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

We employed a parametric psychophysical design in combination with functional imaging to examine the influence of metric changes in perceptual incongruence on perceptual alternation rates and cortical responses. Subjects viewed a bistable stimulus defined by incongruent depth cues; bistability resulted from incongruence between binocular disparity and monocular perspective cues that specify different slants (slant rivalry). Psychophysical results revealed that perceptual alternation rates were positively correlated with the degree of perceived incongruence. Functional imaging revealed systematic increases in activity that paralleled the psychophysical results within anterior intraparietal sulcus, prior to the onset of perceptual alternations. We suggest that this cortical activity predicts the frequency of subsequent alternations, implying a putative causal role for these areas in initiating bistable perception. In contrast, areas implicated in form and depth processing (LOC and V3A) were sensitive to the degree of slant, but failed to show increases in activity when these cues were in conflict.

Abstract

According to theories of embodied cognition, understanding a verb like throw involves unconsciously simulating the action throwing, using areas of the brain that support motor planning. If understanding action words involves mentally simulating our own actions, then the neurocognitive representation of word meanings should differ for people with different kinds of bodies, who perform actions in systematically different ways. In a test of the body-specificity hypothesis (Casasanto, 2009), we used fMRI to compare premotor activity correlated with action verb understanding in right- and left-handers. Right-handers preferentially activated left premotor cortex during lexical decision on manual action verbs (compared with non-manual action verbs), whereas left-handers preferentially activated right premotor areas. This finding helps refine theories of embodied semantics, suggesting that implicit mental simulation during language processing is body-specific: Right and left-handers, who perform actions differently, use correspondingly different areas of the brain for representing action verb meanings.

Abstract

This chapter summarizes the extensive discussions that took place during the Forum as well as the subsequent months thereafter. It assesses current understanding of the neuronal mechanisms that underlie syntactic structure and processing.... It is posited that to understand the neurobiology of syntax, it might be worthwhile to shift the balance from comprehension to syntactic encoding in language production

Abstract

In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, centered on Broca’s region and its homologue, is involved in constructive unification operations during the structure-building process in parsing for comprehension. Tentatively, we provide evidence for a role of the dorsolateral prefrontal cortex centered on BA 9/46 in the control component of the language system. Finally, the left temporo-parietal cortex, in the vicinity of Wernicke’s region, supports the interaction between the syntax of gender agreement and sentence-level semantics.

Abstract

This contribution focuses on the neural infrastructure for parsing and syntactic encoding. From an anatomical point of view, it is argued that Broca's area is an ill-conceived notion. Functionally, Broca's area and adjacent cortex (together Broca's complex) are relevant for language, but not exclusively for this domain of cognition. Its role can be characterized as providing the necessary infrastructure for unification (syntactic and semantic). A general proposal, but with required level of computational detail, is discussed to account for the distribution of labor between different components of the language network in the brain.Arguments are provided for the immediacy principle, which denies a privileged status for syntax in sentence processing. The temporal profile of event-related brain potential (ERP) is suggested to require predictive processing. Finally, since, next to speed, diversity is a hallmark of human languages, the language readiness of the brain might not depend on a universal, dedicated neural machinery for syntax, but rather on a shaping of the neural infrastructure of more general cognitive systems (e.g., memory, unification) in a direction that made it optimally suited for the purpose of communication through language.

Abstract

Language and communication are about the exchange of meaning. A key feature of understanding and producing language is the construction of complex meaning from more elementary semantic building blocks. The functional characteristics of this semantic unification process are revealed by studies using event related brain potentials. These studies have found that word meaning is assembled into compound meaning in not more than 500 ms. World knowledge, information about the speaker, co-occurring visual input and discourse all have an immediate impact on semantic unification, and trigger similar electrophysiological responses as sentence-internal semantic information. Neuroimaging studies show that a network of brain areas, including the left inferior frontal gyrus, the left superior/middle temporal cortex, the left inferior parietal cortex and, to a lesser extent their right hemisphere homologues are recruited to perform semantic unification.

Abstract

How does intention to speak become the action of speaking? It involves the generation of a preverbal message that is tailored to the requirements of a particular language, and through a series of steps, the message is transformed into a linear sequence of speech sounds (1, 2). These steps include retrieving different kinds of information from memory (semantic, syntactic, and phonological), and combining them into larger structures, a process called unification. Despite general agreement about the steps that connect intention to articulation, there is no consensus about their temporal profile or the role of feedback from later steps (3, 4). In addition, since the discovery by the French physician Pierre Paul Broca (in 1865) of the role of the left inferior frontal cortex in speaking, relatively little progress has been made in understanding the neural infrastructure that supports speech production (5). One reason is that the characteristics of natural language are uniquely human, and thus the neurobiology of language lacks an adequate animal model. But on page 445 of this issue, Sahin et al. (6) demonstrate, by recording neuronal activity in the human brain, that different kinds of linguistic information are indeed sequentially processed within Broca's area.

Abstract

Recognizing word boundaries in continuous speech requires detailed knowledge of the native language. In the first year of life, infants acquire considerable word segmentation abilities. Infants at this early stage in word segmentation rely to a large extent on the metrical pattern of their native language, at least in stress-based languages. In Dutch and English (both languages with a preferred trochaic stress pattern), segmentation of strong-weak words develops rapidly between 7 and 10 months of age. Nevertheless, trochaic languages contain not only strong-weak words but also words with a weak-strong stress pattern. In this article, we present electrophysiological evidence of the beginnings of weak-strong word segmentation in Dutch 10-month-olds. At this age, the ability to combine different cues for efficient word segmentation does not yet seem to be completely developed. We provide evidence that Dutch infants still largely rely on strong syllables, even for the segmentation of weak-strong words.

Abstract

Little is known about the genetic contribution to individual differences in neural networks subserving cognition function. In this functional magnetic resonance imaging (fMRI) twin study, we found a significant genetic influence on brain activation in neural networks supporting digit working memory tasks. Participants activating frontal-parietal networks responded faster than individuals relying more on language-related brain networks.There were genetic influences on brain activation in language-relevant brain circuits that were atypical for numerical working memory tasks as such. This suggests that differences in cognition might be related to brain activation patterns that differ qualitatively among individuals.

Abstract

Both local discourse and world knowledge are known to influence sentence processing. We investigated how these two sources of information conspire in language comprehension. Two types of critical sentences, correct and world knowledge anomalies, were preceded by either a neutral or a local context. The latter made the world knowledge anomalies more acceptable or plausible. We predicted that the effect of world knowledge anomalies would be weaker for the local context. World knowledge effects have previously been observed in the left inferior frontal region (Brodmann's area 45/47). In the current study, an effect of world knowledge was present in this region in the neutral context. We also observed an effect in the right inferior frontal gyrus, which was more sensitive to the discourse manipulation than the left inferior frontal gyrus. In addition, the left angular gyrus reacted strongly to the degree of discourse coherence between the context and critical sentence. Overall, both world knowledge and the discourse context affect the process of meaning unification, but do so by recruiting partly different sets of brain areas.

Abstract

The ability to design tailored messages for specific listeners is an important aspect of
human communication. The present study investigates whether a mere belief about an
addressee’s identity influences the generation and production of a communicative message in
a novel, non-verbal communication task. Participants were made to believe they were playing a game with a child or an adult partner, while a confederate acted as both child
and adult partners with matched performance and response times. The participants’ belief
influenced their behavior, spending longer when interacting with the presumed child
addressee, but only during communicative portions of the game, i.e. using time as a tool
to place emphasis on target information. This communicative adaptation attenuated with
experience, and it was related to personality traits, namely Empathy and Need for Cognition
measures. Overall, these findings indicate that novel nonverbal communicative interactions
are selected according to a socio-centric perspective, and they are strongly
influenced by participants’ traits.

Abstract

Human communication has been described as involving the coding-decoding of a conventional symbol system, which could be supported by parts of the human motor system (i.e. the “mirror neurons system”). However, this view does not explain how these conventions could develop in the first place. Here we target the neglected but crucial issue of how people organize their non-verbal behavior to communicate a given intention without pre-established conventions. We have measured behavioral and brain responses in pairs of subjects during communicative exchanges occurring in a real, interactive, on-line social context. In two fMRI studies, we found robust evidence that planning new communicative actions (by a sender) and recognizing the communicative intention of the same actions (by a receiver) relied on spatially overlapping portions of their brains (the right posterior superior temporal sulcus). The response of this region was lateralized to the right hemisphere, modulated by the ambiguity in meaning of the communicative acts, but not by their sensorimotor complexity. These results indicate that the sender of a communicative signal uses his own intention recognition system to make a prediction of the intention recognition performed by the receiver. This finding supports the notion that our communicative abilities are distinct from both sensorimotor processes and language abilities.

Abstract

While autism is one of the most intensively researched psychiatric disorders, little is known about reasoning skills of people with autism. The focus of this study was on defeasible inferences, that is inferences that can be revised in the light of new information. We used a behavioral task to investigate (a) conditional reasoning and (b) the suppression of conditional inferences in high-functioning adults with autism. In the suppression task a possible exception was made salient which could prevent a conclusion from being drawn. We predicted that the autism group would have difficulties dealing with such exceptions because they require mental flexibility to adjust to the context, which is often impaired in autism. The findings confirm our hypothesis that high-functioning adults with autism have a specific difficulty with exception-handling during reasoning. It is suggested that defeasible reasoning is also involved in other cognitive domains. Implications for neural underpinnings of reasoning and autism are discussed.

Abstract

Although people with autism spectrum disorders (ASD) often have severe problems with pragmatic aspects of language, little is known about their pragmatic reasoning. We carried out a behavioral study on highfunctioning adults with autistic disorder (n = 11) and Asperger syndrome (n = 17) and matched controls (n = 28) to investigate whether they are capable of deriving scalar implicatures, which are generally considered to be pragmatic inferences. Participants were presented with underinformative sentences like ‘‘Some sparrows are birds’’. This sentence is logically true, but pragmatically inappropriate if the scalar implicature ‘‘Not all sparrows are birds’’ is derived. The present findings indicate that the combined ASD group was just as likely as controls to derive scalar implicatures, yet there was a difference between participants with autistic disorder and Asperger syndrome, suggesting a potential differentiation between these disorders in pragmatic reasoning. Moreover, our results suggest that verbal intelligence is a constraint for task performance in autistic disorder but not in Asperger syndrome.

Abstract

PET and fMRI experiments have previously shown that several brain regions in the frontal and parietal lobe are involved in working memory maintenance. MEG and EEG experiments have shown parametric increases with load for oscillatory activity in posterior alpha and frontal theta power. In the current study we investigated whether the areas found with fMRI can be associated with these alpha and theta effects by measuring simultaneous EEG and fMRI during a modified Sternberg task This allowed us to correlate EEG at the single trial level with the fMRI BOLD signal by forming a regressor based on single trial alpha and theta
power estimates. We observed a right posterior, parametric alpha power increase, which was functionally related to decreases in BOLD in the primary visual cortex and in the posterior part of the right middle temporal gyrus. We relate this finding to the inhibition of neuronal activity that may interfere with WM maintenance. An observed parametric increase in frontal theta power was correlated to a decrease in BOLD in
regions that together form the default mode network. We did not observe correlations between oscillatory EEG phenomena and BOLD in the traditional WM areas. In conclusion, the study shows that simultaneous EEG fMRI recordings can be successfully used to identify the emergence of functional networks in the brain during the execution of a cognitive task.

Abstract

Sentence comprehension requires the retrieval of single word information from long-term memory, and the integration of this information into multiword representations. The current functional magnetic resonance imaging study explored the hypothesis that the left posterior temporal gyrus supports the retrieval of lexical-syntactic information, whereas left inferior frontal gyrus (LIFG) contributes to syntactic unification. Twenty-eight subjects read sentences and word sequences containing word-category (noun–verb) ambiguous words at critical positions. Regions contributing to the syntactic unification process should show enhanced activation for sentences compared to words, and only within sentences display a larger signal for ambiguous than unambiguous conditions. The posterior LIFG showed exactly this predicted pattern, confirming our hypothesis that LIFG contributes to syntactic unification. The left posterior middle temporal gyrus was activated more for ambiguous than unambiguous conditions (main effect over both sentences and word sequences), as predicted for regions subserving the retrieval of lexical-syntactic information from memory. We conclude that understanding language involves the dynamic interplay between left inferior frontal and left posterior temporal regions.

Abstract

Difficulties with pragmatic aspects of communication are universal across individuals with autism spectrum disorders (ASDs). Here we focused on an aspect of pragmatic language comprehension that is relevant to social interaction in daily life: the integration of speaker characteristics inferred from the voice with the content of a message. Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of the integration of voice-based inferences about the speaker’s age, gender or social background, and sentence content in adults with ASD and matched control participants. Relative to the control group, the ASD group showed increased activation in right inferior frontal gyrus (RIFG; Brodmann area 47) for speakerincongruent sentences compared to speaker-congruent sentences. Given that both groups performed behaviourally at a similar level on a debriefing interview outside the scanner, the increased activation in RIFG for the ASD group was interpreted as being compensatory in nature. It presumably reflects spill-over processing from the language dominant left hemisphere due to higher task demands faced by the participants with ASD when integrating speaker characteristics and the content of a spoken sentence. Furthermore, only the control group showed decreased activation for speaker-incongruent relative to speaker-congruent sentences in right ventral medial prefrontal cortex (vMPFC; Brodmann area 10), including right anterior cingulate cortex (ACC; Brodmann area 24/32). Since vMPFC is involved in self-referential processing related to judgments and inferences about self and others, the absence of such a modulation in vMPFC activation in the ASD group possibly points to atypical default self-referential mental activity in ASD. Our results show that in ASD compensatory mechanisms are necessary in implicit, low-level inferential processes in spoken language understanding. This indicates that pragmatic language problems in ASD are not restricted to high-level inferential processes, but encompass the most basic aspects of pragmatic language processing.

Abstract

When interpreting a message, a listener takes into account several sources of linguistic and extralinguistic information. Here we focused on one particular form of extralinguistic information, certain speaker characteristics as conveyed by the voice. Using functional magnetic resonance imaging, we examined the neural structures involved in the unification of sentence meaning and voice-based inferences about the speaker's age, sex, or social background. We found enhanced activation in the inferior frontal gyrus bilaterally (BA 45/47) during listening to sentences whose meaning was incongruent with inferred speaker characteristics. Furthermore, our results showed an overlap in brain regions involved in unification of speaker-related information and those used for the unification of semantic and world knowledge information [inferior frontal gyrus bilaterally (BA 45/47) and left middle temporal gyrus (BA 21)]. These findings provide evidence for a shared neural unification system for linguistic and extralinguistic sources of information and extend the existing knowledge about the role of inferior frontal cortex as a crucial component for unification during language comprehension.

Abstract

A dominant hypothesis in empirical research on the evolution of language is the following: the fundamental difference between animal and human communication systems is captured by the distinction between regular and more complex non-regular grammars. Studies reporting successful artificial grammar learning of nested recursive structures and imaging studies of the same have methodological shortcomings since they typically allow explicit problem solving strategies and this has been shown to account for the learning effect in subsequent behavioral studies. The present study overcomes these shortcomings by using subtle violations of agreement structure in a preference classification task. In contrast to the studies conducted so far, we use an implicit learning paradigm, allowing the time needed for both abstraction processes and consolidation to take place. Our results demonstrate robust implicit learning of recursively embedded structures (context-free grammar) and recursive structures with cross-dependencies (context-sensitive grammar) in an artificial grammar learning task spanning 9 days. Keywords: Implicit artificial grammar learning; centre embedded; cross-dependency; implicit learning; context-sensitive grammar; context-free grammar; regular grammar; non-regular grammar

Abstract

Next to propositional content, speakers distribute information in their utterances in such a way that listeners can make a distinction between new (focused) and given (non-focused) information. This is referred to as information structure. We measured event-related potentials (ERPs) to explore the role of information structure in semantic processing. Following different questions in wh-question-answer pairs (e.g. What kind of vegetable did Ming buy for cooking today? /Who bought the vegetables for cooking today?), the answer sentences (e.g., Ming bought eggplant/beef to cook today.) contained a critical word, which was either semantically appropriate (eggplant) or inappropriate (beef), and either focus or non-focus. The results showed a full N400 effect only when the critical words were in focus position. In non-focus position a strongly reduced N400 effect was observed, in line with the well-known semantic illusion effect. The results suggest that information structure facilitates semantic processing by devoting more resources to focused information.

Abstract

If motor imagery uses neural structures involved in action execution, then the neural correlates of imagining an action should differ between individuals who tend to execute the action differently. Here we report fMRI data showing that motor imagery is influenced by the way people habitually perform motor actions with their particular bodies; that is, motor imagery is ‘body-specific’ (Casasanto, 2009). During mental imagery for complex hand actions, activation of cortical areas involved in motor planning and execution was left-lateralized in right-handers but right-lateralized in left-handers. We conclude that motor imagery involves the generation of an action plan that is grounded in the participant’s motor habits, not just an abstract representation at the level of the action’s goal. People with different patterns of motor experience form correspondingly different neurocognitive representations of imagined actions.

Abstract

Several studies indicate that both posterior superior temporal sulcus/middle temporal gyrus (pSTS/MTG) and left inferior frontal gyrus (LIFG) are involved in integrating information from different modalities. Here we investigated the respective roles of these two areas in integration of action and language information. We exploited the fact that the semantic relationship between language and different forms of action (i.e. co-speech gestures and pantomimes) is radically different. Speech and co-speech gestures are always produced together, and gestures are not unambiguously understood without speech. On the contrary, pantomimes are not necessarily produced together with speech and can be easily understood without speech. We presented speech together with these two types of communicative hand actions in matching or mismatching combinations to manipulate semantic integration load. Left and right pSTS/MTG were only involved in semantic integration of speech and pantomimes. Left IFG on the other hand was involved in integration of speech and co-speech gestures as well as of speech and pantomimes. Effective connectivity analyses showed that depending upon the semantic relationship between language and action, LIFG modulates activation levels in left pSTS.

This suggests that integration in pSTS/MTG involves the matching of two input streams for which there is a relatively stable common object representation, whereas integration in LIFG is better characterized as the on-line construction of a new and unified representation of the input streams. In conclusion, pSTS/MTG and LIFG are differentially involved in multimodal integration, crucially depending upon the semantic relationship between the input streams.

Abstract

It has been argued that we map observed actions onto our own motor system. Here we added to this issue by investigating whether hand preference influences the neural correlates of action observation of simple, essentially meaningless hand actions. Such an influence would argue for an intricate neural coupling between action production and action observation, which goes beyond effects of motor repertoire or explicit motor training, as has been suggested before. Indeed, parts of the human motor system exhibited a close coupling between action production and action observation. Ventral premotor and inferior and superior parietal cortices showed differential activation for left- and right-handers that was similar during action production as well as during action observation. This suggests that mapping observed actions onto the observer's own motor system is a core feature of action observation - at least for actions that do not have a clear goal or meaning. Basic differences in the way we act upon the world are not only reflected in neural correlates of action production, but can also influence the brain basis of action observation.