High density ERP indices of conscious and unconscious semantic priming

The existence of differential brain mechanisms of conscious and unconscious processing is a matter of debate nowadays. The present experiment explores whether conscious and unconscious semantic priming in a lexical decision task at a long prime-target stimulus onset asynchrony (SOA) correlate with overlapping or different event related potential (ERP) effects. Results show that the N400 effect, which appeared when words were consciously perceived, completely disappeared when primes were masked at a level where the ability of participants to detect the prime was near chance. Instead, a rather different set of ERP effects was found to index unconscious semantic priming. This suggests that the processes at the basis of conscious and unconscious semantic analyses can under some circumstances be rather different. Moreover, our results support the notion that conscious and unconscious processes are at least partially separable in the brain.     

Dissociation of conscious and unconscious repetition priming effect on event-related potentials

Although the neural basis of the unconscious priming effect has previously been investigated, the results of these studies have possibly been contaminated by a conscious priming effect. The aim of the present study was to dissociate the effects of conscious and unconscious priming on event-related potential (ERP) by using the process-dissociation procedure. A prime word was presented briefly, followed by a word-stem, in each trial. Under the inclusion condition, subjects were instructed to complete the word-stem using the prime word, while under the exclusion condition subjects were asked to complete the word-stem with a word not seen as prime. The behavioral priming effect was obtained under both conditions, indicating that the prime words were processed unconsciously and influenced the word-stem completion task. We found that two ERP components were affected by repetition priming. First, the N400 amplitude was decreased by word repetition under the inclusion condition, but not under the exclusion condition. This result suggests that N400 would reflect conscious lexical processing, but not unconscious lexical activation. Second, the negativity at left front lateral region was enhanced by word repetition under the exclusion condition. We discuss this finding herein in relation to the activity of the left inferior prefrontal cortex with regard to word semantic processing.     

The N400 is modulated by unconsciously perceived masked words: further evidence for an automatic spreading activation account of N400 priming effects

It is a matter of debate whether the N400 component of the event-related brain potential (ERP) is sensitive to unconscious automatic priming mechanisms or to strategic mechanisms only. Recent studies demonstrated N400 modulation by masked primes at a short SOA supporting an automatic spreading activation account. However, it cannot be ruled out that strategic mechanisms based upon partial prime identification contributed to the observed priming effects. The present study was set up to substantiate masked N400 priming effects as an index of automatic spreading activation. It was assessed whether partial identification of the masked words due to backward priming could have supported strategic priming to occur. In experiment 1, ERPs were recorded while subjects performed lexical decisions on targets preceded by masked and unmasked primes at an SOA of 67 ms. Masked words, which were not consciously perceived, as well as visible words were shown to modulate the N400 to meaningfully related target words. Experiment 2 required subjects to perform decisions on visual, lexical and semantic features of masked words presented with or without semantically related context words. Subjects performed at chance in all tasks. Furthermore, the results exclude the possibility that backward priming has rendered the masked words partially visible. The present study therefore demonstrates that N400 priming effects can be reliably obtained from unconsciously perceived masked words at a very short SOA and strengthens the notion that the N400 is modulated by automatic spreading activation and not exclusively by strategic semantic processes.     

Neural capacity limits during unconscious semantic processing

A growing body of neuroimaging data suggests that direct measurements of brain activity can reveal subliminal effects that remain invisible with behavior measures alone. We examined whether sentence comprehension processes could be triggered by a sequence of masked words. On each trial, participants viewed a rapid sequence of masked or unmasked words, including a subject noun, three adverbs and followed by a visible target verb. To probe the capacity limits of unconscious processing, we measured event‐related potentials associated with the semantic congruency between the noun and the verb, while varying the subject position in each sentence. Unmasked sentences produced significant behavioral effects of congruency, paralleled by robust N400 effects, independently of subject–verb distance. By contrast, masked sentences produced no behavioral effect and elicited N400 effects only when subjects and verbs were separated by 0 or 1 word. The present results suggest that semantic integration of multiple words can occur unconsciously only if the distance between the words to be integrated does not exceed two words. Although the possibility remains that even longer sequence of invisible words may produce similar neural effects in different experimental settings, our ERP data show that only conscious perception gives access to a buffer that enables robust sentence‐level processing independently of temporal distance.     

Visual semantic features are activated during the processing of concrete words: event-related potential evidence for perceptual semantic priming

There has been conflicting evidence to date regarding the existence of non-strategic semantic priming based on semantic similarity, and in particular on visual-perceptual semantic features (e.g., button-coin: words refer to objects with the same global shape). Both event-related potential (ERP) and reaction time (RT) measures were employed to investigate visual-perceptual semantic priming in a word-pair lexical decision task designed to minimise the contribution of conscious strategic processing. While no RT priming effect was observed, a robust priming effect was obtained on the N400 component of the ERP. This result shows that semantic priming, as indexed by the N400 component, can be supported by nonassociative visual-perceptual semantic relations. The data are consistent with perceptual form information being accessed during the processing of concrete words, and provide support for models of semantic representation which incorporate semantic features and form information.     

Separating phonological and semantic processing in auditory sentence processing: a high-resolution event-related brain potential study

Phonological and semantic processing was studied using high-resolution event-related brain potentials (ERPs) during a sentence-matching task to investigate the spatial distribution of the phonological mismatch negativity (PMN) and the N400 response. It was hypothesized that the two components were spatially separable and that the activity matched prior localization knowledge. Participants examined visual-auditory sentence pairs that related within a semantic hierarchy (e.g., visual: "The man is teaching in the classroom"; Auditory: "The man is in the em leader school/barn"). Semantic congruency was varied for the final words of the spoken sentences. Incongruent words mismatched expectation in terms of both the initial phonological features (unexpected sound) and semantic features (unexpected meaning). In addition, the category-exemplar probability of the final words was either high or low, with low probability words being more difficult to anticipate. Low probability words were predicted to selectively affect PMN activity. We found that incongruent words elicited a PMN (287 msec) and a N400 (424 msec), for both the high and low probability words. As expected, low probability congruent words elicited a small PMN but no N400. In contrast, high probability congruent words elicited neither a detectible PMN nor a N400. The primary PMN sources were in left inferior frontal and inferior parietal lobes. The primary N400 source activation occurred along the left perisylvian cortex, consistent with prior N400 source localization work. From these results, it was concluded that the PMN and N400 were localized to separate cortical language (and memory) regions and had different source activation patterns.     

Repetition-priming modulates category-related effects on event-related potentials: further evidence for multiple cortical semantic systems

In the present study, the significance of category-related brain activations as an index of semantic memory structure was assessed within a repetition-priming paradigm during a lexical decision task. The interpretation of category-related effects has been debated since previous studies observed category-related brain activity mainly in tasks requiring explicit semantic categorization. Furthermore, categories were frequently associated with behavioral performance differences, which could have influenced the pattern of brain activation. Event-related potentials (ERPs) to words denoting objects from artifactual (e.g., tools) and natural categories (e.g., animals) were recorded while subjects were presented with words and pseudowords as distracters, which were repeatedly presented. Category-related ERP differences emerged in the time window of the N400, an electrophysiological index of semantic processing, over occipito-parietal and fronto-central regions as well as in the following window of a late positive potential. Repetition priming modulated these category-related ERP effects whereas behavioral repetition priming (faster reactions to repeated words) was comparable for both categories. Differences in ERP repetition effects were specifically due to diminished category-related activity at repeated presentation. The present results show that category-related brain activation is not confined to tasks requiring explicit semantic categorization. Most importantly, the study demonstrates that category-related brain activation can be specifically modulated by repetition priming in the absence of corresponding behavioral performance differences. These findings therefore substantiate the significance of category-related brain activations as reflections of semantic memory structure and support the notion of multiple cortical semantic systems.     

Automatic activation of the semantic network in schizophrenia: evidence from event-related brain potentials.

BACKGROUND: Language disorder associated with schizophrenia might be due to disturbances in both automatic activation and mechanisms of controlled attention. The contribution of each process to semantic memory dysfunction has not been determined for schizophrenia, and the semantic priming paradigm is well-suited for addressing this question. In the present report, event-related potentials (ERPs) elicited under conditions assumed to reveal automatic activation (short prime-target interval and low proportion of related words) are compared directly with ERPs elicited under conditions associated with controlled processing (long prime-target interval and high proportion of related words). METHODS: Visual ERPs were recorded during a lexical decision task, in which semantic relationship (associated and unassociated words), expectancy (relatedness proportions), and prime-target interval (250- and 850-msec inter-stimulus intervals [ISIs]) were varied. Diagnosis and expectancy were between-subjects factors; semantic relationship and ISI were repeated measures. The N400 priming effect (enhanced negativity to unassociated words) was compared between 34 male normal control subjects tested once and 37 male schizophrenia inpatients evaluated during their participation in a double-blind haloperidol maintenance therapy and placebo replacement protocol. RESULTS: The N400 priming effect for patients was significantly reduced during both pharmacologic phases, compared with controls. During haloperidol treatment, however, patients showed a significant N400 priming effect over the anterior scalp region and additionally under the automatic activation condition. The N400 priming effect was enhanced under the controlled processing condition for control subjects; this effect was not observed for patients. N400 amplitude elicited under the rapid presentation rate (250-msec ISI) differed between medicated patients and controls; groups did not differ for the 850-msec ISI. CONCLUSIONS: Findings suggest that automatic activation and mechanisms of controlled attention are both disrupted during semantic memory access for schizophrenia patients. Pharmacologic agents, such as haloperidol, might enhance automatic activation of the semantic network in this patient population, as indexed by the N400 component of the ERP.     

Is the N400 category-specific? A face and language processing study

N400 event-related potential (ERP) components have been observed during semantic incongruity detection in language, face identity and/or expression. However, it is still unclear whether semantic processing is functionally equivalent, since no study has directly investigated within the same participants the occurrence of the N400s for language and faces. We recorded ERPs while subjects performed incongruity detection on words, facial identities and facial expressions, with conditions matched to involve context integration. N400s were identified on central-parietal electrodes only for language and face identity processing. Scalp topographies of these N400s differed but a LORETA inverse solution identified a common functional generator in the left lateral frontal cortex, suggesting a general role of this brain region in selecting and contextually integrating semantic information.     

Localization of syntactic and semantic brain responses using magnetoencephalography

Electrophysiological methods have been used to study the temporal sequence of syntactic and semantic processing during sentence comprehension. Two responses associated with syntactic violations are the left anterior negativity (LAN) and the P600. A response to semantic violation is the N400. Although the sources of the N400 response have been identified in the left (and right) temporal lobe, the neural signatures of the LAN and P600 have not been revealed. The present study used magnetoencephalography to localize sources of syntactic and semantic activation in Finnish sentence reading. Participants were presented with sentences that ended in normally inf lected nouns, nouns in an unacceptable case, verbs instead of nouns, or nouns that were correctly inflected but made no sense in the context. Around 400 msec, semantically anomalous last words evoked strong activation in the left superior temporal lobe with significant activation also for word class errors (N400). Weaker activation was seen for the semantic errors in the right hemisphere. Later, 600-800 msec after word onset, the strongest activation was seen to word class and morphosyntactic errors (P600). Activation was significantly weaker to semantically anomalous and correct words. The P600 syntactic activation was localized to bilateral sources in the temporal lobe, posterior to the N400 sources. The results suggest that the same general region of the superior temporal cortex gives rise to both LAN and N400 with bilateral reactivity to semantic manipulation and a left hemisphere effect to syntactic manipulation. The bilateral P600 response was sensitive to syntactic but not semantic factors.     

N400 and automatic semantic processing abnormalities in patients with schizophrenia

BACKGROUND: One factor hypothesized to underlie thinking disturbance in patients with schizophrenia is abnormal or disinhibited automatic spreading activation of semantic networks, which can be assessed using the N400 event-related potential. N400 is a negative-going component elicited at about 400 milliseconds following semantic stimuli that are not primed by the preceding context. Semantic priming refers to facilitated semantic processing gained through preexposure to semantic context, which can happen automatically or strategically. Using N400, inferences can be drawn regarding the extent to which a given context primes a word. METHODS: During a picture-word matching task, N400s to primed (exact match) and unprimed (remotely related) words were recorded from 18 healthy control subjects and 18 patients with schizophrenia performing a picture-word matching task. A short interval (325 milliseconds) between picture and word onset (stimulus-onset asynchrony) was used to optimize the role of automatic spreading semantic activation and to minimize the role of attention, expectancy, preparation, and working memory. RESULTS: Despite behavioral evidence of normal semantic priming, patients generated an abnormally small N400 (ie, less negative) to unprimed words. The N400 to primed words was neither larger nor smaller in patients than in controls, suggesting normal use of context. CONCLUSIONS: A reduced N400 to unprimed words in patients with schizophrenia suggests that there was inappropriate priming of words by remotely related semantic contexts. This is consistent with an overly broad automatic spread of activation through semantic networks in patients with schizophrenia.     

Lack of repetition priming effect on visual event-related potentials in schizophrenia.

BACKGROUND: The present study was designed to assess, using event-related potentials, whether aberrant semantic processing reported in schizophrenia results from primary semantic overactivation or contextual dysregulation. METHODS: The visual event-related brain potentials were compared between 9 schizophrenic subjects and 16 normal control subjects performing two kinds of semantic categorization tasks with different nontarget stimuli: 1) nontargets comprising words, pseudowords, and unpronounceable foreign letters and 2) nontargets comprising initial presenting words, immediate repetition words, and delayed repetition words. RESULTS: Schizophrenic subjects showed no evidence suggestive of a greater negative potential associated with words and pseudowords, but they did show a lack of amplitude change associated with immediately repeated words relative to that in control subjects. CONCLUSIONS: These results suggest that aberrant semantic activation in schizophrenia results mainly from a failure to utilize information from preceding words or context, and could explain the increased N400 to the congruent or related words recently reported in this disease.     

N400-like semantic incongruity effect in 19-month-olds: processing known words in picture contexts

To understand mechanisms of early language acquisition, it is important to know whether the child's brain acts in an adult-like manner when processing words in meaningful contexts. The N400, a negative component in the eventrelated potential (ERP) of adults, is a sensitive index of semantic processing reflecting neural mechanisms of semantic integration into context. In the present study, we investigated whether the mechanisms indexed by the N400 are already working during early language acquisition. While 19-month-olds were looking at sequentially presented pictures, they were acoustically presented with words that were either congruous or incongruous to the picture content. The ERP averaged across the group of 55 children revealed an N400-like semantic incongruity effect in addition to an early phonological-lexical priming effect. The results suggest that both lexical expectations facilitating early phonological processing and mechanisms of semantic priming facilitating integration into semantic context are already present in 19-month-olds. The child's specific comprehension abilities are reflected in strength, latency, and hemispheric differences of the semantic incongruity effect. Spatio-temporal differences in that effect, thus, indicate changes in the organization of brain activity correlated with the child's behavioral development.     

Electrophysiological evidence for primary semantic memory functional organization deficits in schizophrenia

N400, an event-related brain potential (ERP) waveform elicited by meaningful stimuli, is normally reduced by stimulus repetition (N400 repetition priming), and relatedness between the eliciting stimulus and preceding ones (relatedness priming). Schizophrenia patients' N400 relatedness priming deficits suggest impairment in using meaningful prime stimuli to facilitate processing of related concepts in semantic memory. To examine whether this deficiency arises from difficulty activating the prime concept per se, as indexed by reduced N400 repetition priming; or from impaired functional connections among concepts in semantic memory, as reflected by reduced relatedness priming but normal repetition priming; we recorded ERPs from 16 schizophrenia patients and 16 controls who viewed prime words each followed at 300- or 750-ms stimulus-onset asynchrony (SOA) by an unrelated, related or repeated target word, or a nonword, in a lexical-decision task. In both groups, N400s were largest (most negative) for unrelated, intermediate for related, and smallest for repeated targets. Schizophrenia patients exhibited subnormal N400 relatedness priming at the 300-ms SOA, but normal repetition priming at both SOAs, suggesting that their impairment in using prime words to activate related concepts results from abnormal functional connections among concepts within semantic memory, rather than inability to activate the prime concept itself.     

Priming illusory words: an ERP approach

Repetition blindness (RB) was used to investigate whether illusory words emerge at a lexical-perceptual or a semantic-reconstructional level. Illusory words were evoked by the rapid serial visual presentation (RSVP) of two real words and a word fragment. The initial words share the same string of letters ("CREEP"-"SHEEP"), producing a free-floating word fragment ("SH"). This fragment is likely to be linked to a subsequently presented fragment ("IFT") if both combine to a meaningful word ("SHIFT"). The processing level of the illusions was probed by prime words preceding the RSVP sequence which were semantically related or unrelated to the second real word or to the illusion. Behavioural and electrophysiological correlates of the semantic priming effect were recorded in 14 subjects. Real words related to the prime were perceived more frequently, and evoked widespread N400-like effect in the event-related brain potentials (ERPs). An ERP effect of the same polarity was obtained for illusory words, however, its latency was delayed and the topographical distribution was restricted to left posterior electrode positions. These differences suggest that priming might affect real and illusory words at different levels of word processing: access to real words is facilitated at a semantic level, whereas lexical activation apparently accompanies the generation of illusory words.     

Attentional modulation of unconscious "automatic" processes: evidence from event-related potentials in a masked priming paradigm

Automatic processes are usually thought to occur independently of any cognitive resources. This traditional view has been recently challenged by showing that temporal attention to a target stimulus is a prerequisite for "automatic" response priming. The event-related potential (ERP) study reported here extends this research by pursuing a somewhat different approach. In two experiments, it was investigated whether masked semantic priming effects can be modulated by temporal attention to the prime using a cueing procedure. We hypothesized that masked priming is amplified when attention is directed to the stimulus stream in the time window of masked prime presentation, even in the absence of any prime awareness. ERPs were recorded while subjects performed a primed lexical decision task. Target words were preceded by semantically related or unrelated masked prime words, which were not consciously identified. A cue stimulus prompted subjects to direct their attention to the stimulus stream either shortly before the masked prime (short cue interval) or a long time interval before. Priming affected the amplitude of the N400 ERP component, an electrophysiological index of semantic processing. Unrelated prime-target pairs elicited a larger N400 than related pairs (N400 priming effect). Most importantly, this masked N400 priming effect was strongest when the cue interval and the stimulus onset asynchronies were short. The present results show that temporal attention to the prime is a prerequisite for obtaining masked N400 priming effects. They also demonstrate that unconscious automatic processes are susceptible to attentional modulation.     

Imaginal, semantic, and surface-level processing of concrete and abstract words: an electrophysiological investigation

Words representing concrete concepts are processed more quickly and efficiently than words representing abstract concepts. Concreteness effects have also been observed in studies using event-related brain potentials (ERPs). The aim of this study was to examine concrete and abstract words using both reaction time (RT) and ERP measurements to determine (1) at what point in the stream of cognitive processing concreteness effects emerge and (2) how different types of cognitive operations influence these concreteness effects. Three groups of subjects performed a sentence verification task in which the final word of each sentence was concrete or abstract. For each group the truthfulness judgment required either (1) image generation, (2) semantic decision, or (3) evaluation of surface characteristics. Concrete and abstract words produced similar RTs and ERPs in the surface task, suggesting that postlexical semantic processing is necessary to elicit concreteness effects. In both the semantic and imagery tasks, RTs were shorter for concrete than for abstract words. This difference was greatest in the imagery task. Also, in both of these tasks concrete words elicited more negative ERPs than abstract words between 300 and 550 msec (N400). This effect was widespread across the scalp and may reflect activation in a linguistic semantic system common to both concrete and abstract words. ERPs were also more negative for concrete than abstract words between 550 and 800 msec. This effect was more frontally distributed and was most evident in the imagery task. We propose that this later anterior effect represents a distinct ERP component (N700) that is sensitive to the use of mental imagery. The N700 may reflect the a access of specific characteristics of the imaged item or activation in a working memory system specific to mental imagery. These results also support the extended dual-coding hypothesis that superior associative connections and the use of mental imagery both contribute to processing advantages for concrete words over abstract words.     

Evidence of conscious and subconscious olfactory information processing during word encoding: a magnetoencephalographic (MEG) study

The present study was meant to distinguish between unconscious and conscious olfactory information processing and to investigate the influence of olfaction on word information processing. Magnetic field changes were recorded in healthy young participants during deep encoding of visually presented words whereby some of the words were randomly associated with an odor. All recorded data were then split into two groups. One group consisted of participants who did not consciously perceive the odor during the whole experiment whereas the other group did report continuous conscious odor perception. The magnetic field changes related to the condition 'words without odor' were subtracted from the magnetic field changes related to the condition 'words with odor' for both groups. First, an odor-induced effect occurred between about 200 and 500 ms after stimulus onset which was similar in both groups. It is interpreted to reflect an activity reduction during word encoding related to the additional olfactory stimulation. Second, a later effect occurred between about 600 and 900 ms after stimulus onset which differed between the two groups. This effect was due to higher brain activity related to the additional olfactory stimulation. It was more pronounced in the group consisting of participants who consciously perceived the odor during the whole experiment as compared to the other group. These results are interpreted as evidence that the later effect is related to conscious odor perception whereas the earlier effect reflects unconscious olfactory information processing. Furthermore, our study provides evidence that only the conscious perception of an odor which is simultaneously presented to the visual presentation of a word reduces its chance to be subsequently recognized.     

Modulation of language processing in schizophrenia: effects of context and haloperidol on the event-related potential.

BACKGROUND: Disturbances in language associations were among the first clinical symptoms reported for individuals described as schizophrenic (Bleuler 1911/1950). Currently, associative language disturbance is a diagnostic feature of schizophrenia (American Psychiatric Association 1994); however, the mechanisms that produce this symptom remain unknown. In the present study, two candidate psychological functions were examined: sensitivity to semantic context and expectancy (attention). METHODS: Visual event-related potentials were recorded during a lexical decision task in which semantic relationship and expectancy (relatedness proportions) were varied. Semantic priming processes were compared between 34 male normal control subjects tested once and 37 male schizophrenic inpatients evaluated during their participation in a double-blind haloperidol maintenance therapy and placebo replacement protocol. RESULTS: Schizophrenic patients failed to discriminate between associated and unassociated words, as measured by the amplitude of the N400 component (i.e., absence of the N400 priming effect); however, the overall mean amplitude of N400 did not differ between patients and control subjects. In addition, patients and control subjects did not differ significantly in the amplitude of N400 elicited to associated words or to unassociated words. Finally, the effect of expectancy-based processing on the magnitude of the N400 priming effect did not differ between patients and control subjects. CONCLUSIONS: On the basis of these findings, a tentative hypothesis is suggested that schizophrenic patients are characterized by a pattern of indiscriminate or random spread of activation in their semantic network during the processing of single-word semantic contexts.     

Auditory semantic networks for words and natural sounds

Does lexical processing rely on a specialized semantic network in the brain, or does it draw on more general semantic resources? The primary goal of this study was to compare behavioral and electrophysiological responses evoked during the processing of words, environmental sounds, and non-meaningful sounds in semantically matching or mismatching visual contexts. A secondary goal was to characterize the dynamic relationship between the behavioral and neural activities related to semantic integration using a novel analysis technique, ERP imaging. In matching trials, meaningful-sound ERPs were characterized by an extended positivity (200-600 ms) that in mismatching trials partly overlapped with centro-parietal N400 and frontal N600 negativities. The mismatch word-N400 peaked later than the environmental sound-N400 and was only slightly more posterior in scalp distribution. Single-trial ERP imaging revealed that for meaningful stimuli, the match-positivity consisted of a sensory P2 (200 ms), a semantic positivity (PS, 300 ms), and a parietal response-related positivity (PR, 500-800 ms). The magnitudes (but not the timing) of the N400 and PS activities correlated with subjects' reaction times, whereas both the latency and magnitude of the PR was correlated with subjects' reaction times. These results suggest that largely overlapping neural networks process verbal and non-verbal semantic information. In addition, it appears that semantic integration operates across different time scales: earlier processes (indexed by the PS and N400) utilize the established meaningful, but not necessarily lexical, semantic representations, whereas later processes (indexed by the PR and N600) are involved in the explicit interpretation of stimulus semantics and possibly of the required response.