The communicative intent of the speakergesturer
was manipulated by varying the informativeness of
the pointing gesture. A second independent variable was the
presence or absence of concurrent speech. As a function of their communicative intent and irrespective of the presence of speech, participants varied the durations of the stroke and the post-stroke hold-phase of their gesture. These findings add to our understanding of how the communicative context influences the form that a gesture takes.

Abstract

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.

Abstract

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.

Abstract

Human referential communication is often thought as coding-decoding a set of symbols, neglecting that establishing shared meanings requires a computational mechanism powerful enough to mutually negotiate them. Sharing the meaning of a novel symbol might rely on similar conceptual inferences across communicators or on statistical similarities in their sensorimotor behaviors. Using magnetoencephalography, we assess spectral, temporal, and spatial characteristics of neural activity evoked when people generate and understand novel shared symbols during live communicative interactions. Solving those communicative problems induced comparable changes in the spectral profile of neural activity of both communicators and addressees. This shared neuronal up-regulation was spatially localized to the right temporal lobe and the ventromedial prefrontal cortex and emerged already before the occurrence of a specific communicative problem. Communicative innovation relies on neuronal computations that are shared across generating and understanding novel shared symbols, operating over temporal scales independent from transient sensorimotor behavior.

Abstract

A forum devoted to the relationship between music and language begins with an implicit assumption: There is at least one common principle that is central to all human musical systems and all languages, but that is not characteristic of (most) other domains. Why else should these two categories be paired together for analysis? We propose that one candidate for a common principle is their structure. In this chapter, we explore the nature of that structure—and its consequences for psychological and neurological processing mechanisms—within and across these two domains.

Abstract

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.

Abstract

Objective Most rhyme awareness assessments do not encompass measures of the global similarity effect (i.e., children who are able to perform simple rhyme judgments get confused when presented with globally similar non-rhyming pairs). The present study examines the neural nature of this effect by studying the N450 rhyme effect. Methods Behavioral and electrophysiological responses of Dutch pre-literate kindergartners and literate second graders were recorded while they made rhyme judgments of word pairs in three conditions; phonologically rhyming (e.g., wijn-pijn), overlapping non-rhyming (e.g., pen-pijn) and unrelated non-rhyming pairs (e.g., boom-pijn). Results Behaviorally, both groups had difficulty judging overlapping but not rhyming and unrelated pairs. The neural data of second graders showed overlapping pairs were processed in a similar fashion as unrelated pairs; both showed a more negative deflection of the N450 component than rhyming items. Kindergartners did not show a typical N450 rhyme effect. However, some other interesting ERP differences were observed, indicating preliterates are sensitive to rhyme at a certain level. Significance Rhyme judgments of globally similar items rely on the same process as rhyme judgments of rhyming and unrelated items. Therefore, incorporating a globally similar condition in rhyme assessments may lead to a more in-depth measure of early phonological awareness skills. Highlights Behavioral and electrophysiological responses were recorded while (pre)literate children made rhyme judgments of rhyming, overlapping and unrelated words. Behaviorally both groups had difficulty judging overlapping pairs as non-rhyming while overlapping and unrelated neural patterns were similar in literates. Preliterates show a different pattern indicating a developing phonological system.

Abstract

Both emotional words and words focused by information structure can capture attention. This study examined the interplay between emotional salience and information structure in modulating attentional resources in the service of integrating emotional words into sentence context. Event-related potentials (ERPs) to affectively negative, neutral, and positive words, which were either focused or nonfocused in question–answer pairs, were evaluated during sentence comprehension. The results revealed an early negative effect (90–200 ms), a P2 effect, as well as an effect in the N400 time window, for both emotional salience and information structure. Moreover, an interaction between emotional salience and information structure occurred within the N400 time window over right posterior electrodes, showing that information structure influences the semantic integration only for neutral words, but not for emotional words. This might reflect the fact that the linguistic salience of emotional words can override the effect of information structure on the integration of words into context. The interaction provides evidence for attention–emotion interactions at a later stage of processing. In addition, the absence of interaction in the early time window suggests that the processing of emotional information is highly automatic and independent of context. The results suggest independent attention capture systems of emotional salience and information structure at the early stage but an interaction between them at a later stage, during the semantic integration of words.

Abstract

Unlike common nouns, person names refer to unique entities and generally have a referring function. We used event-related potentials to investigate the time course of identifying the emotional meaning of nouns and names. The emotional valence of names and nouns were manipulated separately. The results show early N1 effects in response to emotional valence only for nouns. This might reflect automatic attention directed towards emotional stimuli. The absence of such an effect for names supports the notion that the emotional meaning carried by names is accessed after word recognition and person identification. In addition, both names with negative valence and emotional nouns elicited late positive effects, which have been associated with evaluation of emotional significance. This positive effect started earlier for nouns than for names, but with similar durations. Our results suggest that distinct neural systems are involved in the retrieval of names’ and nouns’ emotional meaning.

Abstract

While syntactic reanalysis has been extensively investigated in psycholinguistics, comparatively little is known about reanalysis in the semantic domain. We used event-related brain potentials (ERPs) to keep track of semantic processes involved in understanding short narratives such as ‘The girl was writing a letter when her friend spilled coffee on the paper’. We hypothesize that these sentences are interpreted in two steps: (1) when the progressive clause is processed, a discourse model is computed in which the goal state (a complete letter) is predicted to hold; (2) when the subordinate clause is processed, the initial representation is recomputed to the effect that, in the final discourse structure, the goal state is not satisfied. Critical sentences evoked larger sustained anterior negativities (SANs) compared to controls, starting around 400 ms following the onset of the sentence-final word, and lasting for about 400 ms. The amplitude of the SAN was correlated with the frequency with which participants, in an offline probe-selection task, responded that the goal state was not attained. Our results raise the possibility that the brain supports some form of non-monotonic recomputation to integrate information which invalidates previously held assumptions.

Abstract

An influential hypothesis regarding the neural basis of the mental lexicon is that semantic representations are neurally implemented as distributed networks carrying sensory, motor and/or more abstract functional information. This work investigates whether the semantic properties of words partly determine the topography of such networks. Subjects performed a visual lexical decision task while their EEG was recorded. We compared the EEG responses to nouns with either visual semantic properties (VIS, referring to colors and shapes) or with auditory semantic properties (AUD, referring to sounds). A time–frequency analysis of the EEG revealed power increases in the theta (4–7 Hz) and lower-beta (13–18 Hz) frequency bands, and an early power increase and subsequent decrease for the alpha (8–12 Hz) band. In the theta band we observed a double dissociation: temporal electrodes showed larger theta power increases in the AUD condition, while occipital leads showed larger theta responses in the VIS condition. The results support the notion that semantic representations are stored in functional networks with a topography that reflects the semantic properties of the stored items, and provide further evidence that oscillatory brain dynamics in the theta frequency range are functionally related to the retrieval of lexical semantic information.

Abstract

Several studies have reported an association between dyslexia and implicit learning deficits. It has been suggested that the weakness in implicit learning observed in dyslexic individuals may be related to sequential processing and implicit sequence learning. In the present article, we review the current literature on implicit learning and dyslexia. We describe a novel, forced-choice structural "mere exposure" artificial grammar learning paradigm and characterize this paradigm in normal readers in relation to the standard grammaticality classification paradigm. We argue that preference classification is a more optimal measure of the outcome of implicit acquisition since in the preference version participants are kept completely unaware of the underlying generative mechanism, while in the grammaticality version, the subjects have, at least in principle, been informed about the existence of an underlying complex set of rules at the point of classification (but not during acquisition). On the basis of the "mere exposure effect," we tested the prediction that the development of preference will correlate with the grammaticality status of the classification items. In addition, we examined the effects of grammaticality (grammatical/nongrammatical) and associative chunk strength (ACS; high/low) on the classification tasks (preference/grammaticality). Using a balanced ACS design in which the factors of grammaticality (grammatical/nongrammatical) and ACS (high/low) were independently controlled in a 2 × 2 factorial design, we confirmed our predictions. We discuss the suitability of this task for further investigation of the implicit learning characteristics in dyslexia.

Abstract

Claims that neuroscientific data do not contribute to our understanding of psychological functions have been made recently. Here I argue that these criticisms are solely based on an analysis of functional magnetic resonance imaging (fMRI) studies. However, fMRI is only one of the methods in the toolkit of cognitive neuroscience. I provide examples from research on event-related brain potentials (ERPs) that have contributed to our understanding of the cognitive architecture of human language functions. In addition, I provide evidence of (possible) contributions from fMRI measurements to our understanding of the functional architecture of language processing. Finally, I argue that a neurobiology of human language that integrates information about the necessary genetic and neural infrastructures will allow us to answer certain questions that are not answerable if all we have is evidence from behavior.

Abstract

This paper focuses on what electrical and magnetic recordings of human brain activity reveal about spoken language understanding. Based on the high temporal resolution of these recordings, a fine-grained temporal profile of different aspects of spoken language comprehension can be obtained. Crucial aspects of speech comprehension are lexical access, selection and semantic integration. Results show that for words spoken in context, there is no ‘magic moment’ when lexical selection ends and semantic integration begins. Irrespective of whether words have early or late recognition points, semantic integration processing is initiated before words can be identified on the basis of the acoustic information alone. Moreover, for one particular event-related brain potential (ERP) component (the N400), equivalent impact of sentence- and discourse-semantic contexts is observed. This indicates that in comprehension, a spoken word is immediately evaluated relative to the widest interpretive domain available. In addition, this happens very quickly. Findings are discussed that show that often an unfolding word can be mapped onto discourse-level representations well before the end of the word. Overall, the time course of the ERP effects is compatible with the view that the different information types (lexical, syntactic, phonological, pragmatic) are processed in parallel and influence the interpretation process incrementally, that is as soon as the relevant pieces of information are available. This is referred to as the immediacy principle.

Abstract

In an event-related potential experiment with Chinese discourses as material, we investigated how and when accentuation influences spoken discourse comprehension in relation to the different information states of the critical words. These words could either provide new or old information. It was shown that variation of accentuation influenced the amplitude of the N400, with a larger amplitude for accented than deaccented words. In addition, there was an interaction between accentuation and information state. The N400 amplitude difference between accented and deaccented new information was smaller than that between accented and deaccented old information. The results demonstrate that, during spoken discourse comprehension, listeners rapidly extract the semantic consequences of accentuation in relation to the previous discourse context. Moreover, our results show that the N400 amplitude can be larger for correct (new,accented words) than incorrect (new, deaccented words) information. This, we argue, proves that the N400 does not react to semantic anomaly per se, but rather to semantic integration load, which is higher for new information.

Abstract

Beim Sprechen und beim Sprachverstehen findet man die Wortbedeutung im Gedächtnis auf und kombiniert sie zu größeren Einheiten (Unifikation). Solche Unifikations-Operationen laufen auf unterschiedlichen Ebenen der Sprachverarbeitung ab. In diesem Beitrag wird ein Rahmen vorgeschlagen, in dem psycholinguistische Modelle mit neurobiologischer Sprachbetrachtung in Verbindung gebracht werden. Diesem Vorschlag zufolge spielt der linke inferiore frontale Gyrus (LIFG) eine bedeutende Rolle bei der Unifi kation

Abstract

Functional imaging studies have demonstrated involvement of the anterior temporal cortex in sentence comprehension. It is unclear, however, whether the anterior temporal cortex is essential for this function.We studied two aspects of sentence comprehension, namely syntactic and prosodic comprehension in temporal lobe epilepsy patients who were candidates for resection of the anterior temporal lobe. Methods: Temporal lobe epilepsy patients (n = 32) with normal (left) language dominance were tested on syntactic and prosodic comprehension before and after removal of the anterior temporal cortex. The prosodic comprehension test was also compared with performance of healthy control subjects (n = 47) before surgery. Results: Overall, temporal lobe epilepsy patients did not differ from healthy controls in syntactic and prosodic comprehension before surgery. They did perform less well on an affective prosody task. Post-operative testing revealed that syntactic and prosodic comprehension did not change after removal of the anterior temporal cortex. Discussion: The unchanged performance on syntactic and prosodic comprehension after removal of the anterior temporal cortex suggests that this area is not indispensable for sentence comprehension functions in temporal epilepsy patients. Potential implications for the postulated role of the anterior temporal lobe in the healthy brain are discussed.

Abstract

Chronic fatigue syndrome (CFS) is a disabling disorder, characterized by persistent or relapsing fatigue. Recent studies have detected a decrease in cortical grey matter volume in patients with CFS, but it is unclear whether this cerebral atrophy constitutes a cause or a consequence of the disease. Cognitive behavioural therapy (CBT) is an effective behavioural intervention for CFS, which combines a rehabilitative approach of a graded increase in physical activity with a psychological approach that addresses thoughts and beliefs about CFS which may impair recovery. Here, we test the hypothesis that cerebral atrophy may be a reversible state that can ameliorate with successful CBT. We have quantified cerebral structural changes in 22 CFS patients that underwent CBT and 22 healthy control participants. At baseline, CFS patients had significantly lower grey matter volume than healthy control participants. CBT intervention led to a significant improvement in health status, physical activity and cognitive performance. Crucially, CFS patients showed a significant increase in grey matter volume, localized in the lateral prefrontal cortex. This change in cerebral volume was related to improvements in cognitive speed in the CFS patients. Our findings indicate that the cerebral atrophy associated with CFS is partially reversed after effective CBT. This result provides an example of macroscopic cortical plasticity in the adult human brain, demonstrating a surprisingly dynamic relation between behavioural state and cerebral anatomy. Furthermore, our results reveal a possible neurobiological substrate of psychotherapeutic treatment.

Abstract

Background: Growing evidence for overlap in the syntactic processing of language and music in non-brain-damaged individuals leads to the question of whether aphasic individuals with grammatical comprehension problems in language also have problems processing structural relations in music.

Aims: The current study sought to test musical syntactic processing in individuals with Broca's aphasia and grammatical comprehension deficits, using both explicit and implicit tasks.

Methods & Procedures: Two experiments were conducted. In the first experiment 12 individuals with Broca's aphasia (and 14 matched controls) were tested for their sensitivity to grammatical and semantic relations in sentences, and for their sensitivity to musical syntactic (harmonic) relations in chord sequences. An explicit task (acceptability judgement of novel sequences) was used. The second experiment, with 9 individuals with Broca's aphasia (and 12 matched controls), probed musical syntactic processing using an implicit task (harmonic priming).

Outcomes & Results: In both experiments the aphasic group showed impaired processing of musical syntactic relations. Control experiments indicated that this could not be attributed to low-level problems with the perception of pitch patterns or with auditory short-term memory for tones.

Conclusions: The results suggest that musical syntactic processing in agrammatic aphasia deserves systematic investigation, and that such studies could help probe the nature of the processing deficits underlying linguistic agrammatism. Methodological suggestions are offered for future work in this little-explored area.

Abstract

We used simultaneously recorded EEG and fMRI to investigate in which areas the BOLD signal correlates with frontal theta power changes, while subjects were quietly lying resting in the scanner with their eyes open. To obtain a reliable estimate of frontal theta power we applied ICA on band-pass filtered (2–9 Hz) EEG data. For each subject we selected the component that best matched the mid-frontal scalp topography associated with the frontal theta rhythm. We applied a time-frequency analysis on this component and used the time course of the frequency bin with the highest overall power to form a regressor that modeled spontaneous fluctuations in frontal theta power. No significant positive BOLD correlations with this regressor were observed. Extensive negative correlations were observed in the areas that together form the default mode network. We conclude that frontal theta activity can be seen as an EEG index of default mode network activity.

Abstract

The discovery of mirror neurons in macaques and of a similar system in humans has provided a new and fertile neurobiological ground for rooting a variety of cognitive faculties. Automatic sensorimotor resonance has been invoked as the key elementary process accounting for disparate (dys)functions, like imitation, ideomotor apraxia, autism, and schizophrenia. In this paper, we provide a critical appraisal of three of these claims that deal with the relationship between language and the motor system. Does language comprehension require the motor system? Was there an evolutionary switch from manual gestures to speech as the primary mode of language? Is human communication explained by automatic sensorimotor resonances? A positive answer to these questions would open the tantalizing possibility of bringing language and human communication within the fold of the motor system. We argue that the available empirical evidence does not appear to support these claims, and their theoretical scope fails to account for some crucial features of the phenomena they are supposed to explain. Without denying the enormous importance of the discovery of mirror neurons, we highlight the limits of their explanatory power for understanding language and communication.

Abstract

The human capacity to implicitly acquire knowledge of structured sequences has recently been investigated in artificial grammar learning using functional magnetic resonance imaging. It was found that the left inferior frontal cortex (IFC; Brodmann's area (BA) 44/45) was related to classification performance. The objective of this study was to investigate whether the IFC (BA 44/45) is causally related to classification of artificial syntactic structures by means of an off-line repetitive transcranial magnetic stimulation (rTMS) paradigm. We manipulated the stimulus material in a 2 × 2 factorial design with grammaticality status and local substring familiarity as factors. The participants showed a reliable effect of grammaticality on classification of novel items after 5days of exposure to grammatical exemplars without performance feedback in an implicit acquisition task. The results show that rTMS of BA 44/45 improves syntactic classification performance by increasing the rejection rate of non-grammatical items and by shortening reaction times of correct rejections specifically after left-sided stimulation. A similar pattern of results is observed in FMRI experiments on artificial syntactic classification. These results suggest that activity in the inferior frontal region is causally related to artificial syntax processing.

Abstract

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.

Abstract

The large majority of humankind is more or less fluent in 2 or even more languages. This raises the fundamental question how the language network in the brain is organized such that the correct target language is selected at a particular occasion. Here we present behavioral and functional magnetic resonance imaging data showing that bilingual processing leads to language conflict in the bilingual brain even when the bilinguals’ task only required target language knowledge. This finding demonstrates that the bilingual brain cannot avoid language conflict, because words from the target and nontarget languages become automatically activated during reading. Importantly, stimulus-based language conflict was found in brain regions in the LIPC associated with phonological and semantic processing, whereas response-based language conflict was only found in the pre-supplementary motor area/anterior cingulate cortex when language conflict leads to response conflicts.

Abstract

Understanding language always occurs within a situational context and, therefore, often implies combining streams of information from different domains and modalities. One such combination is that of spoken language and visual information, which are perceived together in a variety of ways during everyday communication. Here we investigate whether and how words and pictures differ in terms of their neural correlates when they are integrated into a previously built-up sentence context. This is assessed in two experiments looking at the time course (measuring event-related potentials, ERPs) and the locus (using functional magnetic resonance imaging, fMRI) of this integration process. We manipulated the ease of semantic integration of word and/or picture to a previous sentence context to increase the semantic load of processing. In the ERP study, an increased semantic load led to an N400 effect which was similar for pictures and words in terms of latency and amplitude. In the fMRI study, we found overlapping activations to both picture and word integration in the left inferior frontal cortex. Specific activations for the integration of a word were observed in the left superior temporal cortex. We conclude that despite obvious differences in representational format, semantic information coming from pictures and words is integrated into a sentence context in similar ways in the brain. This study adds to the growing insight that the language system incorporates (semantic) information coming from linguistic and extralinguistic domains with the same neural time course and by recruitment of overlapping brain areas.

Abstract

Language is often perceived together with visual information. This raises the question on how the brain integrates information conveyed in visual and/or linguistic format during spoken language comprehension. In this study we investigated the dynamics of semantic integration of visual and linguistic information by means of time-frequency analysis of the EEG signal. A modified version of the N400 paradigm with either a word or a picture of an object being semantically incongruous with respect to the preceding sentence context was employed. Event-Related Potential (ERP) analysis showed qualitatively similar N400 effects for integration of either word or picture. Time-frequency analysis revealed early specific decreases in alpha and gamma band power for linguistic and visual information respectively. We argue that these reflect a rapid context-based analysis of acoustic (word) or visual (picture) form information. We conclude that although full semantic integration of linguistic and visual information occurs through a common mechanism, early differences in oscillations in specific frequency bands reflect the format of the incoming information and, importantly, an early context-based detection of its congruity with respect to the preceding language context

Abstract

In the present study, event-related brain potentials (ERP) were recorded to investigate the role of pitch accent and lexical tone in spoken discourse comprehension. Chinese was used as material to explore the potential difference in the nature and time course of brain responses to sentence meaning as indicated by pitch accent and to lexical meaning as indicated by tone. In both cases, the pitch contour of critical words was varied. The results showed that both inconsistent pitch accent and inconsistent lexical tone yielded N400 effects, and there was no interaction between them. The negativity evoked by inconsistent pitch accent had the some topography as that evoked by inconsistent lexical tone violation, with a maximum over central–parietal electrodes. Furthermore, the effect for the combined violations was the sum of effects for pure pitch accent and pure lexical tone violation. However, the effect for the lexical tone violation appeared approximately 90 ms earlier than the effect of the pitch accent violation. It is suggested that there might be a correspondence between the neural mechanism underlying pitch accent and lexical meaning processing in context. They both reflect the integration of the current information into a discourse context, independent of whether the current information was sentence meaning indicated by accentuation, or lexical meaning indicated by tone. In addition, lexical meaning was processed earlier than sentence meaning conveyed by pitch accent during spoken language processing.

Abstract

In this paper presents evidence of the disputed existence of an electrophysiological marker for the lexical-categorical distinction between open- and closed-class words. Event-related brain potentials were recorded from the scalp while subjects read a story. Separate waveforms were computed for open- and closed-class words. Two aspects of the waveforms could be reliably related to vocabulary class. The first was an early negativity in the 230- to 350-msec epoch, with a bilateral anterior predominance. This negativity was elicited by open- and closed-class words alike, was not affected by word frequency or word length, and had an earlier peak latency for closed-class words. The second was a frontal slow negative shift in the 350- to 500-msec epoch, largest over the left side of the scalp. This late negativity was only elicited by closed-class words. Although the early negativity cannot serve as a qualitative marker of the open- and closed-class distinction, it does reflect the earliest electrophysiological manifestation of the availability of categorical information from the mental lexicon. These results suggest that the brain honors the distinction between open- and closed-class words, in relation to the different roles that they play in on-line sentence processing.

Abstract

The central issue of this study concerns the claim that the processing of gender agreement in online sentence comprehension is a syntactic rather than a conceptual/semantic process. This claim was tested for the grammatical gender agreement in Dutch between the definite article and the noun. Subjects read sentences in which the definite article and the noun had the same gender and sentences in which the gender agreement was violated, While subjects read these sentences, their electrophysiological activity was recorded via electrodes placed on the scalp. Earlier research has shown that semantic and syntactic processing events manifest themselves in different event-related brain potential (ERP) effects. Semantic integration modulates the amplitude of the so-called N400.The P600/SPS is an ERP effect that is more sensitive to syntactic processes. The violation of grammatical gender agreement was found to result in a P600/SPS. For violations in sentence-final position, an additional increase of the N400 amplitude was observed. This N400 effect is interpreted as resulting from the consequence of a syntactic violation for the sentence-final wrap-up. The overall pattern of results supports the claim that the on-line processing of gender agreement information is not a content driven but a syntactic-form driven process.

Abstract

Silent reading and reading aloud of German words and pseudowords were used in a PET study using (15O)butanol to examine the neural correlates of reading and of the phonological conversion of legal letter strings, with or without meaning.
The results of 11 healthy, right-handed volunteers in the age range of 25 to 30 years showed activation of the lingual gyri during silent reading in comparison with viewing a fixation cross. Comparisons between the reading of words and pseudowords suggest the involvement of the middle temporal gyri in retrieving both the phonological and semantic code for words. The reading of pseudowords activates the left inferior frontal gyrus, including the ventral part of Broca’s area, to a larger extent than the reading of words. This suggests that this area might be involved in the sublexical conversion of orthographic input strings into phonological output codes. (Pre)motor areas were found to be activated during both silent reading and reading aloud. On the basis of the obtained activation patterns, it is hypothesized that the articulation of high-frequency syllables requires the retrieval of their concomitant articulatory gestures from the SMA and that the articulation of lowfrequency syllables recruits the left medial premotor cortex.

Abstract

Two recent studies (Coulson et al., 1998;Osterhout et al., 1996)examined the
relationship between the event-related brain potential (ERP) responses to linguistic syntactic anomalies (P600/SPS) and domain-general unexpected events (P300). Coulson et al. concluded that these responses are highly similar, whereas Osterhout et al. concluded that they are distinct. In this comment, we evaluate the relativemerits of these claims. We conclude that the available evidence indicates that the ERP response to syntactic anomalies is at least partially distinct from the ERP response to unexpected anomalies that do not involve a grammatical violation

Abstract

This paper presents electrophysiological data on the on-line processing of open- and closed-class words in patients with Broca’s aphasia with agrammatic comprehension. Event-related brain potentials were recorded from the scalp when Broca patients and nonaphasic control subjects were visually presented with a story in which the words appeared one at a time on the screen. Separate waveforms were computed for open- and closed-class words. The non-aphasic control subjects showed clear differences between the processing of open- and closed-class words in an early (210-375 ms) and a late (400-700 ms) time-window.The early electrophysiological differences reflect the first manifestation of the availability of word-category information from the mental lexicon. The late differences presumably relate to post-lexical semantic and syntactic processing. In contrast to the control subjects, the Broca patients showed no early vocabulary class effect and only a limited late effect. The results suggest that an important factor in the agrammatic comprehension deficit of Broca’s aphasics is a delayed and/or incomplete availability of word-class information.

Abstract

An event-related brain potentials experiment was carried out to examine the interplay of referential and structural factors during sentence processing in discourse. Subjects read (Dutch) sentences beginning like “David told the girl that … ” in short story contexts that had introduced either one or two referents for a critical singular noun phrase (“the girl”). The waveforms showed that within 280 ms after onset of the critical noun the reader had already determined whether the noun phrase had a unique referent in earlier discourse. Furthermore, this referential information was immediately used in parsing the rest of the sentence, which was briefly ambiguous between a complement clause (“ … that there would be some visitors”) and a relative clause (“ … that had been on the phone to hang up”). A consistent pattern of P600/SPS effects elicited by various subsequent disambiguations revealed that a two-referent discourse context had led the parser to initially pursue the relative-clause alternative to a larger extent than a one-referent context. Together, the results suggest that during the processing of sentences in discourse, structural and referential sources of information interact on a word-by-word basis.

Abstract

In two ERP experiments we investigated how and when the language comprehension system relates an incoming word to semantic representations of an unfolding local sentence and a wider discourse. In experiment 1, subjects were presented with short stories. The last sentence of these stories occasionally contained a critical word that, although acceptable in the local sentence context, was semantically anomalous with respect to the wider discourse (e.g., "Jane told the brother that he was exceptionally slow" in a discourse context where he had in fact been very quick). Relative to coherent control words (e.g., "quick"), these discourse-dependent semantic anomalies elicited a large N400 effect that began at about 200-250 ms after word onset. In experiment 2, the same sentences were presented without their original story context. Although the words that had previously been anomalous in discourse still elicited a slightly larger average N400 than the coherent words, the resulting N400 effect was much reduced, showing that the large effect observed in stories was related to the wider discourse. In the same experiment, single sentences that contained a clear local semantic anomaly elicited a standard sentence-dependent N400 effect (e.g., Kutas & Hillyard, 1980). The N400 effects elicited in discourse and in single sentences had the same time course, overall morphology, and scalp distribution. We argue that these findings are most compatible with models of language processing in which there is no fundamental distinction between the integration of a word in its local (sentence-level) and its global (discourse-level) semantic context.

Abstract

We review the implications of recent ERP evidence for when and how grammatical gender agreement constrains sentence parsing. In some theories of parsing, gender is assumed to immediately and categorically block gender-incongruent phrase structure alternatives from being pursued. In other theories, the parser initially ignores gender altogether. The ERP evidence we discuss suggests an intermediate position, in which grammatical gender does not immediately block gender-incongruent phrase structures from being considered, but is used to dispose of them shortly thereafter.

Abstract

Motor-related brain potentials were used to examine the time course of grammatical and phonological processes during noun phrase production in Dutch. In the experiments, participants named colored pictures using a no-determiner noun phrase. On half of the trials a syntactic-phonological classification task had to be performed before naming. Depending on the outcome of the classifications, a left or a right push-button response was given (go trials), or no push-button response was given (no-go trials). Lateralized readiness potentials (LRPs) were derived to test whether syntactic and phonological information affected the motor system at separate moments in time. The results showed that when syntactic information determined the response-hand decision, an LRP developed on no-go trials. However, no such effect was observed when phonological information determined response hand. On the basis of the data, it can be estimated that an additional period of at least 40 ms is needed to retrieve a word's initial phoneme once its lemma has been retrieved. These results provide evidence for the view that during speaking, grammatical processing precedes phonological processing in time.