Measuring event-related potentials (ERPs) has been fundamental to our understanding of how language is encoded in the brain. One particular ERP response, the N400 response, has been especially influential as an index of lexical and semantic processing. However, there remains a lack of consensus on the interpretation of this component. Resolving this issue has important consequences for neural models of language comprehension. Here we show that evidence bearing on where the N400 response is generated provides key insights into what it reflects. A neuroanatomical model of semantic processing is used as a guide to interpret the pattern of activated regions in functional MRI, magnetoencephalography and intracranial recordings that are associated with contextual semantic manipulations that lead to N400 effects.

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When a word is preceded by a supportive context such as a semantically associated word or a strongly constraining sentence frame, the N400 component of the ERP is reduced in amplitude. An ongoing debate is the degree to which this reduction reflects a passive spread of activation across long-term semantic memory representations as opposed to specific predictions about upcoming input. We addressed this question by embedding semantically associated prime-target pairs within an experimental context that encouraged prediction to a greater or lesser degree. The proportion of related items was used to manipulate the predictive validity of the prime for the target while holding semantic association constant. A semantic category probe detection task was used to encourage semantic processing and to preclude the need for a motor response on the trials of interest. A larger N400 reduction to associated targets was observed in the high than the low relatedness proportion condition, consistent with the hypothesis that predictions about upcoming stimuli make a substantial contribution to the N400 effect. We also observed an earlier priming effect (205–240 ms) in the high proportion condition, which may reflect facilitation due to form-based prediction. In sum, the results suggest that predictability modulates N400 amplitude to a greater degree than the semantic content of the context.

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Previous studies have found that English speakers experience attraction effects when comprehending subject-verb agreement, showing eased processing of ungrammatical sentences that contain a syntactically unlicensed but number-matching noun. In four selfpaced reading experiments we examine whether attraction effects also occur in Spanish, a language where agreement morphology is richer and functionally more significant. We find that despite having a richer morphology, Spanish speakers show reliable attraction effects in comprehension, and that these effects are strikingly similar to those previously found in English in their magnitude and distributional profile. Further, we use distributional analyses to argue that cue-based memory retrieval is used as an error-driven mechanism in comprehension. We suggest that cross-linguistic similarities in agreement attraction result from speakers deploying repair or error-driven mechanisms uniformly across languages.

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A core property of human semantic processing is the rapid, facilitatory influence of prior input on extracting the meaning of what comes next, even under conditions of minimal awareness. Previous work has shown a number of neurophysiological indices of this facilitation, but the mapping between time course and localization-critical for separating automatic semantic facilitation from other mechanisms-has thus far been unclear. In the current study, we used a multimodal imaging approach to isolate early, bottom-up effects of context on semantic memory, acquiring a combination of electroencephalography (EEG), magnetoencephalography (MEG), and functional magnetic resonance imaging (fMRI) measurements in the same individuals with a masked semantic priming paradigm. Across techniques, the results provide a strikingly convergent picture of early automatic semantic facilitation. Event-related potentials demonstrated early sensitivity to semantic association between 300 and 500 ms; MEG localized the differential neural response within this time window to the left anterior temporal cortex, and fMRI localized the effect more precisely to the left anterior superior temporal gyrus, a region previously implicated in semantic associative processing. However, fMRI diverged from early EEG/MEG measures in revealing semantic enhancement effects within frontal and parietal regions, perhaps reflecting downstream attempts to consciously access the semantic features of the masked prime. Together, these results provide strong evidence that automatic associative semantic facilitation is realized as reduced activity within the left anterior superior temporal cortex between 300 and 500 ms after a word is presented, and emphasize the importance of multimodal neuroimaging approaches in distinguishing the contributions of multiple regions to semantic processing.