Publications

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

We hypothesized that areas in the temporal lobe that have been implicated in the phonological processing of spoken words would also be activated during the generation and phonological processing of imagined speech. We tested this hypothesis using functional magnetic resonance imaging during a behaviorally controlled task of metrical stress evaluation. Subjects were presented with bisyllabic words and had to determine the alternation of strong and weak syllables. Thus, they were required to discriminate between weak-initial words and strong-initial words. In one condition, the stimuli were presented auditorily to the subjects (by headphones). In the other condition the stimuli were presented visually on a screen and subjects were asked to imagine hearing the word. Results showed activation of the supplementary motor area, inferior frontal gyrus (Broca's area) and insula in both conditions. In the superior temporal gyrus (STG) and in the superior temporal sulcus (STS) strong activation was observed during the auditory (perceptual) condition. However, a region located in the posterior part of the STS/STG also responded during the imagery condition. No activation of this same region of the STS was observed during a control condition which also involved processing of visually presented words, but which required a semantic decision from the subject. We suggest that processing of metrical stress, with or without auditory input, relies in part on cortical interface systems located in the posterior part of STS/STG. These results corroborate behavioral evidence regarding phonological loop involvement in auditory–verbal imagery.

Abstract

Oscillatory neuronal dynamics, observed in the human electroencephalogram (EEG) during language processing, have been related to the dynamic formation of functionally coherent networks that serve the role of integrating the different sources of information needed for understanding the linguistic input. To further explore the functional role of oscillatory synchrony during language processing, we quantified event-related EEG power changes induced by the presentation of open-class (OC) words and closed-class (CC) words in a wide range of frequencies (from 1 to 30 Hz), while subjects read a short story. Word presentation induced three oscillatory components: a theta power increase (4–7 Hz), an alpha power decrease (10–12 Hz), and a beta power decrease (16–21 Hz). Whereas the alpha and beta responses showed mainly quantitative differences between the two word classes, the theta responses showed qualitative differences between OC words and CC words: A theta power increase was found over left temporal areas for OC words, but not for CC words. The left temporal theta increase may index the activation of a network involved in retrieving the lexical–semantic properties of the OC items.

Abstract

In speaking and comprehending language, word information is retrieved from memory and combined into larger units (unification). Unification operations take place in parallel at the semantic, syntactic and phonological levels of processing. This article proposes a new framework that connects psycholinguistic models to a neurobiological account of language. According to this proposal the left inferior frontal gyrus (LIFG) plays an important role in unification. Research in other domains of cognition indicates that left prefrontal cortex has the necessary neurobiological characteristics for its involvement in the unification for language. I offer here a psycholinguistic perspective on the nature of language unification and the role of LIFG.

Abstract

Children begin to talk at about age one. The vocabulary they need to do so must be built on perceptual evidence and, indeed, infants begin to recognize spoken words long before they talk. Most of the utterances infants hear, however, are continuous, without pauses between words, so constructing a vocabulary requires them to decompose continuous speech in order to extract the individual words. Here, we present electrophysiological evidence that 10-month-old infants recognize two-syllable words they have previously heard only in isolation when these words are presented anew in continuous speech. Moreover, they only need roughly the first syllable of the word to begin doing this. Thus, prelinguistic infants command a highly efficient procedure for segmentation and recognition of spoken words in the absence of an existing vocabulary, allowing them to tackle effectively the problem of bootstrapping a lexicon out of the highly variable, continuous speech signals in their environment.

Abstract

The chronic fatigue syndrome (CFS) is a disabling disorder of unknown etiology. The symptomatology of CFS (central fatigue, impaired concentration, attention and memory) suggests that this disorder could be related to alterations at the level of the central nervous system. In this study, we have used an automated and unbiased morphometric technique to test whether CFS patients display structural cerebral abnormalities. We mapped structural cerebral morphology and volume in two cohorts of CFS patients (in total 28 patients) and healthy controls (in total 28 controls) from high-resolution structural magnetic resonance images, using voxel-based morphometry. Additionally, we recorded physical activity levels to explore the relation between severity of CFS symptoms and cerebral abnormalities. We observed significant reductions in global gray matter volume in both cohorts of CFS patients, as compared to matched control participants. Moreover, the decline in gray matter volume was linked to the reduction in physical activity, a core aspect of CFS. These findings suggest that the central nervous system plays a key role in the pathophysiology of CFS and point to a new objective and quantitative tool for clinical diagnosis of this disabling disorder.

Abstract

We have used implicit motor imagery to investigate the neural correlates of motor planning independently from actual movements. Subjects were presented with drawings of left or right hands and asked to judge the hand laterality, regardless of the stimulus rotation from its upright orientation. We paired this task with a visual imagery control task, in which subjects were presented with typographical characters and asked to report whether they saw a canonical letter or its mirror image, regardless of its rotation. We measured neurovascular activity with fast event-related fMRI, distinguishing responses parametrically related to motor imagery from responses evoked by visual imagery and other task-related phenomena. By quantifying behavioral and neurovascular correlates of imagery on a trial-by-trial basis, we could discriminate between stimulusrelated, mental rotation-related, and response-related neural activity. We found that specific portions of the posterior parietal and precentral cortex increased their activity as a function of mental rotation only during the motor imagery task. Within these regions, the parietal cortex was visually responsive, whereas the dorsal precentral cortex was not. Response- but not rotation-related activity was found around the left central sulcus (putative primary motor cortex) during both imagery tasks. Our study provides novel evidence on the topography and content of movement representations in the human brain. During intended action, the posterior parietal cortex combines somatosensory and visuomotor information, whereas the dorsal premotor cortex generates the actual motor plan, and the primary motor cortex deals with movement execution. We discuss the relevance of these results in the context of current models of action planning.

Abstract

The authors examined whether people can use their knowledge of the wider discourse rapidly enough to anticipate specific upcoming words as a sentence is unfolding. In an event-related brain potential (ERP) experiment, subjects heard Dutch stories that supported the prediction of a specific noun. To probe whether this noun was anticipated at a preceding indefinite article, stories were continued with a gender-marked adjective whose suffix mismatched the upcoming noun's syntactic gender. Prediction-inconsistent adjectives elicited a differential ERP effect, which disappeared in a no-discourse control experiment. Furthermore, in self-paced reading, prediction-inconsistent adjectives slowed readers down before the noun. These findings suggest that people can indeed predict upcoming words in fluent discourse and, moreover, that these predicted words can immediately begin to participate in incremental parsing operations.

Abstract

An event-related brain potential experiment was carried out to investigate on-line syntactic processing in patients with Broca’s aphasia. Subjects were visually presented with sentences that were either syntactically correct or contained violations of word-category. Three groups of subjects were tested: Broca patients (N=11), non-aphasic patients with a right hemisphere (RH) lesion (N=9), and healthy aged-matched controls (N=15). Both control groups appeared sensitive to the violations of word-category as shown by clear P600/SPS effects. The Broca patients displayed only a very reduced and delayed P600/SPS effect. The results are discussed in the context of a lexicalist parsing model. It is concluded that Broca patients are hindered to detect on-line violations of word-category, if word class information is incomplete or delayed available.