ERP

To comprehend language, we continually use prior context to pre-activate expected upcoming information, resulting in facilitated processing of incoming words that confirm these predictions. But what are the consequences of disconfirming prior predictions? To address this question, most previous studies have examined unpredictable words appearing in contexts that constrain strongly for a single continuation. However, during natural language processing, it is far more common to encounter contexts that constrain for multiple potential continuations, each with some probability. Here, we ask whether and how pre-activating both higher and lower probability alternatives influences the processing of the lower probability incoming word. One possibility is that, similar to language production, there is continuous pressure to select the higher-probability pre-activated alternative through competitive inhibition. During comprehension, this would result in relative costs in processing the lower probability target. A second possibility is that if the two pre-activated alternatives share semantic features, they mutually enhance each other’s pre-activation. This would result in greater facilitation in processing the lower probability target. To distinguish between these accounts, we recorded ERPs as participants read three-sentence scenarios that constrained either for a single word or for two potential continuations – a higher probability expected candidate and a lower probability second-best candidate. We found no evidence that competitive pre-activation between the expected and second-best candidates resulted in costs in processing the second-best target, either during lexico-semantic processing (indexed by the N400) or at later stages of processing (indexed by a later frontal positivity). Instead, we found only benefits of pre-activating multiple alternatives, with evidence of enhanced graded facilitation on lower-probability targets that were semantically related to a higher-probability pre-activated alternative. These findings are consistent with a previous eye-tracking study by Luke and Christianson (2016, Cogn Psychol) using corpus-based materials. They have significant theoretical implications for models of predictive language processing, indicating that routine graded prediction in language comprehension does not operate through the same competitive mechanisms that are engaged in language production. Instead, our results align more closely with hierarchical probabilistic accounts of language comprehension, such as predictive coding.

We used magnetoencephalography (MEG) and event-related potentials (ERPs) to track the time-course and localization of evoked activity produced by expected, unexpected plausible, and implausible words during incremental language comprehension. We suggest that the full pattern of results can be explained within a hierarchical predictive coding framework in which increased evoked activity reflects the activation of residual information that was not already represented at a given level of the fronto-temporal hierarchy (“error” activity). Between 300 and 500 ms, the three conditions produced progressively larger responses within left temporal cortex (lexico-semantic prediction error), whereas implausible inputs produced a selectively enhanced response within inferior frontal cortex (prediction error at the level of the event model). Between 600 and 1,000 ms, unexpected plausible words activated left inferior frontal and middle temporal cortices (feedback activity that produced top-down error), whereas highly implausible inputs activated left inferior frontal cortex, posterior fusiform (unsuppressed orthographic prediction error/reprocessing), and medial temporal cortex (possibly supporting new learning). Therefore, predictive coding may provide a unifying theory that links language comprehension to other domains of cognition.

There is an ongoing controversy over whether readers can access the meaning of multiple words, simultaneously. To date, different experimental methods have generated seemingly contradictory evidence in support of serial or parallel processing accounts. For example, dual-task studies suggest that readers can process a maximum of one word at a time (White, Palmer & Boynton, 2018), while ERP studies have demonstrated neural priming effects that are more consistent with parallel activation (Wen, Snell & Grainger, 2019). To help reconcile these views, I measured neural responses and behavioral accuracy in a dual-task sentence comprehension paradigm. Participants saw masked sentences and two-word phrases and had to judge whether or not they were grammatical. Grammatically correct sentences (This girl is neat) produced smaller N400 responses compared to scrambled sentences (Those girl is fled): an N400 sentence superiority effect. Critically, participants’ grammaticality judgements on the same trials showed striking capacity limitations, with dual-task deficits closely matching the predictions of a serial, all-or-none processing account. Together, these findings suggest that the N400 sentence superiority effect is fully compatible with serial word recognition, and that readers are unable to process multiple sentence positions simultaneously.

The ability to detect and respond to linguistic errors is critical for successful reading comprehension, but these skills can vary considerably across readers. In the current study, healthy adults (age 18-35) read short discourse scenarios for comprehension while monitoring for the presence of semantic anomalies. Using a factor analytic approach, we examined if performance in nonlinguistic conflict monitoring tasks (Stroop, AX-CPT) would predict individual differences in neural and behavioral measures of linguistic error processing. Consistent with this hypothesis, domain-general conflict monitoring predicted both readers’ end-of-trial acceptability judgments and the amplitude of a late neural response (the P600) evoked by linguistic anomalies. The influence on the P600 was nonlinear, suggesting that online neural responses to linguistic errors are influenced by both the effectiveness and efficiency of domain-general conflict monitoring. These relationships were also highly specific and remained after controlling for variability in working memory capacity and verbal knowledge. Finally, we found that domain-general conflict monitoring also predicted individual variability in measures of reading comprehension, and that this relationship was partially mediated by behavioral measures of linguistic error detection. These findings inform our understanding of the role of domain-general executive functions in reading comprehension, with potential implications for the diagnosis and treatment of language impairments. 

Event-related potential (ERP) studies produce large spatiotemporal datasets. These rich datasets are key to the ability of ERP to help us understand cognition and neural processes. However, they can also present a massive multiple comparisons problem, leading to a high Type I error rate. Standard approaches to statistical analysis, which average over time windows and regions of interest, do not always control for Type I error, and their inflexibility can lead to low power to detect true effects. Mass univariate approaches offer an alternative, but have thus far been seen as appropriate only for exploratory analysis and only applicable to simple designs. Here we present new simulation studies showing that permutation-based mass univariate tests can be employed with complex factorial designs. Most importantly, we show that mass univariate approaches provide slightly greater power than traditional spatiotemporal averaging approaches when strong a priori time windows and spatial regions are used, and that power decreases only modestly when more exploratory spatiotemporal parameters are used. We argue that mass univariate approaches are preferable to traditional analysis approaches for most ERP studies.

It has been proposed that people can generate probabilistic predictions at multiple levels of representation during language comprehension. We used magnetoencephalography (MEG) and electroencephalography (EEG), in combination with representational similarity analysis, to seek neural evidence for the prediction of animacy features. In two studies, MEG and EEG activity was measured as human participants (both sexes) read three-sentence scenarios. Verbs in the final sentences constrained for either animate or inanimate semantic features of upcoming nouns, and the broader discourse context constrained for either a specific noun or for multiple nouns belonging to the same animacy category. We quantified the similarity between spatial patterns of brain activity following the verbs until just before the presentation of the nouns. The MEG and EEG datasets revealed converging evidence that the similarity between spatial patterns of neural activity following animate-constraining verbs was greater than following inanimate-constraining verbs. This effect could not be explained by lexical-semantic processing of the verbs themselves. We therefore suggest that it reflected the inherent difference in the semantic similarity structure of the predicted animate and inanimate nouns. Moreover, the effect was present regardless of whether a specific word could be predicted, providing strong evidence for the prediction of coarse-grained semantic features that goes beyond the prediction of individual words.

During language comprehension, online neural processing is strongly influenced by the constraints of the prior context. While the N400 ERP response (300-500ms) is known to be sensitive to a word’s semantic predictability, less is known about a set of late positive-going ERP responses (600-1000ms) that can be elicited when an incoming word violates strong predictions about upcoming content (late frontal positivity) or about what is possible given the prior context (late posterior positivity/P600). Across three experiments, we systematically manipulated the length of the prior context and the source of lexical constraint to determine their influence on comprehenders’ online neural responses to these two types of prediction violations. In Experiment 1, within minimal contexts, both lexical prediction violations and semantically anomalous words produced a larger N400 than expected continuations (James unlocked the door/laptop/gardener), but no late positive effects were observed. Critically, the late posterior positivity/P600 to semantic anomalies appeared when these same sentences were embedded within longer discourse contexts (Experiment 2a), and the late frontal positivity appeared to lexical prediction violations when the preceding context was rich and globally constraining (Experiment 2b). We interpret these findings within a hierarchical generative framework of language comprehension. This framework highlights the role of comprehension goals and broader linguistic context, and how these factors influence both top-down prediction and the decision to update or reanalyze the prior context when these predictions are violated.

It has been proposed that hierarchical prediction is a fundamental computational principle underlying neurocognitive processing. Here we ask whether the brain engages distinct neurocognitive mechanisms in response to inputs that fulfill versus violate strong predictions at different levels of representation during language comprehension. Participants read three-sentence scenarios in which the third sentence constrained for a broad event structure, e.g. Agent caution animate-Patient. High constraint contexts additionally constrained for a specific event/lexical item, e.g. a two-sentence context about a beach, lifeguards and sharks constrained for the event, Lifeguards cautioned Swimmers and the specific lexical item, “swimmers”. Low constraint contexts did not constrain for any specific event/lexical item. We measured ERPs on critical nouns that fulfilled and/or violated each of these constraints. We found clear, dissociable effects to fulfilled semantic predictions (a reduced N400), to event/lexical prediction violations (an increased late frontal positivity), and to event structure/animacy prediction violations (an increased late posterior positivity/P600). We argue that the late frontal positivity reflects a large change in activity associated with successfully updating the comprehender’s current situation model with new unpredicted information. We suggest that the late posterior positivity/P600 is triggered when the comprehender detects a conflict between the input and her model of the communicator and communicative environment. This leads to an initial failure to incorporate the unpredicted input into the situation model, which may be followed by second-pass attempts to make sense of the discourse through reanalysis, repair, or reinterpretation. Together, these findings provide strong evidence that confirmed and violated predictions at different levels of representation manifest as distinct spatiotemporal neural signatures.

It has been proposed that abnormalities in probabilistic prediction and dynamic belief updating explain multiple features of schizophrenia. Here, we used EEG to ask whether these abnormalities can account for the well-established reduction in semantic priming observed in schizophrenia under non-automatic conditions. We isolated predictive contributions to the neural semantic priming effect by manipulating the prime’s predictive validity and minimizing retroactive semantic matching mechanisms. We additionally examined the link between prediction and learning using a Bayesian model that probed dynamic belief updating as participants adapted to the increase in predictive validity. We found that patients were less likely than healthy controls to use the prime to predictively facilitate semantic processing on the target, resulting in a reduced N400 effect. Moreover, the trial-by-trial output of our Bayesian computational model explained between-group differences in trial-by-trial N400 amplitudes as participants transitioned from conditions of lower to higher predictive validity. These findings suggest that, compared to healthy controls, people with schizophrenia are less able to mobilize predictive mechanisms to facilitate processing at the earliest stages of accessing the meanings of incoming words. This deficit may be linked to a failure to adapt to changes in the broader environment. This reciprocal relationship between impairments in probabilistic prediction and Bayesian learning/adaptation may drive a vicious cycle that maintains cognitive disturbances in schizophrenia.

When semantic information is activated by a context prior to new bottom-up input (i.e. when a word is predicted), semantic processing of that incoming word is typically facilitated, attenuating the amplitude of the N400 event related potential (ERP) – a direct neural measure of semantic processing. N400 modulation is observed even when the context is a single semantically related “prime” word. This so-called “N400 semantic priming effect” is sensitive to the probability of encountering a related prime-target pair within an experimental block, suggesting that participants may be adapting the strength of their predictions to the predictive validity of their broader experimental environment. We formalize this adaptation using a Bayesian learning model that estimates and updates the probability of encountering a related versus an unrelated prime-target pair on each successive trial. We found that our model’s trial-by-trial estimates of target word probability accounted for significant variance in the amplitude of the N400 evoked by target words. These findings suggest that Bayesian principles contribute to how comprehenders adapt their semantic predictions to the statistical structure of their broader environment, with implications for the functional significance of the N400 component and the predictive nature of language processing.