Neural Evidence for the Prediction of Animacy Features during Language Comprehension: Evidence from MEG and EEG Representational Similarity Analysis.

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.SIGNIFICANCE STATEMENT Language inputs unfold very quickly during real-time communication. By predicting ahead, we can give our brains a "head start," so that language comprehension is faster and more efficient. Although most contexts do not constrain strongly for a specific word, they do allow us to predict some upcoming information. For example, following the context of "they cautioned the…," we can predict that the next word will be animate rather than inanimate (we can caution a person, but not an object). Here, we used EEG and MEG techniques to show that the brain is able to use these contextual constraints to predict the animacy of upcoming words during sentence comprehension, and that these predictions are associated with specific spatial patterns of neural activity.

A Tale of Two Positivities and the N400: Distinct Neural Signatures Are Evoked by Confirmed and Violated Predictions at Different Levels of Representation.

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, for example, {Agent caution animate-Patient}. High constraint contexts additionally constrained for a specific event/lexical item, for example, 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.

Alpha and theta band dynamics related to sentential constraint and word expectancy.

Despite strong evidence for prediction during language comprehension, the underlying mechanisms, and the extent to which they are specific to language, remain unclear. Re-analyzing an ERP study, we examined responses in the time-frequency domain to expected and unexpected (but plausible) words in strongly and weakly constraining sentences, and found results similar to those reported in nonverbal domains. Relative to expected words, unexpected words elicited an increase in the theta band (4-7 Hz) in strongly constraining contexts, suggesting the involvement of control processes to deal with the consequences of having a prediction disconfirmed. Prior to critical word onset, strongly constraining sentences exhibited a decrease in the alpha band (8-12 Hz) relative to weakly constraining sentences, suggesting that comprehenders can take advantage of predictive sentence contexts to prepare for the input. The results suggest that the brain recruits domain-general preparation and control mechanisms when making and assessing predictions during sentence comprehension.

Time for prediction? The effect of presentation rate on predictive sentence comprehension during word-by-word reading.

Predictive processing is a core component of normal language comprehension, but the brain may not engage in prediction to the same extent in all circumstances. This study investigates the effects of timing on anticipatory comprehension mechanisms. Event-related brain potentials (ERPs) were recorded while participants read two-sentence mini-scenarios previously shown to elicit prediction-related effects for implausible items that are categorically related to expected items ('They wanted to make the hotel look more like a tropical resort. So along the driveway they planted rows of PALMS/PINES/TULIPS.'). The first sentence of every pair was presented in its entirety and was self-paced. The second sentence was presented word-by-word with a fixed stimulus onset asynchrony (SOA) of either 500 msec or 250 msec that was manipulated in a within-subjects blocked design. Amplitudes of the N400 ERP component are taken as a neural index of demands on semantic processing. At 500 msec SOA, implausible words related to predictable words elicited reduced N400 amplitudes compared to unrelated words (PINES vs TULIPS), replicating past studies. At 250 msec SOA this prediction-related semantic facilitation was diminished. Thus, timing is a factor in determining the extent to which anticipatory mechanisms are engaged. However, we found evidence that prediction can sometimes be engaged even under speeded presentation rates. Participants who first read sentences in the 250 msec SOA block showed no effect of semantic similarity for this SOA, although these same participants showed the effect in the second block with 500 msec SOA. However, participants who first read sentences in the 500 msec SOA block continued to show the N400 semantic similarity effect in the 250 msec SOA block. These findings add to results showing that the brain flexibly allocates resources to most effectively achieve comprehension goals given the current processing environment.

Two sides of meaning: the scalp-recorded n400 reflects distinct contributions from the cerebral hemispheres.

The N400, a component of the event-related potential (ERP) associated with the processing of meaning, is sensitive to a wide array of lexico-semantic, sentence-level, and discourse-level manipulations across modalities. In sentence contexts, N400 amplitude varies inversely and nearly linearly with the predictability of a word in its context. However, recent theories and empirical evidence from studies employing the visual half-field technique (to selectively bias processing to one cerebral hemisphere) suggest that the two hemispheres use sentence context information in different ways. Thus, each hemisphere may not respond to manipulations of contextual predictability in an equivalent manner. This possibility was investigated by recording ERPs while presenting [in the left and right visual fields (VFs)] sentence-final words that varied over the full range of sentence-level predictability. RVF/left hemisphere items were facilitated (as evidenced by reduced N400 amplitudes) over a broader range of predictability compared with LVF/right hemisphere items, although both strongly predictable and completely unexpected items evoked similar responses in each VF/hemisphere. Further, the pattern of N400 amplitudes over the full range of predictability significantly differed from a linear response function for both VFs/hemispheres. This suggests that the N400 response recorded with standard central field presentation comprises different contributions from both cerebral hemispheres, neither of which on its own is sensitive to contextual predictability in an evenly graded manner. These data challenge the notion of a singular or unitary mode of comprehension and instead support the view that the left and right hemispheres instantiate unique, complementary language comprehension architectures in parallel.

To predict or not to predict: age-related differences in the use of sentential context.

Older adults (as a group) are less likely than younger adults to engage in an anticipatory mode of language comprehension, failing to successfully preactivate information about upcoming likely (predictable) words during online processing. To assess (within one set of materials) age-related changes in the use of sentential context to affect processing of predictable words and in the consequences of violating predictions, event-related brain potentials were recorded while older adults read sentences that varied in sentence-level constraint and expectancy of sentence-final words. Strongly constraining sentences were completed by their most expected, predictable words and weakly constraining sentences were completed by their most expected, less predictable words. Both types of sentences also were completed by unexpected (but plausible) words. Older adults showed reduced and delayed effects of sentential context on processing predictable words. Whereas younger adults elicit an enhanced positive ERP (starting around 500 ms poststimulus onset, largest over prefrontal electrode sites), specifically for unexpected words that violate strong expectancies for a different word, older adults as a group did not exhibit this neural consequence of disconfirmed predictions. Older adults were instead more likely to show a left-lateralized frontal negativity for predictable items. This ERP response has been attributed to processes needed to revisit contextual material in forming an interpretation of message-level meaning, which may be more likely when anticipatory modes of comprehension are not engaged. Taken together, the results suggest that normal aging can affect allocation of resources to different cognitive and neural pathways in achieving comprehension outcomes.

So that's what you meant! Event-related potentials reveal multiple aspects of context use during construction of message-level meaning.

Factors that modulate the influence of contextual information on semantic processing in language comprehension have been thoroughly investigated with the N400 component of the event-related potential (ERP), a direct measure of initial contact with semantic memory. Although context has a strong and immediate impact on processing, multiple mechanisms contribute to the construction of message-level representations during normal comprehension. Some of these may be engaged after or concurrent with the formation of an initial meaning representation, and can then serve to revise or reshape meaning. In this study, ERPs were recorded while participants read plausible sentences that continuously varied in the amount of contextual constraint for the sentence-final word, defined via extensive norming data including the range of possible alternative completions for the contexts. Consistent with numerous past studies, the amplitude of the N400 was graded with expectancy, as amplitudes decreased with increasing constraint. Additionally, a left-lateralized, broad, slow negativity onsetting around 400-500 ms was largest for sentences with moderately strong constraint. Within this range of constraint, the negativity was larger for sentences with fewer alternative completions compared to those with many different ones. The timing and scalp distribution of the effect resemble brain responses linked to engagement of working memory resources, ambiguity resolution, and comprehension of jokes. Similar to cases of "frame-shifting" in non-literal language, this effect may reflect processing associated with reinterpretation or reconsideration of contextual material when multiple interpretations of a sentence were likely.

Age-related changes in the impact of contextual strength on multiple aspects of sentence comprehension.

Contextual information influences multiple aspects of language comprehension extended over time. To determine how age-related changes impact normal comprehension, effects of contextual strength were examined with event-related potentials. Increased contextual constraint facilitated semantic processing (reduced N400s). Effects were smaller and delayed for older adults, and sensitivity to contextual information was diminished for weak contexts. Both groups elicited a later left-lateralized frontal negativity associated with reinterpretation of context when multiple interpretations of a sentence were likely. Older adults evidenced the frontal negativity over a wider range of constraint. The change in the use of contextual information across age is attributed to decreased reliance on predictive processing for older adults. Thus, age-related changes in comprehension lead to differential engagement of processing resources over time for older adults.

Language of the aging brain: Event-related potential studies of comprehension in older adults.

Normal aging brings increased richness in knowledge and experience as well as declines in cognitive abilities. Event-related brain potential (ERP) studies of language comprehension corroborate findings showing that the structure and organization of semantic knowledge remains relatively stable with age. Highlighting the advantages of the temporal and functional specificity of ERPs, this survey focuses on age-related changes in higher-level processes required for the successful comprehension of meaning representations built from multiple words. Older adults rely on different neural pathways and cognitive processes during normal, everyday comprehension, including a shift away from the predictive use of sentential context, differential recruitment of neural resources, and reduced engagement of controlled processing. Within age groups, however, there are important individual differences that, for example, differentiate a subset of older adults whose processing patterns more closely resemble that of young adults, providing a window into cognitive skills and abilities that may mediate or moderate age-related declines.

What's "right" in language comprehension: ERPs reveal right hemisphere language capabilities.

Although the term "nonverbal" is often applied to the right cerebral hemisphere (RH), a growing body of work indicates that the RH can comprehend language and, indeed, that it makes critical contributions to normal language functions. Reviewed here are studies that have examined RH language capabilities by combining visual half-field presentation methods with event-related potential (ERP) measures. Because they afford temporal and functional specificity and can be obtained as participants simply process language for meaning, ERPs provide especially valuable insights into RH language functions. Such studies suggest that the RH appreciates word and message-level meaning information, and that it may play a particularly important role in the processing of relatively unpredictable semantic relationships. In addition, this work suggests that patterns observed for everyday language processing may often be an emergent property of multiple, distinct mechanisms operating in parallel as the left and right hemispheres jointly comprehend language.

Finding the right word: hemispheric asymmetries in the use of sentence context information.

The cerebral hemispheres have been shown to be differentially sensitive to sentence-level information; in particular, it has been suggested that only the left hemisphere (LH) makes predictions about upcoming items, whereas the right (RH) processes words in a more integrative fashion. The current study used event-related potentials to jointly examine the effects of expectancy and sentential constraint on word processing. Expected and unexpected but plausible words matched for contextual fit were inserted into strongly and weakly constraining sentence frames and presented to the left and right visual fields (LVF and RVF). Consistent with the prediction/integration view, the P2 was sensitive to constraint: words in strongly constraining contexts elicited larger P2s than those in less predictive contexts, for RVF/LH presentation only. N400 responses for both VFs departed from the typical pattern of amplitudes graded by cloze probability. Expected endings in strongly and weakly constraining contexts were facilitated to a similar degree with RVF/LH presentation, and expected endings in weakly constraining contexts were not facilitated compared to unexpected endings in those contexts for LVF/RH presentation. These data suggest that responses seen for central presentation reflect contributions from both hemispheres. Finally, a late positivity, larger for unexpected endings in strongly constraining contexts, observed for these stimuli with central presentation was not seen here for either VF. Thus, some phenomena observed with central presentation may be an emergent property of mechanisms that require interhemispheric cooperation. These data highlight the importance of understanding hemispheric asymmetries and their implications for normal language processing.

Multiple effects of sentential constraint on word processing.

Behavioral and electrophysiological studies have uncovered different patterns of constraint effects on the processing of words in sentences. Whereas response time measures have indicated a reduced scope of facilitation from strongly constraining contexts, event-related brain potential (ERP) measures have instead revealed enhanced facilitation for semantically related endings in such sentences. Given this disparity, and the concomitant possibility of functionally separable stages of context effects, the current study jointly examined expectancy (cloze probability) and constraint effects on the ERP response to words. Expected and unexpected (but plausible) words completed strongly and weakly constraining sentences; unexpected items were matched for contextual fit across the two levels of constraint and were semantically unrelated to the most expected endings. N400 amplitudes were graded by expectancy but unaffected by constraint and seemed to index the benefit of contextual information. However, a later effect, in the form of increased frontal positivity from 500 to 900 ms post-stimulus-onset, indicated a possible cost associated with the processing of unexpected words in strongly constraining contexts.

Word learning and individual differences in word learning reflected in event-related potentials.

Adults learned the meanings of rare words (e.g., gloaming) and then made meaning judgments on pairs of words. The 1st word was a trained rare word, an untrained rare word, or an untrained familiar word. Event-related potentials distinguished trained rare words from both untrained rare and familiar words, first at 140 ms and again at 400-600 ms after onset of the 1st word. These results may point to an episodic memory effect. The 2nd word produced an N400 that distinguished trained and familiar word pairs that were related in meaning from unrelated word pairs. Skilled comprehenders learned more words than less skilled comprehenders and showed a stronger episodic memory effect at 400-600 ms on the 1st word and a stronger N400 effect on the 2nd word. These results suggest that superior word learning among skilled comprehenders may arise from a stronger episodic trace that includes orthographic and meaning information and illustrate, how an episodic theory of word identification can explain reading skill.