Publications

Abstract

At least three cognitive brain components are necessary in order for us to be able to produce and comprehend language: a Memory repository for the lexicon, a Unification buffer where lexical information is combined into novel structures, and a Control apparatus presiding over executive function in language. Here we describe the brain networks that support Memory and Unification in semantics. A dynamic account of their interactions is presented, in which a balance between the two components is sought at each word-processing step. We use the theory to provide an explanation of the N400 effect.

Abstract

Many children with Specific Language Impairment (SLI) show impairments in discriminating auditorily presented stimuli. The present study investigates whether these discrimination problems are speech specific or of a general auditory nature. This was studied by using a linguistic and nonlinguistic contrast that were matched for acoustic complexity in an active behavioral task and a passive ERP paradigm, known to elicit the mismatch negativity (MMN). In addition, attention skills and a variety of language skills were measured. Participants were 25 five-year-old Dutch children with SLI having receptive as well as productive language problems and 25 control children with typical speechand language development. At the behavioral level, the SLI group was impaired in discriminating the linguistic contrast as compared to the control group, while both groups were unable to distinguish the non-linguistic contrast. Moreover, the SLI group tended to have impaired attention skills which correlated with performance on most of the language tests. At the neural level, the SLI group, in contrast to the control group, did not show an MMN in response to either the linguistic or nonlinguistic contrast. The MMN data are consistent with an account that relates the symptoms in children with SLI to non-speech processing difficulties.

Abstract

The first objective of this study was to compare the brain network engaged by preference classification and the standard grammaticality classification after implicit artificial syntax acquisition by re-analyzing previously reported event-related fMRI data. The results show that preference and grammaticality classification engage virtually identical brain networks, including Broca’s region, consistent with previous behavioral findings. Moreover, the results showed that the effects related to artificial syntax in Broca’s region were essentially the same when masked with variability related to natural syntax processing in the same participants. The second objective was to explore CNTNAP2-related effects in implicit artificial syntax learning by analyzing behavioral and event-related fMRI data from a subsample. The CNTNAP2 gene has been linked to specific language impairment and is controlled by the FOXP2 transcription factor. CNTNAP2 is expressed in language related brain networks in the developing human brain and the FOXP2–CNTNAP2 pathway provides a mechanistic link between clinically distinct syndromes involving disrupted language. Finally, we discuss the implication of taking natural language to be a neurobiological system in terms of bounded recursion and suggest 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.

Abstract

During face-to-face communication, one does not only hear speech but also see a speaker's communicative hand movements. It has been shown that such hand gestures play an important role in communication where the two modalities influence each other's interpretation. A gesture typically temporally overlaps with coexpressive speech, but the gesture is often initiated before (but not after) the coexpressive speech. The present ERP study investigated what degree of asynchrony in the speech and gesture onsets are optimal for semantic integration of the concurrent gesture and speech. Videos of a person gesturing were combined with speech segments that were either semantically congruent or incongruent with the gesture. Although gesture and speech always overlapped in time, gesture and speech were presented with three different degrees of asynchrony. In the SOA 0 condition, the gesture onset and the speech onset were simultaneous. In the SOA 160 and 360 conditions, speech was delayed by 160 and 360 msec, respectively. ERPs time locked to speech onset showed a significant difference between semantically congruent versus incongruent gesture–speech combinations on the N400 for the SOA 0 and 160 conditions. No significant difference was found for the SOA 360 condition. These results imply that speech and gesture are integrated most efficiently when the differences in onsets do not exceed a certain time span because of the fact that iconic gestures need speech to be disambiguated in a way relevant to the speech context.

Abstract

Whether the brain’s speech-production system is also involved in speech comprehension is a topic of much debate. Research has focused on whether motor areas are involved in listening, but overlap between speaking and listening might occur not only at primary sensory and motor levels, but also at linguistic levels (where semantic, lexical, and syntactic processes occur). Using functional MRI adaptation during speech comprehension and production, we found that the brain areas involved in semantic, lexical, and syntactic processing are mostly the same for speaking and for listening. Effects of primary processing load (indicative of sensory and motor processes) overlapped in auditory cortex and left inferior frontal cortex, but not in motor cortex, where processing load affected activity only in speaking. These results indicate that the linguistic parts of the language system are used for both speaking and listening, but that the motor system does not seem to provide a crucial contribution to listening.

Abstract

Defeasible inferences are inferences that can be revised in the light of new information. Although defeasible inferences are pervasive in everyday communication, little is known about how and when they are processed by the brain. This study examined the electrophysiological signature of defeasible reasoning using a modified version of the suppression task. Participants were presented with conditional inferences (of the type “if p, then q; p, therefore q”) that were preceded by a congruent or a disabling context. The disabling context contained a possible exception or precondition that prevented people from drawing the conclusion. Acceptability of the conclusion was indeed lower in the disabling condition compared to the congruent condition. Further, we found a large sustained negativity at the conclusion of the disabling condition relative to the congruent condition, which started around 250 msec and was persistent throughout the entire epoch. Possible accounts for the observed effect are discussed.

Abstract

Work on animals indicates that BOLD is preferentially sensitive to local field potentials, and that it correlates most strongly with gamma band neuronal synchronization. Here we investigate how the BOLD signal in humans performing a cognitive task is related to neuronal synchronization across different frequency bands. We simultaneously recorded EEG and BOLD while subjects engaged in a visual attention task known to induce sustained changes in neuronal synchronization across a wide range of frequencies. Trial-by-trial BOLD luctuations correlated positively with trial-by-trial fluctuations in high-EEG gamma power (60–80 Hz) and negatively with alpha and beta power. Gamma power on the one hand, and alpha and beta power on the other hand, independently contributed to explaining BOLD variance. These results indicate that the BOLD-gamma coupling observed in animals can be extrapolated to humans performing a task and that neuronal dynamics underlying high- and low-frequency synchronization contribute independently to the BOLD signal.

Abstract

Speakers tend to repeat syntactic structures across sentences, a phenomenon called syntactic priming. Although it has been suggested that repeating syntactic structures should result in speeded responses, previous research has focused on effects in response tendencies. We investigated syntactic priming effects simultaneously in response tendencies and response latencies for active and passive transitive sentences in a picture description task. In Experiment 1, there were priming effects in response tendencies for passives and in response latencies for actives. However, when participants' pre-existing preference for actives was altered in Experiment 2, syntactic priming occurred for both actives and passives in response tendencies as well as in response latencies. This is the first investigation of the effects of structure frequency on both response tendencies and latencies in syntactic priming. We discuss the implications of these data for current theories of syntactic processing.

Abstract

In individuals with ASD, difficulties with language comprehension are most evident when higher-level semantic-pragmatic language processing is required, for instance when context has to be used to interpret the meaning of an utterance. Until now, it is unclear at what level of processing and for what type of context these difficulties in language comprehension occur. Therefore, in the current fMRI study, we investigated the neural correlates of the integration of contextual information during auditory language comprehension in 24 adults with ASD and 24 matched control participants. Different levels of context processing were manipulated by using spoken sentences that were correct or contained either a semantic or world knowledge anomaly. Our findings demonstrated significant differences between the groups in inferior frontal cortex that were only present for sentences with a world knowledge anomaly. Relative to the ASD group, the control group showed significantly increased activation in left inferior frontal gyrus (LIFG) for sentences with a world knowledge anomaly compared to correct sentences. This effect possibly indicates reduced integrative capacities of the ASD group. Furthermore, world knowledge anomalies elicited significantly stronger activation in right inferior frontal gyrus (RIFG) in the control group compared to the ASD group. This additional RIFG activation probably reflects revision of the situation model after new, conflicting information. The lack of recruitment of RIFG is possibly related to difficulties with exception handling in the ASD group.

Abstract

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.

Abstract

To highlight relevant information in dialogues, both wh-question context and pitch accent in answers can be used, such that focused information gains more attention and is processed more elaborately. To evaluate the relative influence of context and pitch accent on the depth of semantic processing, we measured Event-Related Potentials (ERPs) to auditorily presented wh-question-answer pairs. A semantically incongruent word in the answer occurred either in focus or non-focus position as determined by the context, and this word was either accented or unaccented. Semantic incongruency elicited different N400 effects in different conditions. The largest N400 effect was found when the question-marked focus was accented, while the other three conditions elicited smaller N400 effects. The results suggest that context and accentuation interact. Thus accented focused words were processed more deeply compared to conditions where focus and accentuation mismatched, or when the new information had no marking. In addition, there seems to be sex differences in the depth of semantic processing. For males, a significant N400 effect was observed only when the question-marked focus was accented, reduced N400 effects were found in the other dialogues. In contrast, females produced similar N400 effects in all the conditions. These results suggest that regardless of external cues, females tend to engage in more elaborate semantic processing compared to males.

Abstract

We investigated how visual and linguistic information interact in the perception of emotion. We borrowed a phenomenon from film theory which states that presentation of an as such neutral visual scene intensifies the percept of fear or suspense induced by a different channel of information, such as language. Our main aim was to investigate how neutral visual scenes can enhance responses to fearful language content in parts of the brain involved in the perception of emotion. Healthy participants’ brain activity was measured (using functional magnetic resonance imaging) while they read fearful and less fearful sentences presented with or without a neutral visual scene. The main idea is that the visual scenes intensify the fearful content of the language by subtly implying and concretizing what is described in the sentence. Activation levels in the right anterior temporal pole were selectively increased when a neutral visual scene was paired with a fearful sentence, compared to reading the sentence alone, as well as to reading of non-fearful sentences presented with the same neutral scene. We conclude that the right anterior temporal pole serves a binding function of emotional information across domains such as visual and linguistic information.

Abstract

A debated issue in the relationship between language and thought is how our linguistic abilities are involved in understanding the intentions of others (‘mentalizing’). The results of both theoretical and empirical work have been used to argue that linguistic, and more specifically, grammatical, abilities are crucial in representing the mental states of others. Here we contribute to this debate by investigating how damage to the language system influences the generation and understanding of intentional communicative behaviors. Four patients with pervasive language difficulties (severe global or agrammatic aphasia) engaged in an experimentally controlled non-verbal communication paradigm, which required signaling and understanding a communicative message. Despite their profound language problems they were able to engage in recipient design as well as intention recognition, showing similar indicators of mentalizing as have been observed in the neurologically healthy population. Our results show that aspects of the ability to communicate remain present even when core capacities of the language system are dysfunctional

Abstract

Language comprehension involves two basic operations: the retrieval of lexical information (such as phonologic, syntactic, and semantic information) from long-term memory, and the unification of this information into a coherent representation of the overall utterance. Neuroimaging studies using hemo¬dynamic measures such as PET and fMRI have provided detailed information on which areas of the brain are involved in these language-related memory and unification operations. However, much less is known about the dynamics of the brain's language network. This chapter presents a literature review of the oscillatory neuronal dynamics of EEG and MEG data that can be observed during language comprehen¬sion tasks. From a detailed review of this (rapidly growing) literature the following picture emerges: memory retrieval operations are mostly accompanied by increased neuronal synchronization in the theta frequency range (4-7 Hz). Unification operations, in contrast, induce high-frequency neuronal synchro¬nization in the beta (12-30 Hz) and gamma (above 30 Hz) frequency bands. A desynchronization in the (upper) alpha frequency band is found for those studies that use secondary tasks, and seems to correspond with attentional processes, and with the behavioral consequences of the language comprehension process. We conclude that it is possible to capture the dynamics of the brain's language network by a careful analysis of the event-related changes in power and coherence of EEG and MEG data in a wide range of frequencies, in combination with subtle experimental manipulations in a range of language comprehension tasks. It appears then that neuronal synchrony is a mechanism by which the brain integrates the different types of information about language (such as phonological, orthographic, semantic, and syntactic infor¬mation) represented in different brain areas.

Abstract

The human brain supports acquisition mechanisms that extract structural regularities implicitly from experience without the induction of an explicit model. It has been argued that the capacity to generalize to new input is based on the acquisition of abstract representations, which reflect underlying structural regularities in the input ensemble. In this study, we explored the outcome of this acquisition mechanism, and to this end, we investigated the neural correlates of artificial syntactic classification using event-related functional magnetic resonance imaging. The participants engaged once a day during an 8-day period in a short-term memory acquisition task in which consonant-strings generated from an artificial grammar were presented in a sequential fashion without performance feedback. They performed reliably above chance on the grammaticality classification tasks on days 1 and 8 which correlated with a corticostriatal processing network, including frontal, cingulate, inferior parietal, and middle occipital/occipitotemporal regions as well as the caudate nucleus. Part of the left inferior frontal region (BA 45) was specifically related to syntactic violations and showed no sensitivity to local substring familiarity. In addition, the head of the caudate nucleus correlated positively with syntactic correctness on day 8 but not day 1, suggesting that this region contributes to an increase in cognitive processing fluency.

Abstract

We explore the nature of the oscillatory dynamics in the EEG of subjects reading sentences that contain a semantic violation. More specifically, we examine whether increases in theta (≈3–7 Hz) and gamma (around 40 Hz) band power occur in response to sentences that were either semantically correct or contained a semantically incongruent word (semantic violation). ERP results indicated a classical N400 effect. A wavelet-based time-frequency analysis revealed a theta band power increase during an interval of 300–800 ms after critical word onset, at temporal electrodes bilaterally for both sentence conditions, and over midfrontal areas for the semantic violations only. In the gamma frequency band, a predominantly frontal power increase was observed during the processing of correct sentences. This effect was absent following semantic violations. These results provide a characterization of the oscillatory brain dynamics, and notably of both theta and gamma oscillations, that occur during language comprehension.

Abstract

When faced with the noun phrase (NP) versus sentence (S) coordination ambiguity as in, for example, The thief shot the jeweller and the cop hellip, readers prefer the reading with NP-coordination (e.g., "The thief shot the jeweller and the cop yesterday") over one with two conjoined sentences (e.g., "The thief shot the jeweller and the cop panicked"). A corpus study is presented showing that NP-coordinations are produced far more often than S-coordinations, which in frequency-based accounts of parsing might be taken to explain the NP-coordination preference. In addition, we describe an eye-tracking experiment investigating S-coordinated sentences such as Jasper sanded the board and the carpenter laughed, where the poor thematic fit between carpenter and sanded argues against NP-coordination. Our results indicate that information regarding poor thematic fit was used rapidly, but not without leaving some residual processing difficulty. This is compatible with claims that thematic information can reduce but not completely eliminate garden-path effects.

Abstract

This study investigates functional interpretations of left
anterior negativities (LANs), a language-related electroencephalogram effect that has been found for syntactic and morphological violations. We focus on three possible interpretations of LANs caused by the replacement of irregular affixes with regular affixes: misapplication of morphological rules, mismatch of the presented form with analogy-based expectations, and mismatch of the presented form with stored representations. Event-related brain potentials were recorded during the visual presentation of existing and novel Dutch compounds. Existing compounds contained correct or replaced interfixes (dame + s + salons > damessalons vs. *dame + n + salons > *damensalons ‘‘women’s hairdresser salons’’), whereas novel Dutch compounds contained interfixes that were either supported or not supported by analogy to similar existing compounds
(kruidenkelken vs. ?kruidskelken ‘‘herb chalices’’); earlier studies had shown that interfixes are selected by analogy instead of rules. All compounds were presented with correct or incorrect regular plural suffixes (damessalons vs. *damessalonnen). Replacing suffixes or interfixes in existing compounds both led to increased (L)ANs between 400 and 700 msec without any evidence for different scalp distributions for interfixes and suffixes. There was no evidence for a negativity when manipulating the analogical support for interfixes in novel compounds. Together with earlier studies, these results suggest that LANs had been caused by the mismatch of the presented forms with stored forms. We discuss these findings with respect to the single/dual-route debate of morphology and LANs found for the misapplication of syntactic rules.

Abstract

The recognition of a word makes available its semantic and
syntactic properties. Using electrophysiological recordings, we
investigated whether one set of these properties is available
earlier than the other set. Dutch participants saw nouns on a
computer screen and performed push-button responses: In
one task, grammatical gender determined response hand
(left/right) and semantic category determined response execution
(go/no-go). In the other task, response hand depended
on semantic category, whereas response execution depended
on gender. During the latter task, response preparation occurred
on no-go trials, as measured by the lateralized
readiness potential: Semantic information was used for
response preparation before gender information inhibited
this process. Furthermore, an inhibition-related N2 effect
occurred earlier for inhibition by semantics than for inhibition
by gender. In summary, electrophysiological measures
of both response preparation and inhibition indicated that
the semantic word property was available earlier than the
syntactic word property when participants read single
words.

Abstract

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.