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.

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.

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.

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.

Context-dependent semantic processing in the human brain: evidence from idiom comprehension.

Language comprehension involves activating word meanings and integrating them with the sentence context. This study examined whether these routines are carried out even when they are theoretically unnecessary, namely, in the case of opaque idiomatic expressions, for which the literal word meanings are unrelated to the overall meaning of the expression. Predictable words in sentences were replaced by a semantically related or unrelated word. In literal sentences, this yielded previously established behavioral and electrophysiological signatures of semantic processing: semantic facilitation in lexical decision, a reduced N400 for semantically related relative to unrelated words, and a power increase in the gamma frequency band that was disrupted by semantic violations. However, the same manipulations in idioms yielded none of these effects. Instead, semantic violations elicited a late positivity in idioms. Moreover, gamma band power was lower in correct idioms than in correct literal sentences. It is argued that the brain's semantic expectancy and literal word meaning integration operations can, to some extent, be "switched off" when the context renders them unnecessary. Furthermore, the results lend support to models of idiom comprehension that involve unitary idiom representations.

ERP evidence on the interaction between information structure and emotional salience of words.

Both emotional words and words focused by information structure can capture attention. This study examined the interplay between emotional salience and information structure in modulating attentional resources in the service of integrating emotional words into sentence context. Event-related potentials (ERPs) to affectively negative, neutral, and positive words, which were either focused or nonfocused in question-answer pairs, were evaluated during sentence comprehension. The results revealed an early negative effect (90-200 ms), a P2 effect, as well as an effect in the N400 time window, for both emotional salience and information structure. Moreover, an interaction between emotional salience and information structure occurred within the N400 time window over right posterior electrodes, showing that information structure influences the semantic integration only for neutral words, but not for emotional words. This might reflect the fact that the linguistic salience of emotional words can override the effect of information structure on the integration of words into context. The interaction provides evidence for attention-emotion interactions at a later stage of processing. In addition, the absence of interaction in the early time window suggests that the processing of emotional information is highly automatic and independent of context. The results suggest independent attention capture systems of emotional salience and information structure at the early stage but an interaction between them at a later stage, during the semantic integration of words.

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.

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.

The anterior left inferior frontal gyrus contributes to semantic unification.

Semantic unification, the process by which small blocks of semantic information are combined into a coherent utterance, has been studied with various types of tasks. However, whether the brain activations reported in these studies are attributed to semantic unification per se or to other task-induced concomitant processes still remains unclear. The neural basis for semantic unification in sentence comprehension was examined using event-related potentials (ERP) and functional Magnetic Resonance Imaging (fMRI). The semantic unification load was manipulated by varying the goodness of fit between a critical word and its preceding context (in high cloze, low cloze and violation sentences). The sentences were presented in a serial visual presentation mode. The participants were asked to perform one of three tasks: semantic congruency judgment (SEM), silent reading for comprehension (READ), or font size judgment (FONT), in separate sessions. The ERP results showed a similar N400 amplitude modulation by the semantic unification load across all of the three tasks. The brain activations associated with the semantic unification load were found in the anterior left inferior frontal gyrus (aLIFG) in the FONT task and in a widespread set of regions in the other two tasks. These results suggest that the aLIFG activation reflects a semantic unification, which is different from other brain activations that may reflect task-specific strategic processing.

Beta oscillations relate to the N400m during language comprehension.

The relationship between the evoked responses (ERPs/ERFs) and the event-related changes in EEG/MEG power that can be observed during sentence-level language comprehension is as yet unclear. This study addresses a possible relationship between MEG power changes and the N400m component of the event-related field. Whole-head MEG was recorded while subjects listened to spoken sentences with incongruent (IC) or congruent (C) sentence endings. A clear N400m was observed over the left hemisphere, and was larger for the IC sentences than for the C sentences. A time-frequency analysis of power revealed a decrease in alpha and beta power over the left hemisphere in roughly the same time range as the N400m for the IC relative to the C condition. A linear regression analysis revealed a positive linear relationship between N400m and beta power for the IC condition, not for the C condition. No such linear relation was found between N400m and alpha power for either condition. The sources of the beta decrease were estimated in the LIFG, a region known to be involved in semantic unification operations. One source of the N400m was estimated in the left superior temporal region, which has been related to lexical retrieval. We interpret our data within a framework in which beta oscillations are inversely related to the engagement of task-relevant brain networks. The source reconstructions of the beta power suppression and the N400m effect support the notion of a dynamic communication between the LIFG and the left superior temporal region during language comprehension.

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.

Syntactic unification operations are reflected in oscillatory dynamics during on-line sentence comprehension.

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.

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.

Trial-by-trial coupling between EEG and BOLD identifies networks related to alpha and theta EEG power increases during working memory maintenance.

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

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.

Frontal theta EEG activity correlates negatively with the default mode network in resting state.

We used simultaneously recorded EEG and fMRI to investigate in which areas the BOLD signal correlates with frontal theta power changes, while subjects were quietly lying resting in the scanner with their eyes open. To obtain a reliable estimate of frontal theta power we applied ICA on band-pass filtered (2-9 Hz) EEG data. For each subject we selected the component that best matched the mid-frontal scalp topography associated with the frontal theta rhythm. We applied a time-frequency analysis on this component and used the time course of the frequency bin with the highest overall power to form a regressor that modeled spontaneous fluctuations in frontal theta power. No significant positive BOLD correlations with this regressor were observed. Extensive negative correlations were observed in the areas that together form the default mode network. We conclude that frontal theta activity can be seen as an EEG index of default mode network activity.

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.

Oscillatory neuronal dynamics during language comprehension.

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

EEG theta and gamma responses to semantic violations in online sentence processing.

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