Is beta in agreement with the relatives? Using relative clause sentences to investigate MEG beta power dynamics during sentence comprehension.

There remains some debate about whether beta power effects observed during sentence comprehension reflect ongoing syntactic unification operations (beta-syntax hypothesis), or instead reflect maintenance or updating of the sentence-level representation (beta-maintenance hypothesis). In this study, we used magnetoencephalography to investigate beta power neural dynamics while participants read relative clause sentences that were initially ambiguous between a subject- or an object-relative reading. An additional condition included a grammatical violation at the disambiguation point in the relative clause sentences. The beta-maintenance hypothesis predicts a decrease in beta power at the disambiguation point for unexpected (and less preferred) object-relative clause sentences and grammatical violations, as both signal a need to update the sentence-level representation. While the beta-syntax hypothesis also predicts a beta power decrease for grammatical violations due to a disruption of syntactic unification operations, it instead predicts an increase in beta power for the object-relative clause condition because syntactic unification at the point of disambiguation becomes more demanding. We observed decreased beta power for both the agreement violation and object-relative clause conditions in typical left hemisphere language regions, which provides compelling support for the beta-maintenance hypothesis. Mid-frontal theta power effects were also present for grammatical violations and object-relative clause sentences, suggesting that violations and unexpected sentence interpretations are registered as conflicts by the brain's domain-general error detection system.

Neural responses to facial attractiveness: Event-related potentials differentiate between salience and valence effects.

We examined the neural correlates of facial attractiveness by presenting pictures of male or female faces (neutral expression) with low/intermediate/high attractiveness to 48 male or female participants while recording their electroencephalogram (EEG). Subjective attractiveness ratings were used to determine the 10% highest, 10% middlemost, and 10% lowest rated faces for each individual participant to allow for high contrast comparisons. These were then split into preferred and dispreferred gender categories. ERP components P1, N1, P2, N2, early posterior negativity (EPN), P300 and late positive potential (LPP) (up until 3000 ms post-stimulus), and the face specific N170 were analysed. A salience effect (attractive/unattractive > intermediate) in an early LPP interval (450-850 ms) and a long-lasting valence related effect (attractive > unattractive) in a late LPP interval (1000-3000 ms) were elicited by the preferred gender faces but not by the dispreferred gender faces. Multi-variate pattern analysis (MVPA)-classifications on whole-brain single-trial EEG patterns further confirmed these salience and valence effects. It is concluded that, facial attractiveness elicits neural responses that are indicative of valenced experiences, but only if these faces are considered relevant. These experiences take time to develop and last well beyond the interval that is commonly explored.

Decoding the neural responses to experiencing disgust and sadness.

Being able to classify experienced emotions by identifying distinct neural responses has tremendous value in both fundamental research (e.g. positive psychology, emotion regulation theory) and in applied settings (clinical, healthcare, commercial). We aimed to decode the neural representation of the experience of two discrete emotions: sadness and disgust, devoid of differences in valence and arousal. In a passive viewing paradigm, we showed emotion evoking images from the International Affective Picture System to participants while recording their EEG. We then selected a subset of those images that were distinct in evoking either sadness or disgust (20 for each), yet were indistinguishable on normative valence and arousal. Event-related potential analysis of 69 participants showed differential responses in the N1 and EPN components and a support-vector machine classifier was able to accurately classify (58%) whole-brain EEG patterns of sadness and disgust experiences. These results support and expand on earlier findings that discrete emotions do have differential neural responses that are not caused by differences in valence or arousal.

On the Neuronal Dynamics of Aesthetic Experience: Evidence from Electroencephalographic Oscillatory Dynamics.

Aesthetic experiences have an influence on many aspects of life. Interest in the neural basis of aesthetic experiences has grown rapidly in the past decade, and fMRI studies have identified several brain systems supporting aesthetic experiences. Work on the rapid neuronal dynamics of aesthetic experience, however, is relatively scarce. This study adds to this field by investigating the experience of being aesthetically moved by means of ERP and time-frequency analysis. Participants' EEG was recorded while they viewed a diverse set of artworks and evaluated the extent to which these artworks moved them. Results show that being aesthetically moved is associated with a sustained increase in gamma activity over centroparietal regions. In addition, alpha power over right frontocentral regions was reduced in high- and low-moving images, compared to artworks given intermediate ratings. We interpret the gamma effect as an indication for sustained savoring processes for aesthetically moving artworks compared to aesthetically less-moving artworks. The alpha effect is interpreted as an indication of increased attention for aesthetically salient images. In contrast to previous works, we observed no significant effects in any of the established ERP components, but we did observe effects at latencies longer than 1 sec. We conclude that EEG time-frequency analysis provides useful information on the neuronal dynamics of aesthetic experience.

Are alpha oscillations instrumental in multisensory synchrony perception?

Different inputs from a multisensory object or event are often integrated into a coherent and unitary percept, despite differences in sensory formats, neural pathways, and processing times of the involved modalities. Presumably, multisensory integration occurs if the cross-modal inputs are presented within a certain window of temporal integration where inputs are perceived as being simultaneous. Here, we examine the role of ongoing neuronal alpha (i.e. 10-Hz) oscillations in multimodal synchrony perception. While EEG was measured, participants performed a simultaneity judgement task with visual stimuli preceding auditory ones. At stimulus onset asynchronies (SOA's) of 160-200 ms, simultaneity judgements were around 50%. For trials with these SOA's, occipital alpha power was smaller preceding correct judgements, and the individual alpha frequency was correlated with the size of the temporal window of integration. In addition, simultaneity judgements were modulated as a function of oscillatory phase at 12.5 Hz, but the latter effect was only marginally significant. These results support the notion that oscillatory neuronal activity in the alpha frequency range, which has been taken to shape perceptual cycles, is instrumental in multisensory perception.

From Experience to Memory: On the Robustness of the Peak-and-End-Rule for Complex, Heterogeneous Experiences.

Memory forms the input for future behavior. Therefore, how individuals remember a certain experience may be just as important as the experience itself. The peak-and-end-rule (PE-rule) postulates that remembered experiences are best predicted by the peak emotional valence and the emotional valence at the end of an experience in the here and now. The PE-rule, however, has mostly been assessed in experimental paradigms that induce relatively simple, one-dimensional experiences (e.g., experienced pain in a clinical setting). This hampers generalizations of the PE-rule to the experiences in everyday life. This paper evaluates the generalizability of the PE-rule to more complex and heterogeneous experiences by examining the PE-rule in a virtual reality (VR) experience, as VR combines improved ecological validity with rigorous experimental control. Findings indicate that for more complex and heterogeneous experiences, peak and end emotional valence are inferior to other measures (such as averaged valence and arousal ratings over the entire experiential episode) in predicting remembered experience. These findings suggest that the PE-rule cannot be generalized to ecologically more valid experiential episodes.

Assessing the utility of frequency tagging for tracking memory-based reactivation of word representations.

Reinstatement of memory-related neural activity measured with high temporal precision potentially provides a useful index for real-time monitoring of the timing of activation of memory content during cognitive processing. The utility of such an index extends to any situation where one is interested in the (relative) timing of activation of different sources of information in memory, a paradigm case of which is tracking lexical activation during language processing. Essential for this approach is that memory reinstatement effects are robust, so that their absence (in the average) definitively indicates that no lexical activation is present. We used electroencephalography to test the robustness of a reported subsequent memory finding involving reinstatement of frequency-specific entrained oscillatory brain activity during subsequent recognition. Participants learned lists of words presented on a background flickering at either 6 or 15 Hz to entrain a steady-state brain response. Target words subsequently presented on a non-flickering background that were correctly identified as previously seen exhibited reinstatement effects at both entrainment frequencies. Reliability of these statistical inferences was however critically dependent on the approach used for multiple comparisons correction. We conclude that effects are not robust enough to be used as a reliable index of lexical activation during language processing.

Control adjustments in speaking: Electrophysiology of the Gratton effect in picture naming.

Accumulating evidence suggests that spoken word production requires different amounts of top-down control depending on the prevailing circumstances. For example, during Stroop-like tasks, the interference in response time (RT) is typically larger following congruent trials than following incongruent trials. This effect is called the Gratton effect, and has been taken to reflect top-down control adjustments based on the previous trial type. Such control adjustments have been studied extensively in Stroop and Eriksen flanker tasks (mostly using manual responses), but not in the picture-word interference (PWI) task, which is a workhorse of language production research. In one of the few studies of the Gratton effect in PWI, Van Maanen and Van Rijn (2010) examined the effect in picture naming RTs during dual-task performance. Based on PWI effect differences between dual-task conditions, they argued that the functional locus of the PWI effect differs between post-congruent trials (i.e., locus in perceptual and conceptual encoding) and post-incongruent trials (i.e., locus in word planning). However, the dual-task procedure may have contaminated the results. We therefore performed an electroencephalography (EEG) study on the Gratton effect in a regular PWI task. We observed a PWI effect in the RTs, in the N400 component of the event-related brain potentials, and in the midfrontal theta power, regardless of the previous trial type. Moreover, the RTs, N400, and theta power reflected the Gratton effect. These results provide evidence that the PWI effect arises at the word planning stage following both congruent and incongruent trials, while the amount of top-down control changes depending on the previous trial type.

Using Brain Potentials to Functionally Localise Stroop-Like Effects in Colour and Picture Naming: Perceptual Encoding versus Word Planning.

The colour-word Stroop task and the picture-word interference task (PWI) have been used extensively to study the functional processes underlying spoken word production. One of the consistent behavioural effects in both tasks is the Stroop-like effect: The reaction time (RT) is longer on incongruent trials than on congruent trials. The effect in the Stroop task is usually linked to word planning, whereas the effect in the PWI task is associated with either word planning or perceptual encoding. To adjudicate between the word planning and perceptual encoding accounts of the effect in PWI, we conducted an EEG experiment consisting of three tasks: a standard colour-word Stroop task (three colours), a standard PWI task (39 pictures), and a Stroop-like version of the PWI task (three pictures). Participants overtly named the colours and pictures while their EEG was recorded. A Stroop-like effect in RTs was observed in all three tasks. ERPs at centro-parietal sensors started to deflect negatively for incongruent relative to congruent stimuli around 350 ms after stimulus onset for the Stroop, Stroop-like PWI, and the Standard PWI tasks: an N400 effect. No early differences were found in the PWI tasks. The onset of the Stroop-like effect at about 350 ms in all three tasks links the effect to word planning rather than perceptual encoding, which has been estimated in the literature to be finished around 200-250 ms after stimulus onset. We conclude that the Stroop-like effect arises during word planning in both Stroop and PWI.

A Predictive Coding Perspective on Beta Oscillations during Sentence-Level Language Comprehension.

Oscillatory neural dynamics have been steadily receiving more attention as a robust and temporally precise signature of network activity related to language processing. We have recently proposed that oscillatory dynamics in the beta and gamma frequency ranges measured during sentence-level comprehension might be best explained from a predictive coding perspective. Under our proposal we related beta oscillations to both the maintenance/change of the neural network configuration responsible for the construction and representation of sentence-level meaning, and to top-down predictions about upcoming linguistic input based on that sentence-level meaning. Here we zoom in on these particular aspects of our proposal, and discuss both old and new supporting evidence. Finally, we present some preliminary magnetoencephalography data from an experiment comparing Dutch subject- and object-relative clauses that was specifically designed to test our predictive coding framework. Initial results support the first of the two suggested roles for beta oscillations in sentence-level language comprehension.

Frequency-based Segregation of Syntactic and Semantic Unification during Online Sentence Level Language Comprehension.

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.

A predictive coding framework for rapid neural dynamics during sentence-level language comprehension.

There is a growing literature investigating the relationship between oscillatory neural dynamics measured using electroencephalography (EEG) and/or magnetoencephalography (MEG), and sentence-level language comprehension. Recent proposals have suggested a strong link between predictive coding accounts of the hierarchical flow of information in the brain, and oscillatory neural dynamics in the beta and gamma frequency ranges. We propose that findings relating beta and gamma oscillations to sentence-level language comprehension might be unified under such a predictive coding account. Our suggestion is that oscillatory activity in the beta frequency range may reflect both the active maintenance of the current network configuration responsible for representing the sentence-level meaning under construction, and the top-down propagation of predictions to hierarchically lower processing levels based on that representation. In addition, we suggest that oscillatory activity in the low and middle gamma range reflect the matching of top-down predictions with bottom-up linguistic input, while evoked high gamma might reflect the propagation of bottom-up prediction errors to higher levels of the processing hierarchy. We also discuss some of the implications of this predictive coding framework, and we outline ideas for how these might be tested experimentally.

The influence of emotional salience on the integration of person names into context.

Previous event-related potentials (ERP) studies on the processing of emotional information in sentence/discourse context have yielded inconsistent findings. An important reason for the discrepancies is the different lexico-semantic properties of the emotional words. The present study controlled for the lexico-semantic meaning of emotional information by endowing the same person names with either positive or negative valence. ERPs were computed for positively and negatively valenced person names that were either congruent or incongruent to previous emotional contexts. We found that positive names elicited an N400 effect while negative names elicited a P600 effect in response to the incongruence. These results suggest that the integration of positive and negative information into emotional context exhibits different time courses, with a relatively delayed integration for negative information. Our study demonstrates that using person names constitutes a new and improved tool for investigating the integration of emotional information into context.

Fast oscillatory dynamics during language comprehension: Unification versus maintenance and prediction?

The role of neuronal oscillations during language comprehension is not yet well understood. In this paper we review and reinterpret the functional roles of beta- and gamma-band oscillatory activity during language comprehension at the sentence and discourse level. We discuss the evidence in favor of a role for beta and gamma in unification (the unification hypothesis), and in light of mounting evidence that cannot be accounted for under this hypothesis, we explore an alternative proposal linking beta and gamma oscillations to maintenance and prediction (respectively) during language comprehension. Our maintenance/prediction hypothesis is able to account for most of the findings that are currently available relating beta and gamma oscillations to language comprehension, and is in good agreement with other proposals about the roles of beta and gamma in domain-general cognitive processing. In conclusion we discuss proposals for further testing and comparing the prediction and unification hypotheses.

Brain dynamics in the comprehension of action-related language. A time-frequency analysis of mu rhythms.

EEG mu rhythms (8-13 Hz) recorded at fronto-central electrodes are generally considered as markers of motor cortical activity in humans, because they are modulated when participants perform an action, when they observe another's action or even when they imagine performing an action. In this study, we analyzed the time-frequency (TF) modulation of mu rhythms while participants read action language ("You will cut the strawberry cake"), abstract language ("You will doubt the patient's argument"), and perceptive language ("You will notice the bright day"). The results indicated that mu suppression at fronto-central sites is associated with action language rather than with abstract or perceptive language. Also, the largest difference between conditions occurred quite late in the sentence, while reading the first noun, (contrast Action vs. Abstract), or the second noun following the action verb (contrast Action vs. Perceptive). This suggests that motor activation is associated with the integration of words across the sentence beyond the lexical processing of the action verb. Source reconstruction localized mu suppression associated with action sentences in premotor cortex (BA 6). The present study suggests (1) that the understanding of action language activates motor networks in the human brain, and (2) that this activation occurs online based on semantic integration across multiple words in the sentence.

ERP responses to person names as a measure of trait inference in person perception.

Using event-related potentials (ERPs), this study examines how trait information inferred from behaviors is associated with person names. In linguistic discourses, person names were associated with descriptions of either positive or negative behaviors. In a subsequent explicit evaluation task, the previously described person names were presented in isolation, and the participants were asked to judge the emotional valence of these names. We found that the names associated with positive descriptions elicited a larger positivity in the ERP than the names associated with negative descriptions. The results indicate that the emotional valence of person names attached to person perception can be dynamically influenced by short descriptions of the target person, probably due to trait inference based on the provided behavioral descriptions.

Oscillatory brain dynamics associated with the automatic processing of emotion in words.

This study examines the automaticity of processing the emotional aspects of words, and characterizes the oscillatory brain dynamics that accompany this automatic processing. Participants read emotionally negative, neutral and positive nouns while performing a color detection task in which only perceptual-level analysis was required. Event-related potentials and time frequency representations were computed from the concurrently measured EEG. Negative words elicited a larger P2 and a larger late positivity than positive and neutral words, indicating deeper semantic/evaluative processing of negative words. In addition, sustained alpha power suppressions were found for the emotional compared to neutral words, in the time range from 500 to 1000ms post-stimulus. These results suggest that sustained attention was allocated to the emotional words, whereas the attention allocated to the neutral words was released after an initial analysis. This seems to hold even when the emotional content of the words is task-irrelevant.

Early anticipation lies behind the speed of response in conversation.

RTs in conversation, with average gaps of 200 msec and often less, beat standard RTs, despite the complexity of response and the lag in speech production (600 msec or more). This can only be achieved by anticipation of timing and content of turns in conversation, about which little is known. Using EEG and an experimental task with conversational stimuli, we show that estimation of turn durations are based on anticipating the way the turn would be completed. We found a neuronal correlate of turn-end anticipation localized in ACC and inferior parietal lobule, namely a beta-frequency desynchronization as early as 1250 msec, before the end of the turn. We suggest that anticipation of the other's utterance leads to accurately timed transitions in everyday conversations.

Sit down and read on: working memory and long-term memory in particle-verb processing.

Particle verbs (e.g., look up) are lexical items for which particle and verb share a single lexical entry. Using event-related brain potentials, we examined working memory and long-term memory involvement in particle-verb processing. Dutch participants read sentences with head verbs that allow zero, two, or more than five particles to occur downstream. Additionally, sentences were presented for which the encountered particle was semantically plausible, semantically implausible, or forming a non-existing particle verb. An anterior negativity was observed at the verbs that potentially allow for a particle downstream relative to verbs that do not, possibly indexing storage of the verb until the dependency with its particle can be closed. Moreover, a graded N400 was found at the particle (smallest amplitude for plausible particles and largest for particles forming non-existing particle verbs), suggesting that lexical access to a shared lexical entry occurred at two separate time points.

Recognizing the emotional valence of names: an ERP study.

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.