Publications

Abstract

Studies of perception have long shown that the brain adds information to its sensory analysis of the physical environment. A touchstone example for humans is language use: to comprehend a physical signal like speech, the brain must add linguistic knowledge, including syntax. Yet, syntactic rules and representations are widely assumed to be atemporal (i.e., abstract and not bound by time), so they must be translated into time-varying signals for speech comprehension and production. Here, we test 3 different models of the temporal spell-out of syntactic structure against brain activity of people listening to Dutch stories: an integratory bottom-up parser, a predictive top-down parser, and a mildly predictive left-corner parser. These models build exactly the same structure but differ in when syntactic information is added by the brain—this difference is captured in the (temporal distribution of the) complexity metric “incremental node count.” Using temporal response function models with both acoustic and information-theoretic control predictors, node counts were regressed against source-reconstructed delta-band activity acquired with magnetoencephalography. Neural dynamics in left frontal and temporal regions most strongly reflect node counts derived by the top-down method, which postulates syntax early in time, suggesting that predictive structure building is an important component of Dutch sentence comprehension. The absence of strong effects of the left-corner model further suggests that its mildly predictive strategy does not represent Dutch language comprehension well, in contrast to what has been found for English. Understanding when the brain projects its knowledge of syntax onto speech, and whether this is done in language-specific ways, will inform and constrain the development of mechanistic models of syntactic structure building in the brain.

Abstract

Face-to-face communication is not only about ‘what’ is said but also ‘how’ it is said, both in speech and bodily signals. Beat gestures are rhythmic hand movements that typically accompany prosodic prominence in con-versation. Yet, it is still unclear how beat gestures influence language comprehension. On the one hand, beat gestures may share the same functional role of focus markers as prosodic prominence. Accordingly, they would drive attention towards the concurrent speech and highlight its content. On the other hand, beat gestures may trigger inferences of high speaker confidence, generate the expectation that the sentence content is correct and thereby elicit the commitment to the truth of the statement. This study directly disentangled the two hypotheses by evaluating additive and interactive effects of prosodic prominence and beat gestures on language comprehension. Participants watched videos of a speaker uttering sentences and judged whether each sentence was true or false. Sentences sometimes contained a world knowledge violation that may go unnoticed (‘semantic illusion’). Combining beat gestures with prosodic prominence led to a higher degree of semantic illusion, making more world knowledge violations go unnoticed during language comprehension. These results challenge current theories proposing that beat gestures are visual focus markers. To the contrary, they suggest that beat gestures automatically trigger inferences of high speaker confidence and thereby elicit the commitment to the truth of the statement, in line with Grice’s cooperative principle in conversation. More broadly, our findings also highlight the influence of metacognition on language comprehension in face-to-face ommunication.

Abstract

The increased interest in developing next-gen social robots has raised questions about the factors affecting the perception of robot emotions. This study investigates the impact of robot appearances (human-like, mechanical) and face regions (full-face, eye-region) on human perception of robot emotions. A between-subjects user study (N = 305) was conducted where participants were asked to identify the emotions being displayed in videos of robot faces, as well as a human baseline. Our findings reveal three important insights for effective social robot face design in Human-Robot Interaction (HRI): Firstly, robots equipped with a back-projected, fully animated face – regardless of whether they are more human-like or more mechanical-looking – demonstrate a capacity for emotional expression comparable to that of humans. Secondly, the recognition accuracy of emotional expressions in both humans and robots declines when only the eye region is visible. Lastly, within the constraint of only the eye region being visible, robots with more human-like features significantly enhance emotion recognition.

Abstract

To date, research has reliably shown that language can engage and modify perceptual processes in a top-down manner. However, our understanding of the cognitive and neural mechanisms underlying such top-down influences is still under debate. In this review, we provide an overview of findings from literature investigating the organization of semantic networks in the brain (spontaneous engagement of the visual system while processing linguistic information), and linguistic cueing studies (looking at the immediate effects of language on the perception of a visual target), in an effort to isolate such mechanisms. Additionally, we connect the findings from linguistic cueing studies to those reported in (nonlinguistic) literature on priors in perception, in order to find commonalities in neural processes allowing for top-down influences on perception. In doing so, we discuss the effects of language on perception in the context of broader, general cognitive and neural principles. Finally, we propose a way forward in the study of linguistic influences on perception.

Abstract

Prosody underpins various linguistic domains ranging from semantics and syntax to discourse. For instance, prosodic information in the form of lexical stress modifies meanings and, as such, syntactic contexts of words as in Turkish kaz-má "pickaxe" (noun) versus káz-ma "do not dig" (imperative). Likewise, prosody indicates the focused constituent of an utterance as the noun phrase filling the wh-spot in a dialogue like What did you eat? I ate----. In the present study, we investigated the relevance of such prosodic variations for discourse comprehension in Turkish. We aimed at answering how lexical stress and prosodic focus mismatches on critical noun phrases-resulting in grammatical anomalies involving both semantics and syntax and discourse-level anomalies, respectively-affect the perceived correctness of an answer to a question in a given context. To that end, 80 native speakers of Turkish, 40 participating in a psychometric experiment and 40 participating in an EEG experiment, were asked to judge the acceptability of prosodic mismatches that occur either separately or concurrently. Psychometric results indicated that lexical stress mismatch led to a lower correctness score than prosodic focus mismatch, and combined mismatch received the lowest score. Consistent with the psychometric data, EEG results revealed an N400 effect to combined mismatch, and this effect was followed by a P600 response to lexical stress mismatch. Conjointly, these results suggest that every source of prosodic information is immediately available and codetermines the interpretation of an utterance; however, semantically and syntactically relevant lexical stress information is assigned more significance by the language comprehension system compared with prosodic focus information.

Abstract

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

Abstract

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

Abstract

Heschl's gyrus (HG) is a core region of the auditory cortex whose morphology is highly variable across individuals. This variability has been linked to sound perception ability in both speech and music domains. Previous studies show that variations in morphological features of HG, such as cortical surface area and thickness, are heritable. To identify genetic variants that affect HG morphology, we conducted a genome-wide association scan (GWAS) meta-analysis in 3054 healthy individuals using HG surface area and thickness as quantitative traits. None of the single nucleotide polymorphisms (SNPs) showed association P values that would survive correction for multiple testing over the genome. The most significant association was found between right HG area and SNP rs72932726 close to gene DCBLD2 (3q12.1; P=2.77x10(-7)). This SNP was also associated with other regions involved in speech processing. The SNP rs333332 within gene KALRN (3q21.2; P=2.27x10(-6)) and rs143000161 near gene COBLL1 (2q24.3; P=2.40x10(-6)) were associated with the area and thickness of left HG, respectively. Both genes are involved in the development of the nervous system. The SNP rs7062395 close to the X-linked deafness gene POU3F4 was associated with right HG thickness (Xq21.1; P=2.38x10(-6)). This is the first molecular genetic analysis of variability in HG morphology

Abstract

Language and action systems are highly interlinked. A critical piece of evidence is that speech and its accompanying gestures are tightly synchronized. Five experiments were conducted to test 2 hypotheses about the synchronization of speech and gesture. According to the interactive view, there is continuous information exchange between the gesture and speech systems, during both their planning and execution phases. According to the ballistic view, information exchange occurs only during the planning phases of gesture and speech, but the 2 systems become independent once their execution has been initiated. In all experiments, participants were required to point to and/or name a light that had just lit up. Virtual reality and motion tracking technologies were used to disrupt their gesture or speech execution. Participants delayed their speech onset when their gesture was disrupted. They did so even when their gesture was disrupted at its late phase and even when they received only the kinesthetic feedback of their gesture. Also, participants prolonged their gestures when their speech was disrupted. These findings support the interactive view and add new constraints on models of speech and gesture production

Abstract

There are increasing reports that individual variation in behavioral and neurophysiological measures of infant speech processing predicts later language outcomes, and specifically concurrent or subsequent vocabulary size. If such findings are held up under scrutiny, they could both illuminate theoretical models of language development and contribute to the prediction of communicative disorders. A qualitative, systematic review of this emergent literature illustrated the variety of approaches that have been used and highlighted some conceptual problems regarding the measurements. A quantitative analysis of the same data established that the bivariate relation was significant, with correlations of similar strength to those found for well-established nonlinguistic predictors of language. Further exploration of infant speech perception predictors, particularly from a methodological perspective, is recommended.

Abstract

Numerous experiments show that space and musical pitch are
closely linked in people's minds. However, the exact nature of
space-pitch associations and their neuronal underpinnings are
not well understood. In an fMRI experiment we investigated
different types of spatial representations that may underlie
musical pitch. Participants judged stimuli that varied in
spatial height in both the visual and tactile modalities, as well
as auditory stimuli that varied in pitch height. In order to
distinguish between unimodal and multimodal spatial bases of
musical pitch, we examined whether pitch activations were
present in modality-specific (visual or tactile) versus
multimodal (visual and tactile) regions active during spatial
height processing. Judgments of musical pitch were found to
activate unimodal visual areas, suggesting that space-pitch
associations may involve modality-specific spatial
representations, supporting a key assumption of embodied
theories of metaphorical mental representation.

Abstract

The left and right sides of the human brain are specialized for different kinds of information processing, and much of our cognition is lateralized to an extent towards one side or the other. Handedness is a reflection of nervous system lateralization. Roughly ten percent of people are mixed- or left-handed, and they show an elevated rate of reductions or reversals of some cerebral functional asymmetries compared to right-handers. Brain anatomical correlates of left-handedness have also been suggested. However, the relationships of left-handedness to brain structure and function remain far from clear. We carried out a comprehensive analysis of cortical surface area differences between 106 left-handed subjects and 1960 right-handed subjects, measured using an automated method of regional parcellation (FreeSurfer, Destrieux atlas). This is the largest study sample that has so far been used in relation to this issue. No individual cortical region showed an association with left-handedness that survived statistical correction for multiple testing, although there was a nominally significant association with the surface area of a previously implicated region: the left precentral sulcus. Identifying brain structural correlates of handedness may prove useful for genetic studies of cerebral asymmetries, as well as providing new avenues for the study of relations between handedness, cerebral lateralization and cognition.

Abstract

Functional and anatomical asymmetries are prevalent features of the human brain, linked to gender, handedness, and cognition. However, little is known about the neurodevelopmental processes involved. In zebrafish, asymmetries arise in the diencephalon before extending within the central nervous system. We aimed to identify genes involved in the development of subtle, left-right volumetric asymmetries of human subcortical structures using large datasets. We first tested the feasibility of measuring left-right volume differences in such large-scale samples, as assessed by two automated methods of subcortical segmentation (FSL|FIRST and FreeSurfer), using data from 235 subjects who had undergone MRI twice. We tested the agreement between the first and second scan, and the agreement between the segmentation methods, for measures of bilateral volumes of six subcortical structures and the hippocampus, and their volumetric asymmetries. We also tested whether there were biases introduced by left-right differences in the regional atlases used by the methods, by analyzing left-right flipped images. While many bilateral volumes were measured well (scan-rescan r = 0.6-0.8), most asymmetries, with the exception of the caudate nucleus, showed lower repeatabilites. We meta-analyzed genome-wide association scan results for caudate nucleus asymmetry in a combined sample of 3,028 adult subjects but did not detect associations at genome-wide significance (P < 5 × 10-8). There was no enrichment of genetic association in genes involved in left-right patterning of the viscera. Our results provide important information for researchers who are currently aiming to carry out large-scale genome-wide studies of subcortical and hippocampal volumes, and their asymmetries

Abstract

Current views on the neurobiological underpinnings of language are discussed that deviate in a number of ways from the classical Wernicke–Lichtheim–Geschwind model. More areas than Broca's and Wernicke's region are involved in language. Moreover, a division along the axis of language production and language comprehension does not seem to be warranted. Instead, for central aspects of language processing neural infrastructure is shared between production and comprehension. Three different accounts of the role of Broca's area in language are discussed. Arguments are presented in favor of a dynamic network view, in which the functionality of a region is co-determined by the network of regions in which it is embedded at particular moments in time. Finally, core regions of language processing need to interact with other networks (e.g. the attentional networks and the ToM network) to establish full functionality of language and communication.

Abstract

A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.

Abstract

The use of virtual reality (VR) as a methodological tool is
becoming increasingly popular in behavioural research due
to its seemingly limitless possibilities. This new method has
not been used frequently in the field of psycholinguistics,
however, possibly due to the assumption that humancomputer
interaction does not accurately reflect human-human
interaction. In the current study we compare participants’
language behaviour in a syntactic priming task with human
versus avatar partners. Our study shows comparable priming
effects between human and avatar partners (Human: 12.3%;
Avatar: 12.6% for passive sentences) suggesting that VR is a
valid platform for conducting language research and studying
dialogue interactions.

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 different modalities during comprehension, and how perceived communicative intentions, often signaled through visual signals, influence this process. We explored this question by simulating a multi-party communication context in which a speaker alternated her gaze between two recipients. Participants viewed speech-only or speech + gesture object-related messages when being addressed (direct gaze) or unaddressed (gaze averted to other participant). They were then asked to choose which of two object images matched the speaker’s preceding message. Unaddressed recipients responded significantly more slowly than addressees for speech-only utterances. However, perceiving the same speech accompanied by gestures sped unaddressed recipients up to a level identical to that of addressees. That is, when unaddressed recipients’ speech processing suffers, gestures can enhance the comprehension of a speaker’s message. We discuss our findings with respect to two hypotheses attempting to account for how social eye gaze may modulate multi-modal language comprehension.

Abstract

Most words that infants hear occur within fluent speech. To compile a vocabulary, infants therefore need to segment words from speech contexts. This study is the first to investigate whether infants (here: 10-month-olds) can recognize words when both initial exposure and test presentation are in continuous speech. Electrophysiological evidence attests that this indeed occurs: An increased extended negativity (word recognition effect) appears for familiarized target words relative to control words. This response proved constant at the individual level: Only infants who showed this negativity at test had shown such a response, within six repetitions after first occurrence, during familiarization.

Abstract

Morphology is the aspect of language concerned with the internal structure of words. In the past decades, a large body of masked priming (behavioral and neuroimaging) data has suggested that the visual word recognition system automatically decomposes any morphologically complex word into a stem and its constituent morphemes. Yet the reliance of morphology on other reading processes (e.g., orthography and semantics), as well as its underlying neuronal mechanisms are yet to be determined. In the current magnetoencephalography study, we addressed morphology from the perspective of the unification framework, that is, by applying the Hold/Release paradigm, morphological unification was simulated via the assembly of internal morphemic units into a whole word. Trials representing real words were divided into words with a transparent (true) or a nontransparent (pseudo) morphological relationship. Morphological unification of truly suffixed words was faster and more accurate and additionally enhanced induced oscillations in the narrow gamma band (60–85 Hz, 260–440 ms) in the left posterior occipitotemporal junction. This neural signature could not be explained by a mere automatic lexical processing (i.e., stem perception), but more likely it related to a semantic access step during the morphological unification process. By demonstrating the validity of unification at the morphological level, this study contributes to the vast empirical evidence on unification across other language processes. Furthermore, we point out that morphological unification relies on the retrieval of lexical semantic associations via induced gamma band oscillations in a cerebral hub region for visual word form processing.

Abstract

We report on an fMRI syntactic priming experiment in which we measure brain activity for participants who communicate with another participant outside the scanner. We investigated whether syntactic processing during overt language production and comprehension is influenced by having a (shared) goal to communicate. Although theory suggests this is true, the nature of this influence remains unclear. Two hypotheses are tested: i. syntactic priming effects (fMRI and RT) are stronger for participants in the communicative context than for participants doing the same experiment in a non-communicative context, and ii. syntactic priming magnitude (RT) is correlated with the syntactic priming magnitude of the speaker’s communicative partner. Results showed that across conditions, participants were faster to produce sentences with repeated syntax, relative to novel syntax. This behavioral result converged with the fMRI data: we found repetition suppression effects in the left insula extending into left inferior frontal gyrus (BA 47/45), left middle temporal gyrus (BA 21), left inferior parietal cortex (BA 40), left precentral gyrus (BA 6), bilateral precuneus (BA 7), bilateral supplementary motor cortex (BA 32/8) and right insula (BA 47). We did not find support for the first hypothesis: having a communicative intention does not increase the magnitude of syntactic priming effects (either in the brain or in behavior) per se. We did find support for the second hypothesis: if speaker A is strongly/weakly primed by speaker B, then speaker B is primed by speaker A to a similar extent. We conclude that syntactic processing is influenced by being in a communicative context, and that the nature of this influence is bi-directional: speakers are influenced by each other.

Abstract

Speakers sometimes repeat syntactic structures across sentences, a phenomenon called syntactic priming. We investigated the influence of verb-bound syntactic preferences on syntactic priming effects in response choices and response latencies for German ditransitive sentences. In the response choices we found inverse preference effects: There were stronger syntactic priming effects for primes in the less preferred structure, given the syntactic preference of the prime verb. In the response latencies we found positive preference effects: There were stronger syntactic priming effects for primes in the more preferred structure, given the syntactic preference of the prime verb. These findings provide further support for the idea that syntactic processing is lexically guided.

Abstract

An ability to decode semantic information from fMRI spatial patterns has been demonstrated in previous studies mostly for 1 specific input modality. In this study, we aimed to decode semantic category independent of the modality in which an object was presented. Using a searchlight method, we were able to predict the stimulus category from the data while participants performed a semantic categorization task with 4 stimulus modalities (spoken and written names, photographs, and natural sounds). Significant classification performance was achieved in all 4 modalities. Modality-independent decoding was implemented by training and testing the searchlight method across modalities. This allowed the localization of those brain regions, which correctly discriminated between the categories, independent of stimulus modality. The analysis revealed large clusters of voxels in the left inferior temporal cortex and in frontal regions. These voxels also allowed category discrimination in a free recall session where subjects recalled the objects in the absence of external stimuli. The results show that semantic information can be decoded from the fMRI signal independently of the input modality and have clear implications for understanding the functional mechanisms of semantic memory.

Abstract

How can we understand each other during communicative interactions? An influential suggestion holds that communicators are primed by each other’s behaviors, with associative mechanisms automatically coordinating the production of communicative signals and the comprehension of their meanings. An alternative suggestion posits that mutual understanding requires shared conceptualizations of a signal’s use, i.e., “conceptual pacts” that are abstracted away from specific experiences. Both accounts predict coherent neural dynamics across communicators, aligned either to the occurrence of a signal or to the dynamics of conceptual pacts. Using coherence spectral-density analysis of cerebral activity simultaneously measured in pairs of communicators, this study shows that establishing mutual understanding of novel signals synchronizes cerebral dynamics across communicators’ right temporal lobes. This interpersonal cerebral coherence occurred only within pairs with a shared communicative history, and at temporal scales independent from signals’ occurrences. These findings favor the notion that meaning emerges from shared conceptualizations of a signal’s use.

Abstract

Despite the ambiguity inherent in human communication, people are remarkably efficient in establishing mutual understanding. Studying how people communicate in novel settings provides a window into the mechanisms supporting the human competence to rapidly generate and understand novel shared symbols, a fundamental property of human communication. Previous work indicates that the right posterior superior temporal sulcus (pSTS) is involved when people understand the intended meaning of novel communicative actions. Here, we set out to test whether normal functioning of this cerebral structure is required for understanding novel communicative actions using inhibitory low-frequency repetitive transcranial magnetic stimulation (rTMS). A factorial experimental design contrasted two tightly matched stimulation sites (right pSTS vs. left MT+, i.e. a contiguous homotopic task-relevant region) and tasks (a communicative task vs. a visual tracking task that used the same sequences of stimuli). Overall task performance was not affected by rTMS, whereas changes in task performance over time were disrupted according to TMS site and task combinations. Namely, rTMS over pSTS led to a diminished ability to improve action understanding on the basis of recent communicative history, while rTMS over MT+ perturbed improvement in visual tracking over trials. These findings qualify the contributions of the right pSTS to human communicative abilities, showing that this region might be necessary for incorporating previous knowledge, accumulated during interactions with a communicative partner, to constrain the inferential process that leads to action understanding.

Abstract

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

Abstract

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