N400 Time Window Research Articles

Cross-modal symbolic processing can elicit either an N2 or a protracted N2/N400 response.

A cross-modal symbolic paradigm was used to elicit EEG activity related to semantic incongruence. Twenty-five undergraduate students viewed pairings of visual lexical cues (e.g., DOG) with congruent (50% of trials) or incongruent (50%) auditory nonlexical stimuli (animal vocalizations; e.g., sound of a dog woofing or a cat meowing). In one condition, many different pairs of congruent/incongruent stimuli were shown, whereas in a second condition only two pairs of stimuli were repeatedly shown. A typical N400-like pattern of incongruence-related activity (including activity in the N2 time window) was evident in the condition using many stimuli, whereas the incongruence-related activity in the two-stimuli condition was confined to differential N2-like activity. A supplementary analysis excluded stimulus characteristics as the source of this differential activity between conditions. We found that a single individual performing a fixed task can demonstrate either a protracted N400-like pattern of activity or a more temporally focused N2-like pattern of activity in response to the same stimulus, which suggests that the N2 may be a precursor to the protracted N400 response.

Categorization Method Affects the Typicality Effect: ERP Evidence from a Category-Inference Task.

The typicality effect during categorization describes a phenomenon whereby typical items are more easily judged as members of a category than atypical items. Prior studies of the typicality effect have often used an inclusion task, which asks participants to assess whether an item belongs to a category. However, the correct exclusion of non-members is also an important component of effective categorization, which has yet to be directly investigated. Thus, the present study investigated how categorization method (inclusion vs. exclusion) modulates the typicality effect via behavioral and electrophysiological measures. Thirty-two participants (16 in the inclusion and 16 in the exclusion group) were shown six consecutive words that all shared one feature. Then, a seventh word was presented. The inclusion group judged whether the seventh word also possessed the feature, whereas the exclusion group judged whether the word did not possess the feature. The seventh word could be typical, atypical, or a nonmember of the category. Behavioral and event-related potential (ERP) data were collected. Behavioral results showed that the two groups did not differ in accuracy. However, typical items elicited shorter response times than atypical items, and this effect was more pronounced in the inclusion than the exclusion group. With regard to ERPs, interactions between item type and group were shown for the P2, N2, and N400 periods. Within the inclusion group, a typicality effect (indicated by a main effect of item type) was present in the P2 and N400 time windows, while the exclusion group elicited a typicality effect only in the N2 time window. These results provide electrophysiological evidence that an inclusion judgment task is more sensitive to category typicality than is an exclusion task.

Imageability and semantic association in the representation and processing of event verbs.

This study examined the relative salience of imageability (the degree to which a word evokes mental imagery) versus semantic association (the density of semantic network in which a word is embedded) in the representation and processing of four types of event verbs: sensory, cognitive, speech, and motor verbs. ERP responses were recorded, while 34 university students performed on a lexical decision task. Analysis focused primarily on amplitude differences across verb conditions within the N400 time window where activities are considered representing meaning activation. Variation in N400 amplitude across four types of verbs was found significantly associated with the level of imageability, but not the level of semantic association. The findings suggest imageability as a more salient factor relative to semantic association in the processing of these verbs. The role of semantic association and the representation of speech verbs are also discussed.

The Effect of Distance on Moral Engagement: Event Related Potentials and Alpha Power are Sensitive to Perspective in a Virtual Shooting Task.

In a shooting video game we investigated whether increased distance reduces moral conflict. We measured and analyzed the event related potential (ERP), including the N2 component, which has previously been linked to cognitive conflict from competing decision tendencies. In a modified Go/No-go task designed to trigger moral conflict participants had to shoot suddenly appearing human like avatars in a virtual reality scene. The scene was seen either from an ego perspective with targets appearing directly in front of the participant or from a bird's view, where targets were seen from above and more distant. To control for low level visual features, we added a visually identical control condition, where the instruction to “shoot” was replaced by an instruction to “detect.” ERP waveforms showed differences between the two tasks as early as in the N1 time-range, with higher N1 amplitudes for the close perspective in the “shoot” task. Additionally, we found that pre-stimulus alpha power was significantly decreased in the ego, compared to the bird's view only for the “shoot” but not for the “detect” task. In the N2 time window, we observed main amplitude effects for response (No-go > Go) and distance (ego > bird perspective) but no interaction with task type (shoot vs. detect). We argue that the pre-stimulus and N1 effects can be explained by reduced attention and arousal in the distance condition when people are instructed to “shoot.” These results indicate a reduced moral engagement for increased distance. The lack of interaction in the N2 across tasks suggests that at that time point response execution dominates. We discuss potential implications for real life shooting situations, especially considering recent developments in drone shootings which are per definition of a distant view.

Impaired speech perception leads to poor semantic access: ERP evidence from individuals with pathologically or artificially impaired speech perception.

Impaired speech perception leads to poor semantic access: ERP evidence from individuals with pathologically or artificially impaired speech perception.

When Feelings Arise with Meanings: How Emotion and Meaning of a Native Language Affect Second Language Processing in Adult Learners.

To determine when and how L2 learners start to process L2 words affectively and semantically, we conducted a longitudinal study on their interaction in adult L2 learners. In four test sessions, spanning half a year of L2 learning, we monitored behavioral and ERP learning-related changes for one and the same set of words by means of a primed lexical-decision paradigm with L1 primes and L2 targets. Sensitivity rates, accuracy rates, RTs, and N400 amplitude to L2 words and pseudowords improved significantly across sessions. A semantic priming effect (e.g, prime “driver”facilitating response to target “street”) was found in accuracy rates and RTs when collapsing Sessions 1 to 4, while this effect modulated ERP amplitudes within the first 300 ms of L2 target processing. An overall affective priming effect (e.g., “sweet” facilitating”taste”) was also found in RTs and ERPs (posterior P1). Importantly, the ERPs showed an L2 valence effect across sessions (e.g., positive words were easier to process than neutral words), indicating that L2 learners were sensitive to L2 affective meaning. Semantic and affective priming interacted in the N400 time-window only in Session 4, implying that they affected meaning integration during L2 immersion together. The results suggest that L1 and L2 are initially processed semantically and affectively via relatively separate channels that are more and more linked contingent on L2 exposure.

Perceived arousal of facial expressions of emotion modulates the N170, regardless of emotional category: Time domain and time–frequency dynamics

Findings concerning the emotional modulation of the N170 component of the visual event-related potential are mixed. In the present report we tested the hypothesis that the emotional modulation of the N170 may be driven by the perceived emotional arousal of the stimuli, rather than by specific emotional categories.Fifty-four participants viewed facial expressions of anger, disgust, fear and happiness, plus low arousal neutral faces. All emotional categories were matched in arousal, while stimuli within each category varied parametrically in this dimension. The modulation of the electrocortical activity on the N170 time-window was analyzed in the time domain and via time–frequency decomposition. The effects of emotion and arousal were analyzed separately.In the time domain N170 amplitudes co-varied parametrically with perceived arousal, regardless of emotional category. This modulation was linearly associated with the power of the theta, alpha, and beta frequency bands. Moreover, fear was associated with a trend for increased N170 amplitudes, enhanced alpha power, and increased broad band inter-trial phase coherence.These results support the views that a) the activity in N170 time window is fundamentally modulated by perceived arousal, b) the modulation of the N170 may be the product of an increased evoked response, rather than the result of phase resetting processes, and c) facial expressions of fear retain some processing primacy, that may be related to their increased value as environmental cues.

Early perceptual anomaly of negative facial expression in depression: An event-related potential study

Depressed patients have a demonstrated cognitive bias in emotional information processing. However, it is unknown how early perceptual processing is modulated by emotional stimuli in depression. To examine this question, we studied 22 depressed patients and 22 healthy controls performing a cued target-response task with emotional facial expression as the cue. The early perceptual processes were examined using event-related potential (ERP) components, i.e., P1 and N170. Results showed that depressed patients had larger P1 amplitudes than healthy controls, implying that early perceptual abnormality for face processing in depression may occur as early as the P1 stage. There was no significant interaction between emotion types and groups on P1 amplitudes, which suggested that cognitive biases in depression might not yet have arisen. Following the P1 stage, N170 amplitudes for sad faces were larger than for other emotion types in depressed patients, whereas N170 amplitudes for happy faces were larger than for other emotion types in healthy controls. These results implied that depressed patients might have a perceptual bias associated with sad emotions, which may be detectable from the N170 time window. In summary, this study provides new insights for understanding the negative cognitive bias in depression using the electroneurophysiological biomarker N170.

Finding indexes of spontaneous brain-to-brain communications when looking for a cause of the similarity of qualia assumed across individuals

Qualia, the individual instances of subjective conscious experience, are private events. However, in everyday life, we assume qualia of others and their perceptual worlds, to be similar to ours. One way this similarity is possible is if qualia of others somehow contribute to the production of qualia by our own brain and vice versa. To test this hypothesis, we focused on the mean voltages of event-related potentials (ERPs) in the time-window of the P600 component, whose amplitude correlates positively with conscious awareness. These ERPs were elicited by stimuli of the international affective picture system in 16 pairs of friends, siblings or couples going side by side through hyperscanning without having to interact. Each of the 32 members of these 16 pairs faced one half of the screen and could not see what the other member was presented with on the other half. One stimulus occurred on each half simultaneously. The sameness of these stimulus pairs was manipulated as well as the participants’ belief in that sameness by telling subjects’ pairs that they were going to be presented with the same stimuli in two blocks and with different ones in the two others. In the P600 time window, belief, and thus social cognition, was found to have an effect on ERPs only at left anterior electrode sites. In contrast, ERPs were more positive at all electrode subsets for stimulus pairs that were inconsistent with the belief than for those that were consistent. In the N400 time window, at frontal electrode sites, ERPs were again more positive for inconsistent than for consistent stimuli. As participants had no way to see the stimulus their partner was presented with, and thus no way to detect inconsistence, we proposed that these data could support the existence of spontaneous brain-to-brain communications. Such communications might provide a research avenue when trying to explain the similarity of qualia across individuals, which is assumed in virtually all instants of every day life.

Looking for effects of qualia on event-related brain potentials of close others in search for a cause of the similarity of qualia assumed across individuals

Qualia, the individual instances of subjective conscious experience, are private events. However, in everyday life, we assume qualia of others and their perceptual worlds, to be similar to ours. One way this similarity is possible is if qualia of others somehow contribute to the production of qualia by our own brain and vice versa. To test this hypothesis, we focused on the mean voltages of event-related potentials (ERPs) in the time-window of the P600 component, whose amplitude correlates positively with conscious awareness. These ERPs were elicited by images of the international affective picture system in 16 pairs of friends, siblings or couples going side by side through hyperscanning without having to interact. Each of the 32 members of these 16 pairs faced one half of the screen and could not see what the other member was presented with on the other half. One stimulus occurred on each half simultaneously. The sameness of these stimulus pairs was manipulated as well as the participants’ belief in that sameness by telling subjects’ pairs that they were going to be presented with the same stimuli in two blocks and with different ones in the two others. ERPs were more positive at all electrode subsets for stimulus pairs that were inconsistent with the belief than for those that were consistent. In the N400 time window, at frontal electrode sites, ERPs were again more positive for inconsistent than for consistent stimuli. As participants had no way to see the stimulus their partner was presented with and thus no way to detect inconsistence, these data might reveal an impact of the qualia of a person on the brain activity of another. Such impact could provide a research avenue when trying to explain the similarity of qualia across individuals.

Is Semantic Processing During Sentence Reading Autonomous or Controlled? Evidence from the N400 Component in a Dual Task Paradigm.

The present study contributes to the discussion on the automaticity of semantic processing. Whereas most previous research investigated semantic processing at word level, the present study addressed semantic processing during sentence reading. A dual task paradigm was combined with the recording of event-related brain potentials. Previous research at word level processing reported different patterns of interference with the N400 by additional tasks: attenuation of amplitude or delay of latency. In the present study, we presented Spanish sentences that were semantically correct or contained a semantic violation in a critical word. At different intervals preceding the critical word a tone was presented that required a high-priority choice response. At short intervals/high temporal overlap between the tasks mean amplitude of the N400 was reduced relative to long intervals/low temporal overlap, but there were no shifts of peak latency. We propose that processing at sentence level exerts a protective effect against the additional task. This is in accord with the attentional sensitization model (Kiefer & Martens, 2010), which suggests that semantic processing is an automatic process that can be enhanced by the currently activated task set. The present experimental sentences also induced a P600, which is taken as an index of integrative processing. Additional task effects are comparable to those in the N400 time window and are briefly discussed.

Neuroelectric Correlates of Pragmatic Emotional Incongruence Processing: Empathy Matters.

The emotions people feel can be simulated internally based on emotional situational contexts. In the present study, we assessed the behavioral and neuroelectric effects of seeing an unexpected emotional facial expression. We investigated the correct answer rate, response times and Event-Related Potential (ERP) effects during an incongruence paradigm between emotional faces and sentential contexts allowing emotional inferences. Most of the 36 healthy participants were recruited from a larger population (1 463 subjects), based on their scores on the Empathy Questionnaire (EQ). Regression analyses were conducted on these ratings using EQ factors as predictors (cognitive empathy, emotional reactivity and social skills). Recognition of pragmatic emotional incongruence was less accurate (P < .05) and slower (P < .05) than recognition of congruence. The incongruence effect on response times was inversely predicted by social skills. A significant N400 incongruence effect was found at the centro-parietal (P < .001) and centro-posterior midline (P < .01) electrodes. Cognitive empathy predicted the incongruence effect in the left occipital region, in the N400 time window. Finally, incongruence effects were also found on the LPP wave, in frontal midline and dorso-frontal regions, (P < .05), with no modulation by empathy. Processing pragmatic emotional incongruence is more cognitively demanding than congruence (as reflected by both behavioral and ERP data). This processing shows modulation by personality factors at the behavioral (through self-reported social skills) and neuroelectric levels (through self-reported cognitive empathy).

N400-like effect of stroke-based neighborhood size in Chinese characters

Word-to-letter neighborhood feedback has been proposed to account for the lexical facilitation effect of orthographic neighborhood size (N). In the present study, a new N of Chinese characters was manipulated. Neighbors were created by randomly replacing about one third of the total strokes of characters while keeping the structure and the orientation of its remaining strokes constant. Participants were asked to match probe characters with the preceding target characters while event-related brain potentials elicited by target characters were recorded. In the N400 time window, characters with many neighbors produced more negative amplitudes than characters with few neighbors. The N400-like effect supports the lateral inhibition component of the interactive activation model where the non-target candidates are inhibited; lateral inhibition contributes to the N400-like effect of N more than does character-to-stroke feedback. Likewise, a large neighborhood size yielded higher N250 amplitudes, thus possibly indicating greater summed mismatches between the numerous neighbors and their targets.

Selective attention modulates early human evoked potentials during emotional face-voice processing.

Recent findings on multisensory integration suggest that selective attention influences cross-sensory interactions from an early processing stage. Yet, in the field of emotional face-voice integration, the hypothesis prevails that facial and vocal emotional information interacts preattentively. Using ERPs, we investigated the influence of selective attention on the perception of congruent versus incongruent combinations of neutral and angry facial and vocal expressions. Attention was manipulated via four tasks that directed participants to (i) the facial expression, (ii) the vocal expression, (iii) the emotional congruence between the face and the voice, and (iv) the synchrony between lip movement and speech onset. Our results revealed early interactions between facial and vocal emotional expressions, manifested as modulations of the auditory N1 and P2 amplitude by incongruent emotional face-voice combinations. Although audiovisual emotional interactions within the N1 time window were affected by the attentional manipulations, interactions within the P2 modulation showed no such attentional influence. Thus, we propose that the N1 and P2 are functionally dissociated in terms of emotional face-voice processing and discuss evidence in support of the notion that the N1 is associated with cross-sensory prediction, whereas the P2 relates to the derivation of an emotional percept. Essentially, our findings put the integration of facial and vocal emotional expressions into a new perspective-one that regards the integration process as a composite of multiple, possibly independent subprocesses, some of which are susceptible to attentional modulation, whereas others may be influenced by additional factors.

Re-examination of Chinese semantic processing and syntactic processing: evidence from conventional ERPs and reconstructed ERPs by residue iteration decomposition (RIDE).

A number of studies have explored the time course of Chinese semantic and syntactic processing. However, whether syntactic processing occurs earlier than semantics during Chinese sentence reading is still under debate. To further explore this issue, an event-related potentials (ERPs) experiment was conducted on 21 native Chinese speakers who read individually-presented Chinese simple sentences (NP1+VP+NP2) word-by-word for comprehension and made semantic plausibility judgments. The transitivity of the verbs was manipulated to form three types of stimuli: congruent sentences (CON), sentences with a semantically violated NP2 following a transitive verb (semantic violation, SEM), and sentences with a semantically violated NP2 following an intransitive verb (combined semantic and syntactic violation, SEM+SYN). The ERPs evoked from the target NP2 were analyzed by using the Residue Iteration Decomposition (RIDE) method to reconstruct the ERP waveform blurred by trial-to-trial variability, as well as by using the conventional ERP method based on stimulus-locked averaging. The conventional ERP analysis showed that, compared with the critical words in CON, those in SEM and SEM+SYN elicited an N400–P600 biphasic pattern. The N400 effects in both violation conditions were of similar size and distribution, but the P600 in SEM+SYN was bigger than that in SEM. Compared with the conventional ERP analysis, RIDE analysis revealed a larger N400 effect and an earlier P600 effect (in the time window of 500–800 ms instead of 570–810ms). Overall, the combination of conventional ERP analysis and the RIDE method for compensating for trial-to-trial variability confirmed the non-significant difference between SEM and SEM+SYN in the earlier N400 time window. Converging with previous findings on other Chinese structures, the current study provides further precise evidence that syntactic processing in Chinese does not occur earlier than semantic processing.

Loving yourself more than your neighbor: ERPs reveal online effects of a self-positivity bias.

A large body of social psychological research suggests that we think quite positively of ourselves, often unrealistically so. Research on this 'self-positivity bias' has relied mainly on self-report and behavioral measures, but these can suffer from a number of problems including confounds that arise from the desire to present oneself well. What has not been clearly assessed is whether the self-positivity bias influences the processing of incoming information as it unfolds in real time. In this study, we used event-related potentials to address this question. Participants read two-sentence social vignettes that were either self- or other-relevant. Pleasant words in self-relevant contexts evoked a smaller negativity between 300 and 500 ms (the N400 time window) than the same words in other-relevant contexts, suggesting that comprehenders were more likely to expect positive information when a scenario referred to themselves. This finding indicates that the self-positivity bias is available online, acting as a general schema that directly influences real-time comprehension.

An ERP investigation of orthographic priming with superset primes

Prime stimuli formed by inserting unrelated letters in a given target word (called “superset” primes) provide a means to modify the relative positions of the letters shared by prime and target. Here we examined the time-course of superset priming effects in an ERP study using the sandwich-priming paradigm. We compared the effects of superset primes formed by the insertion of unrelated letters (e.g., maurkdet–MARKET), or by the insertion of hyphens (e.g., ma-rk-et–MARKET), with identity priming (e.g., market–MARKET), all measured relative to unrelated control primes. Behavioral data revealed significantly greater priming in the hyphen-insert condition compared with the letter-insert condition. In the ERP signal, letter-insert priming emerged later than hyphen-insert priming and produced a reversed priming effect in the N400 time-window compared with the more typical N400 priming effects seen for both hyphen-insert priming and identity priming. The different pattern of priming effects seen for letter-insert primes and hyphen-insert primes suggests that compared with identity priming, letter superset priming reflects the joint influence of: (1) a disruption in letter position information, and (2) an inhibitory influence of mismatching letters.

Interaction Between Words and Symbolic Gestures as Revealed By N400.

What happens if you see a person pronouncing the word "go" after having gestured "stop"? Differently from iconic gestures, that must necessarily be accompanied by verbal language in order to be unambiguously understood, symbolic gestures are so conventionalized that they can be effortlessly understood in the absence of speech. Previous studies proposed that gesture and speech belong to a unique communication system. From an electrophysiological perspective the N400 modulation was considered the main variable indexing the interplay between two stimuli. However, while many studies tested this effect between iconic gestures and speech, little is known about the capability of an emblem to modulate the neural response to subsequently presented words. Using high-density EEG, the present study aimed at evaluating the presence of an N400 effect and its spatiotemporal dynamics, in terms of cortical activations, when emblems primed the observation of words. Participants were presented with symbolic gestures followed by a semantically congruent or incongruent verb. A N400 modulation was detected, showing larger negativity when gesture and words were incongruent. The source localization during N400 time window evidenced the activation of different portions of temporal cortex according to the gesture and word congruence. Our data provide further evidence of how the observation of an emblem influences verbal language perception, and of how this interplay is mainly instanced by different portions of the temporal cortex.

On the time-course of adjacent and non-adjacent transposed-letter priming

We compared effects of adjacent (e.g. atricle-ARTICLE) and non-adjacent (e.g. actirle-ARTICLE) transposed-letter (TL) primes in an event-related potential (ERP) study using the sandwich priming technique. TL priming was measured relative to the standard double-substitution condition. We found significantly stronger priming effects for adjacent transpositions than non-adjacent transpositions (with two intervening letters) in behavioural responses (lexical decision latencies), and the adjacent priming effects emerged earlier in the ERP signal, at around 200-ms post-target onset. Non-adjacent priming effects emerged about 50 ms later and were short-lived, being significant only in the 250- to 300-ms time-window. Adjacent transpositions on the other hand continued to produce priming in the N400 time-window (300- to 500-ms post-target onset). This qualitatively different pattern of priming effects for adjacent and non-adjacent transpositions is discussed in the light of different accounts of letter transposition effects and the utility of drawing a distinction between positional flexibility and positional noise.

Sustained posterior contralateral activity indicates re-entrant target processing in visual change detection: an EEG study.

The present study investigated the neural mechanisms that contribute to the detection of visual feature changes between stimulus displays by means of event-related lateralizations of the electroencephalogram (EEG). Participants were instructed to respond to a luminance change in either of two lateralized stimuli that could randomly occur alone or together with an irrelevant orientation change of the same or contralateral stimulus. Task performance based on response times and accuracy was decreased compared to the remaining stimulus conditions when relevant and irrelevant feature changes were presented contralateral to each other (contralateral distractor condition). The sensory response to the feature changes was reflected in a posterior contralateral positivity at around 100 ms after change presentation and a posterior contralateral negativity in the N1 time window (N1pc). N2pc reflected a subsequent attentional bias in favor of the relevant luminance change. The continuation of the sustained posterior contralateral negativity (SPCN) following N2pc covaried with response times within feature change conditions and revealed a posterior topography comparable to the earlier components associated with sensory and attentional mechanisms. Therefore, this component might reflect the re-processing of information based on sustained short-term memory representations in the visual system until a stable target percept is created that can serve as the perceptual basis for response selection and the initiation of goal-directed behavior.