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According to embodied theories of language, people understand a verb like "throw", at least in part, by mentally simulating "throwing". This implicit simulation is often assumed to be similar or identical to motor imagery. Here we used fMRI to test whether implicit simulations of actions during language understanding involve the same cortical motor regions as explicit motor imagery. Healthy participants were presented with verbs related to hand actions (e.g., to "throw") and nonmanual actions (e.g., to "kneel"). They either read these verbs (lexical decision task) or actively imagined performing the actions named by the verbs (imagery task). Primary motor cortex showed effector-specific activation during imagery, but not during lexical decision. Parts of premotor cortex distinguished manual from nonmanual actions during both lexical decision and imagery, but there was no overlap or correlation between regions activated during the two tasks. These dissociations suggest that implicit simulation and explicit imagery cued by action verbs may involve different types of motor representations and that the construct of "mental simulation" should be distinguished from "mental imagery" in embodied theories of language.

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., (© 2024 Massachusetts Institute of Technology.)

Words are not processed in isolation; instead, they are commonly embedded in phrases and sentences. The sentential context influences the perception and processing of a word. However, how this is achieved by brain processes and whether predictive mechanisms underlie this process remain a debated topic. Here, we employed an experimental paradigm in which we orthogonalized sentence context constraints and predictive validity, which was defined as the ratio of congruent to incongruent sentence endings within the experiment. While recording electroencephalography, participants read sentences with three levels of sentential context constraints (high, medium, and low). Participants were also separated into two groups that differed in their ratio of valid congruent to incongruent target words that could be predicted from the sentential context. For both groups, we investigated modulations of alpha power before, and N400 amplitude modulations after target word onset. The results reveal that the N400 amplitude gradually decreased with higher context constraints and cloze probability. In contrast, alpha power was not significantly affected by context constraint. Neither the N400 nor alpha power were significantly affected by changes in predictive validity., (© 2023 Massachusetts Institute of Technology.)

This neuroimaging study investigated the neural infrastructure of sentence-level language production. We compared brain activation patterns, as measured with BOLD-fMRI, during production of sentences that differed in verb argument structures (intransitives, transitives, ditransitives) and the lexical status of the verb (known verbs or pseudoverbs). The experiment consisted of 30 mini-blocks of six sentences each. Each mini-block started with an example for the type of sentence to be produced in that block. On each trial in the mini-blocks, participants were first given the (pseudo-)verb followed by three geometric shapes to serve as verb arguments in the sentences. Production of sentences with known verbs yielded greater activation compared to sentences with pseudoverbs in the core language network of the left inferior frontal gyrus, the left posterior middle temporal gyrus, and a more posterior middle temporal region extending into the angular gyrus, analogous to effects observed in language comprehension. Increasing the number of verb arguments led to greater activation in an overlapping left posterior middle temporal gyrus/angular gyrus area, particularly for known verbs, as well as in the bilateral precuneus. Thus, producing sentences with more complex structures using existing verbs leads to increased activation in the language network, suggesting some reliance on memory retrieval of stored lexical-syntactic information during sentence production. This study thus provides evidence from sentence-level language production in line with functional models of the language network that have so far been mainly based on single-word production, comprehension, and language processing in aphasia.

Recent neuroimaging evidence suggests that the frequency of entrained oscillations in auditory cortices influences the perceived duration of speech segments, impacting word perception [Kösem, A., Bosker, H. R., Takashima, A., Meyer, A., Jensen, O., & Hagoort, P. Neural entrainment determines the words we hear. Current Biology , 28 , 2867-2875, 2018]. We further tested the causal influence of neural entrainment frequency during speech processing, by manipulating entrainment with continuous transcranial alternating current stimulation (tACS) at distinct oscillatory frequencies (3 and 5.5 Hz) above the auditory cortices. Dutch participants listened to speech and were asked to report their percept of a target Dutch word, which contained a vowel with an ambiguous duration. Target words were presented either in isolation (first experiment) or at the end of spoken sentences (second experiment). We predicted that the tACS frequency would influence neural entrainment and therewith how speech is perceptually sampled, leading to a perceptual overestimation or underestimation of the vowel's duration. Whereas results from Experiment 1 did not confirm this prediction, results from Experiment 2 suggested a small effect of tACS frequency on target word perception: Faster tACS leads to more long-vowel word percepts, in line with the previous neuroimaging findings. Importantly, the difference in word perception induced by the different tACS frequencies was significantly larger in Experiment 1 versus Experiment 2, suggesting that the impact of tACS is dependent on the sensory context. tACS may have a stronger effect on spoken word perception when the words are presented in continuous speech as compared to when they are isolated, potentially because prior (stimulus-induced) entrainment of brain oscillations might be a prerequisite for tACS to be effective.

Perceiving speech requires the integration of different speech cues, that is, formants. When the speech signal is split so that different cues are presented to the right and left ear (dichotic listening), comprehension requires the integration of binaural information. Based on prior electrophysiological evidence, we hypothesized that the integration of dichotically presented speech cues is enabled by interhemispheric phase synchronization between primary and secondary auditory cortex in the gamma frequency band. We tested this hypothesis by applying transcranial alternating current stimulation (TACS) bilaterally above the superior temporal lobe to induce or disrupt interhemispheric gamma-phase coupling. In contrast to initial predictions, we found that gamma TACS applied in-phase above the two hemispheres (interhemispheric lag 0°) perturbs interhemispheric integration of speech cues, possibly because the applied stimulation perturbs an inherent phase lag between the left and right auditory cortex. We also observed this disruptive effect when applying antiphasic delta TACS (interhemispheric lag 180°). We conclude that interhemispheric phase coupling plays a functional role in interhemispheric speech integration. The direction of this effect may depend on the stimulation frequency.

Using magnetoencephalography, the current study examined gamma activity associated with language prediction. Participants read high- and low-constraining sentences in which the final word of the sentence was either expected or unexpected. Although no consistent gamma power difference induced by the sentence-final words was found between the expected and unexpected conditions, the correlation of gamma power during the prediction and activation intervals of the sentence-final words was larger when the presented words matched with the prediction compared with when the prediction was violated or when no prediction was available. This suggests that gamma magnitude relates to the match between predicted and perceived words. Moreover, the expected words induced activity with a slower gamma frequency compared with that induced by unexpected words. Overall, the current study establishes that prediction is related to gamma power correlations and a slowing of the gamma frequency.

Readers and listeners actively predict upcoming words during language processing. These predictions might serve to support the unification of incoming words into sentence context and thus rely on interactions between areas in the language network. In the current magnetoencephalography study, participants read sentences that varied in contextual constraints so that the predictability of the sentence-final words was either high or low. Before the sentence-final words, we observed stronger alpha power suppression for the highly compared with low constraining sentences in the left inferior frontal cortex, left posterior temporal region, and visual word form area. Importantly, the temporal and visual word form area alpha power correlated negatively with left frontal gamma power for the highly constraining sentences. We suggest that the correlation between alpha power decrease in temporal language areas and left prefrontal gamma power reflects the initiation of an anticipatory unification process in the language network.

Listeners interpret utterances by integrating information from multiple sources including word level semantics and world knowledge. When the semantics of an expression is inconsistent with their knowledge about the world, the listener may have to search through the conceptual space for alternative possible world scenarios that can make the expression more acceptable. Such cognitive exploration requires considerable computational resources and might depend on motivational factors. This study explores whether and how oxytocin, a neuropeptide known to influence social motivation by reducing social anxiety and enhancing affiliative tendencies, can modulate the integration of world knowledge and sentence meanings. The study used a between-participant double-blind randomized placebo-controlled design. Semantic integration, indexed with magnetoencephalography through the N400m marker, was quantified while 45 healthy male participants listened to sentences that were either congruent or incongruent with facts of the world, after receiving intranasally delivered oxytocin or placebo. Compared with congruent sentences, world knowledge incongruent sentences elicited a stronger N400m signal from the left inferior frontal and anterior temporal regions and medial pFC (the N400m effect) in the placebo group. Oxytocin administration significantly attenuated the N400m effect at both sensor and cortical source levels throughout the experiment, in a state-like manner. Additional electrophysiological markers suggest that the absence of the N400m effect in the oxytocin group is unlikely due to the lack of early sensory or semantic processing or a general downregulation of attention. These findings suggest that oxytocin drives listeners to resolve challenges of semantic integration, possibly by promoting the cognitive exploration of alternative possible world scenarios.

Communication is facilitated when listeners allocate their attention to important information (focus) in the message, a process called "information structure." Linguistic cues like the preceding context and pitch accent help listeners to identify focused information. In multimodal communication, relevant information can be emphasized by nonverbal cues like beat gestures, which represent rhythmic nonmeaningful hand movements. Recent studies have found that linguistic and nonverbal attention cues are integrated independently in single sentences. However, it is possible that these two cues interact when information is embedded in context, because context allows listeners to predict what information is important. In an ERP study, we tested this hypothesis and asked listeners to view videos capturing a dialogue. In the critical sentence, focused and nonfocused words were accompanied by beat gestures, grooming hand movements, or no gestures. ERP results showed that focused words are processed more attentively than nonfocused words as reflected in an N1 and P300 component. Hand movements also captured attention and elicited a P300 component. Importantly, beat gesture and focus interacted in a late time window of 600-900 msec relative to target word onset, giving rise to a late positivity when nonfocused words were accompanied by beat gestures. Our results show that listeners integrate beat gesture with the focus of the message and that integration costs arise when beat gesture falls on nonfocused information. This suggests that beat gestures fulfill a unique focusing function in multimodal discourse processing and that they have to be integrated with the information structure of the message.

In everyday human communication, we often express our communicative intentions by manually pointing out referents in the material world around us to an addressee, often in tight synchronization with referential speech. This study investigated whether and how the kinematic form of index finger pointing gestures is shaped by the gesturer's communicative intentions and how this is modulated by the presence of concurrently produced speech. Furthermore, we explored the neural mechanisms underpinning the planning of communicative pointing gestures and speech. Two experiments were carried out in which participants pointed at referents for an addressee while the informativeness of their gestures and speech was varied. Kinematic and electrophysiological data were recorded online. It was found that participants prolonged the duration of the stroke and poststroke hold phase of their gesture to be more communicative, in particular when the gesture was carrying the main informational burden in their multimodal utterance. Frontal and P300 effects in the ERPs suggested the importance of intentional and modality-independent attentional mechanisms during the planning phase of informative pointing gestures. These findings contribute to a better understanding of the complex interplay between action, attention, intention, and language in the production of pointing gestures, a communicative act core to human interaction.

During sentence level language comprehension, semantic and syntactic unification are functionally distinct operations. Nevertheless, both recruit roughly the same brain areas (spatially overlapping networks in the left frontotemporal cortex) and happen at the same time (in the first few hundred milliseconds after word onset). We tested the hypothesis that semantic and syntactic unification are segregated by means of neuronal synchronization of the functionally relevant networks in different frequency ranges: gamma (40 Hz and up) for semantic unification and lower beta (10-20 Hz) for syntactic unification. EEG power changes were quantified as participants read either correct sentences, syntactically correct though meaningless sentences (syntactic prose), or sentences that did not contain any syntactic structure (random word lists). Other sentences contained either a semantic anomaly or a syntactic violation at a critical word in the sentence. Larger EEG gamma-band power was observed for semantically coherent than for semantically anomalous sentences. Similarly, beta-band power was larger for syntactically correct sentences than for incorrect ones. These results confirm the existence of a functional dissociation in EEG oscillatory dynamics during sentence level language comprehension that is compatible with the notion of a frequency-based segregation of syntactic and semantic unification.

This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, "The boy fell asleep and never woke up again," and their neutral counterparts "The boy stood up and grabbed his bag." This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.

Perception does not function as an isolated module but is tightly linked with other cognitive functions. Several studies have demonstrated an influence of language on motion perception, but it remains debated at which level of processing this modulation takes place. Some studies argue for an interaction in perceptual areas, but it is also possible that the interaction is mediated by "language areas" that integrate linguistic and visual information. Here, we investigated whether language-perception interactions were specific to the language-dominant left hemisphere by comparing the effects of language on visual material presented in the right (RVF) and left visual fields (LVF). Furthermore, we determined the neural locus of the interaction using fMRI. Participants performed a visual motion detection task. On each trial, the visual motion stimulus was presented in either the LVF or in the RVF, preceded by a centrally presented word (e.g., "rise"). The word could be congruent, incongruent, or neutral with regard to the direction of the visual motion stimulus that was presented subsequently. Participants were faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. Interestingly, the speed benefit was present only for motion stimuli that were presented in the RVF. We observed a neural counterpart of the behavioral facilitation effects in the left middle temporal gyrus, an area involved in semantic processing of verbal material. Together, our results suggest that semantic information about motion retrieved in language regions may automatically modulate perceptual decisions about motion.

In this study, we explore the possibility to predict the semantic category of words from brain signals in a free word generation task. Participants produced single words from different semantic categories in a modified semantic fluency task. A Bayesian logistic regression classifier was trained to predict the semantic category of words from single-trial MEG data. Significant classification accuracies were achieved using sensor-level MEG time series at the time interval of conceptual preparation. Semantic category prediction was also possible using source-reconstructed time series, based on minimum norm estimates of cortical activity. Brain regions that contributed most to classification on the source level were identified. These were the left inferior frontal gyrus, left middle frontal gyrus, and left posterior middle temporal gyrus. Additionally, the temporal dynamics of brain activity underlying the semantic preparation during word generation was explored. These results provide important insights about central aspects of language production.

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 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 the 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.

Research from the past decade has shown that understanding the meaning of words and utterances (i.e., abstracted symbols) engages the same systems we used to perceive and interact with the physical world in a content-specific manner. For example, understanding the word "grasp" elicits activation in the cortical motor network, that is, part of the neural substrate involved in planned and executing a grasping action. In the embodied literature, cortical motor activation during language comprehension is thought to reflect motor simulation underlying conceptual knowledge [note that outside the embodied framework, other explanations for the link between action and language are offered, e.g., Mahon, B. Z., & Caramazza, A. A critical look at the embodied cognition hypothesis and a new proposal for grouding conceptual content. Journal of Physiology, 102, 59-70, 2008; Hagoort, P. On Broca, brain, and binding: A new framework. Trends in Cognitive Sciences, 9, 416-423, 2005]. Previous research has supported the view that the coupling between language and action is flexible, and reading an action-related word form is not sufficient for cortical motor activation [Van Dam, W. O., van Dijk, M., Bekkering, H., & Rueschemeyer, S.-A. Flexibility in embodied lexical-semantic representations. Human Brain Mapping, doi: 10.1002/hbm.21365, 2011]. The current study goes one step further by addressing the necessity of action-related word forms for motor activation during language comprehension. Subjects listened to indirect requests (IRs) for action during an fMRI session. IRs for action are speech acts in which access to an action concept is required, although it is not explicitly encoded in the language. For example, the utterance "It is hot here!" in a room with a window is likely to be interpreted as a request to open the window. However, the same utterance in a desert will be interpreted as a statement. The results indicate (1) that comprehension of IR sentences activates cortical motor areas reliably more than comprehension of sentences devoid of any implicit motor information. This is true despite the fact that IR sentences contain no lexical reference to action. (2) Comprehension of IR sentences also reliably activates substantial portions of the theory of mind network, known to be involved in making inferences about mental states of others. The implications of these findings for embodied theories of language are discussed.

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.

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.

Research in psycholinguistics and in the cognitive neuroscience of language has suggested that semantic and syntactic processing are associated with different neurophysiologic correlates, such as the N400 and the P600 in the ERPs. However, only a handful of studies have investigated the neural basis of the syntax-semantics interface, and even fewer experiments have dealt with the cases in which semantic composition can proceed independently of the syntax. Here we looked into one such case-complement coercion--using ERPs. We compared sentences such as, "The journalist wrote the article" with "The journalist began the article." The second sentence seems to involve a silent semantic element, which is expressed in the first sentence by the head of the verb phrase (VP) "wrote the article." The second type of construction may therefore require the reader to infer or recover from memory a richer event sense of the VP "began the article," such as began writing the article, and to integrate that into a semantic representation of the sentence. This operation is referred to as "complement coercion." Consistently with earlier reading time, eye tracking, and MEG studies, we found traces of such additional computations in the ERPs: Coercion gives rise to a long-lasting negative shift, which differs at least in duration from a standard N400 effect. Issues regarding the nature of the computation involved are discussed in the light of a neurocognitive model of language processing and a formal semantic analysis of coercion.

There is growing evidence suggesting that synchronization changes in the oscillatory neuronal dynamics in the EEG or MEG reflect the transient coupling and uncoupling of functional networks related to different aspects of language comprehension. In this work, we examine how sentence-level syntactic unification operations are reflected in the oscillatory dynamics of the MEG. Participants read sentences that were either correct, contained a word category violation, or were constituted of random word sequences devoid of syntactic structure. A time-frequency analysis of MEG power changes revealed three types of effects. The first type of effect was related to the detection of a (word category) violation in a syntactically structured sentence, and was found in the alpha and gamma frequency bands. A second type of effect was maximally sensitive to the syntactic manipulations: A linear increase in beta power across the sentence was present for correct sentences, was disrupted upon the occurrence of a word category violation, and was absent in syntactically unstructured random word sequences. We therefore relate this effect to syntactic unification operations. Thirdly, we observed a linear increase in theta power across the sentence for all syntactically structured sentences. The effects are tentatively related to the building of a working memory trace of the linguistic input. In conclusion, the data seem to suggest that syntactic unification is reflected by neuronal synchronization in the lower-beta frequency band.

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.

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.

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

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

When do listeners take into account who the speaker is? We asked people to listen to utterances whose content sometimes did not match inferences based on the identity of the speaker (e.g., "If only I looked like Britney Spears" in a male voice, or "I have a large tattoo on my back" spoken with an upper-class accent). Event-related brain responses revealed that the speaker's identity is taken into account as early as 200-300 msec after the beginning of a spoken word, and is processed by the same early interpretation mechanism that constructs sentence meaning based on just the words. This finding is difficult to reconcile with standard "Gricean" models of sentence interpretation in which comprehenders initially compute a local, context-independent meaning for the sentence ("semantics") before working out what it really means given the wider communicative context and the particular speaker ("pragmatics"). Because the observed brain response hinges on voice-based and usually stereotype-dependent inferences about the speaker, it also shows that listeners rapidly classify speakers on the basis of their voices and bring the associated social stereotypes to bear on what is being said. According to our event-related potential results, language comprehension takes very rapid account of the social context, and the construction of meaning based on language alone cannot be separated from the social aspects of language use. The linguistic brain relates the message to the speaker immediately.

During language comprehension, listeners use the global semantic representation from previous sentence or discourse context to immediately integrate the meaning of each upcoming word into the unfolding message-level representation. Here we investigate whether communicative gestures that often spontaneously co-occur with speech are processed in a similar fashion and integrated to previous sentence context in the same way as lexical meaning. Event-related potentials were measured while subjects listened to spoken sentences with a critical verb (e.g., knock), which was accompanied by an iconic co-speech gesture (i.e., KNOCK). Verbal and/or gestural semantic content matched or mismatched the content of the preceding part of the sentence. Despite the difference in the modality and in the specificity of meaning conveyed by spoken words and gestures, the latency, amplitude, and topographical distribution of both word and gesture mismatches are found to be similar, indicating that the brain integrates both types of information simultaneously. This provides evidence for the claim that neural processing in language comprehension involves the simultaneous incorporation of information coming from a broader domain of cognition than only verbal semantics. The neural evidence for similar integration of information from speech and gesture emphasizes the tight interconnection between speech and co-speech gestures.

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.

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.

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

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

An event-related brain potential experiment was carried out to investigate the influence of semantic and syntactic context constraints on lexical selection and integration in spoken-word comprehension. Subjects were presented with constraining spoken sentences that contained a critical word that was either (a) congruent, (b) semantically and syntactically incongruent, but beginning with the same initial phonemes as the congruent critical word, or (c) semantically and syntactically incongruent, beginning with phonemes that differed from the congruent critical word. Relative to the congruent condition, an N200 effect reflecting difficulty in the lexical selection process was obtained in the semantically and syntactically incongruent condition where word onset differed from that of the congruent critical word. Both incongruent conditions elicited a large N400 followed by a left anterior negativity (LAN) time-locked to the moment of word category violation and a P600 effect. These results would best fit within a cascaded model of spoken-word processing, proclaiming an optimal use of contextual information during spoken-word identification by allowing for semantic and syntactic processing to take place in parallel after bottom-up activation of a set of candidates, and lexical integration to proceed with a limited number of candidates that still match the acoustic input.

This article presents electrophysiological data on on-line syntactic processing during auditory sentence comprehension in patients with Broca's aphasia. Event-related brain potentials (ERPs) were recorded from the scalp while subjects listened to sentences that were either syntactically correct or contained violations of subject-verb agreement. Three groups of subjects were tested: Broca patients (n = 10), nonaphasic patients with a right-hemisphere (RH) lesion (n = 5), and healthy aged-matched controls (n = 12). The healthy, control subjects showed a P600/SPS effect as response to the agreement violations. The nonaphasic patients with an RH lesion showed essentially the same pattern. The overall group of Broca patients did not show this sensitivity. However, the sensitivity was modulated by the severity of the syntactic comprehension impairment. The largest deviation from the standard P600/SPS effect was found in the patients with the relatively more severe syntactic comprehension impairment. In addition, ERPs to tones in a classical tone oddball paradigm were also recorded. Similar to the normal control subjects and RH patients, the group of Broca patients showed a P300 effect in the tone oddball condition. This indicates that aphasia in itself does not lead to a general reduction in all cognitive ERP effects. It was concluded that deviations from the standard P600/SPS effect in the Broca patients reflected difficulties with on-line maintaining of number information across clausal boundaries for establishing subject-verb agreement.

This study investigated the effects of combined semantic and syntactic violations in relation to the effects of single semantic and single syntactic violations on language-related event-related brain potential (ERP) effects (N400 and P600/SPS). Syntactic violations consisted of a mismatch in grammatical gender or number features of the definite article and the noun in sentence-internal or sentence-final noun phrases (NPs). Semantic violations consisted of semantically implausible adjective-noun combinations in the same NPs. Combined syntactic and semantic violations were a summation of these two respective violation types. ERPs were recorded while subjects read the sentences with the different types of violations and the correct control sentences. ERP effects were computed relative to ERPs elicited by the sentence-internal or sentence-final nouns. The size of the N400 effect to the semantic violation was increased by an additional syntactic violation (the syntactic boost). In contrast, the size of the P600/SPS to the syntactic violation was not affected by an additional semantic violation. This suggests that in the absence of syntactic ambiguity, the assignment of syntactic structure is independent of semantic context. However, semantic integration is influenced by syntactic processing. In the sentence-final position, additional global processing consequences were obtained as a result of earlier violations in the sentence. The resulting increase in the N400 amplitude to sentence-final words was independent of the nature of the violation. A speeded anomaly detection task revealed that it takes substantially longer to detect semantic than syntactic anomalies. These results are discussed in relation to the latency and processing characteristics of the N400 and P600/SPS effects. Overall, the results reveal an asymmetry in the interplay between syntax and semantics during on-line sentence comprehension.

An event-related brain potential experiment was carried out to investigate the time course of contextual influences on spoken-word recognition. Subjects were presented with spoken sentences that ended with a word that was either (a) congruent, (b) semantically anomalous, but beginning with the same initial phonemes as the congruent completion, or (c) semantically anomalous beginning with phonemes that differed from the congruent completion. In addition to finding an N400 effect in the two semantically anomalous conditions, we obtained an early negative effect in the semantically anomalous condition where word onset differed from that of the congruent completions. It was concluded that the N200 effect is related to the lexical selection process, where word-form information resulting from an initial phonological analysis and content information derived from the context interact.

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

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

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

In this study the N400 component of the event-related potential was used to investigate spoken sentence understanding in Broca's and Wernicke's aphasics. The aim of the study was to determine whether spoken sentence comprehension problems in these patients might result from a deficit in the on-line integration of lexical information. Subjects listened to sentences spoken at a normal rate. In half of these sentences, the meaning of the final word of the sentence matched the semantic specifications of the preceding sentence context. In the other half of the sentences, the sentence-final word was anomalous with respect to the preceding sentence context. The N400 was measured to the sentence-final words in both conditions. The results for the aphasic patients (n = 14) were analyzed according to the severity of their comprehension deficit and compared to a group of 12 neurologically unimpaired age-matched controls, as well as a group of 6 nonaphasic patients with a lesion in the right hemisphere. The nonaphasic brain damaged patients and the aphasic patients with a light comprehension deficit (high comprehenders, n = 7) showed an N400 effect that was comparable to that of the neurologically unimpaired subjects. In the aphasic patients with a moderate to severe comprehension deficit (low comprehenders, n = 7), a reduction and delay of the N400 effect was obtained. In addition, the P300 component was measured in a classical oddball paradigm, in which subjects were asked to count infrequent low tones in a random series of high and low tones. No correlation was found between the occurrence of N400 and P300 effects, indicating that changes in the N400 results were related to the patients' language deficit. Overall, the pattern of results was compatible with the idea that aphasic patients with moderate to severe comprehension problems are impaired in the integration of lexical information into a higher order representation of the preceding sentence context.