The Impact of Linguistic Prediction Violations on Downstream Recognition Memory and Sentence Recall.

Predicting upcoming words during language comprehension not only affects processing in the moment but also has consequences for memory, although the source of these memory effects (e.g., whether driven by lingering pre-activations, re-analysis following prediction violations, or other mechanisms) remains underspecified. Here, we investigated downstream impacts of prediction on memory in two experiments. First, we recorded EEG as participants read strongly and weakly constraining sentences with expected, unexpected but plausible, or semantically anomalous endings ("He made a holster for his gun / father / train") and were tested on their recognition memory for the sentence endings. Participants showed similar rates of false alarms for predicted but never presented sentence endings whether the prediction violation was plausible or anomalous, suggesting that these arise from pre-activation of the expected words during reading. During sentence reading, especially in strongly constraining sentences, plausible prediction violations elicited an anterior positivity; anomalous endings instead elicited a posterior positivity, whose amplitude was predictive of later memory for those anomalous words. ERP patterns at the time of recognition differentiated plausible and anomalous sentence endings: Words that had been plausible prediction violations elicited enhanced late positive complex amplitudes, suggesting greater episodic recollection, whereas anomalous sentence endings elicited greater N1 amplitudes, suggesting attentional tagging. In a follow-up behavioral study, a separate group of participants read the same sentence stimuli and were tested for sentence-level recall. We found that recall of full sentences was impaired when sentences ended with a prediction violation. Taken together, the results suggest that prediction violations draw attention and affect encoding of the violating word, in a manner that depends on plausibility, and that this, in turn, may impair future memory of the gist of the sentence.

Differential Recruitment of Inhibitory Control Processes by Directed Forgetting and Thought Substitution.

Humans have the ability to intentionally forget information via different strategies, included suppression of encoding (directed forgetting) and mental replacement of the item to encode (thought substitution). These strategies may rely on different neural mechanisms; namely, encoding suppression may induce prefrontally mediated inhibition, whereas thought substitution is potentially accomplished through modulating contextual representations. Yet, few studies have directly related inhibitory processing to encoding suppression, or tested its involvement in thought substitution. Here, we directly tested whether encoding suppression recruits inhibitory mechanisms with a cross-task design, relating the behavioral and neural data from male and female participants in a Stop Signal task (a task specifically testing inhibitory processing) to a directed forgetting task with both encoding suppression (Forget) and thought substitution (Imagine) cues. Behaviorally, Stop Signal task performance (stop signal reaction times) was related to the magnitude of encoding suppression, but not thought substitution. Two complementary neural analyses corroborated the behavioral result. Namely, brain-behavior analysis demonstrated that the magnitude of right-frontal beta activity following stop signals was related to stop signal reaction times and successful encoding suppression, but not thought substitution; and classifiers trained to discriminate successful and unsuccessful stopping in the Stop Signal task could also classify successful and unsuccessful forgetting following Forget cues, but not Imagine cues. Importantly, inhibitory neural mechanisms were engaged following Forget cues at a later time than motor stopping. These findings not only support an inhibitory account of directed forgetting, and that thought substitution engages separate mechanisms, but also potentially identify a specific time in which inhibition occurs when suppressing encoding.SIGNIFICANCE STATEMENT Forgetting often seems like an unintended experience, but forgetting can be intentional, and can be accomplished with multiple strategies. These strategies, including encoding suppression and thought substitution, may rely on different neural mechanisms. Here, we test the hypothesis that encoding suppression engages domain-general prefrontally driven inhibitory control mechanisms, while thought substitution does not. Using cross-task analyses, we provide evidence that encoding suppression engages the same inhibitory mechanisms used for stopping motor actions, but these mechanisms are not engaged by thought substitution. These findings not only support the notion that mnemonic encoding processes can be directly inhibited, but also have broad relevance, as certain populations with disrupted inhibitory processing may be more successful accomplishing intentional forgetting through thought substitution strategies.

Brain connectivity alterations during sleep by closed-loop transcranial neurostimulation predict metamemory sensitivity.

Metamemory involves the ability to correctly judge the accuracy of our memories. The retrieval of memories can be improved using transcranial electrical stimulation (tES) during sleep, but evidence for improvements to metamemory sensitivity is limited. Applying tES can enhance sleep-dependent memory consolidation, which along with metamemory requires the coordination of activity across distributed neural systems, suggesting that examining functional connectivity is important for understanding these processes. Nevertheless, little research has examined how functional connectivity modulations relate to overnight changes in metamemory sensitivity. Here, we developed a closed-loop short-duration tES method, time-locked to up-states of ongoing slow-wave oscillations, to cue specific memory replays in humans. We measured electroencephalographic (EEG) coherence changes following stimulation pulses, and characterized network alterations with graph theoretic metrics. Using machine learning techniques, we show that pulsed tES elicited network changes in multiple frequency bands, including increased connectivity in the theta band and increased efficiency in the spindle band. Additionally, stimulation-induced changes in beta-band path length were predictive of overnight changes in metamemory sensitivity. These findings add new insights into the growing literature investigating increases in memory performance through brain stimulation during sleep, and highlight the importance of examining functional connectivity to explain its effects.

Separable neural mechanisms support intentional forgetting and thought substitution.

Psychological and neuroscientific experiments have established that people can intentionally forget information via different strategies: direct suppression and thought substitution. However, few studies have directly compared the effectiveness of these strategies in forgetting specific items, and it remains an open question if the neural mechanisms supporting these strategies differ. Here, we developed a novel item-method directed forgetting paradigm with Remember, Forget, and Imagine cues, and recorded EEG to directly compare these strategies. Behaviorally, Forget and Imagine cues produced similar forgetting compared to Remember cues, but through separable neural processes; Forget cues elicited frontal oscillatory power changes that were predictive of future forgetting, whereas item-cue representational similarity was predictive of later accuracy for Imagine cues. These results suggest that both strategies can lead to intentional forgetting, but directed forgetting may rely on frontally-mediated suppression, while thought substitution may lead to contextual shifting, impairing successful retrieval.

Dividing attention influences contextual facilitation and revision during language comprehension.

Although we often seem to successfully comprehend language in the face of distraction, few studies have examined the role of sustained attention in critical components of sentence processing, such as integrating information over a sentence and revising predictions when unexpected information is encountered. The current study investigated the impact of attention on sentence processing using a novel dual-task paradigm. Participants read weakly and strongly constraining sentences with expected or unexpected endings while also tracking the motion of dots in the background, and their EEG was recorded. Under full attention, the amplitude of the N400 component of the ERP, a measure of semantic access, was reduced (facilitated) in a graded fashion by contextual strength and fit. This context-based facilitation was attenuated when attention was divided, suggesting that sustained attention is important for building up message-level representations. In contrast, the post-N400 frontal positivity that has been observed to prediction violations and associated with revision processes was unaffected by dividing attention. However, under divided attention, participants also elicited posteriorly-distributed effects to these violations. Thus, predictive processes seem to be engaged even when attention is divided, but additional resources may then be required to process unexpected information.

Representational Pattern Similarity of Electrical Brain Activity Reveals Rapid and Specific Prediction during Language Comprehension.

Predicting upcoming events is a critical function of the brain, and language provides a fertile testing ground for studying prediction, as comprehenders use context to predict features of upcoming words. Many aspects of the mechanisms of prediction remain elusive, partly due to a lack of methodological tools to probe prediction formation in the moment. To elucidate what features are neurally preactivated and when, we used representational similarity analysis on previously collected sentence reading data. We compared EEG activity patterns elicited by expected and unexpected sentence final words to patterns from the preceding words of the sentence, in both strongly and weakly constraining sentences. Pattern similarity with the final word was increased in an early time window following the presentation of the pre-final word, and this increase was modulated by both expectancy and constraint. This was not seen at earlier words, suggesting that predictions were precisely timed. Additionally, pre-final word activity-the predicted representation-had negative similarity with later final word activity, but only for strongly expected words. These findings shed light on the mechanisms of prediction in the brain: rapid preactivation occurs following certain cues, but the predicted features may receive reduced processing upon confirmation.

Adult Age Differences in the Use of Conceptual Combination as an Associative Encoding Strategy.

It is well-established that aging impairs memory for associations more than it does memory for single items. Aging also impacts processes involved in online language comprehension, including the ability to form integrated, message-level representations. These changes in comprehension processes could impact older adults' associative memory performance, perhaps by reducing or altering the effectiveness of encoding strategies that encourage semantic integration. The present study examined age differences in the use of a strategy termed conceptual combination, which involves integrating two words (e.g., "winter" and "salad") into a single concept ("a salad for winter"). We recorded ERPs while participants studied unrelated noun pairs using a strategy that either did or did not encourage conceptual combination. We also varied the concreteness of the first noun in each pair in order to measure compositional concreteness effects, or ERP differences at the second noun due to the concreteness of the first noun. At the first nouns, older adults showed word-level concreteness effects that were similar to those of younger adults. However, compositional concreteness effects were diminished in older adults, consistent with reduced semantic integration. Older adults' associative memory performance was better for word pairs studied during the conceptual combination task versus the non-combinatory encoding task; however, the magnitude of the age-related associative memory deficit did not differ between tasks. Finally, analyses of both memory accuracy and trial-by-trial ratings of perceived combination success suggested that older adults had disproportionate difficulty applying the conceptual combination strategy to word pairs that began with abstract nouns. Overall, these results indicate that changes to integrative language processing that occur with age are not independent of - and may sometimes exacerbate - age-related memory decline.

Downstream Behavioral and Electrophysiological Consequences of Word Prediction on Recognition Memory.

When people process language, they can use context to predict upcoming information, influencing processing and comprehension as seen in both behavioral and neural measures. Although numerous studies have shown immediate facilitative effects of confirmed predictions, the downstream consequences of prediction have been less explored. In the current study, we examined those consequences by probing participants' recognition memory for words after they read sets of sentences. Participants read strongly and weakly constraining sentences with expected or unexpected endings ("I added my name to the list/basket"), and later were tested on their memory for the sentence endings while EEG was recorded. Critically, the memory test contained words that were predictable ("list") but were never read (participants saw "basket"). Behaviorally, participants showed successful discrimination between old and new items, but false alarmed to the expected-item lures more often than to new items, showing that predicted words or concepts can linger, even when predictions are disconfirmed. Although false alarm rates did not differ by constraint, event-related potentials (ERPs) differed between false alarms to strongly and weakly predictable words. Additionally, previously unexpected (compared to previously expected) endings that appeared on the memory test elicited larger N1 and LPC amplitudes, suggesting greater attention and episodic recollection. In contrast, highly predictable sentence endings that had been read elicited reduced LPC amplitudes during the memory test. Thus, prediction can facilitate processing in the moment, but can also lead to false memory and reduced recollection for predictable information.

One-Shot Tagging During Wake and Cueing During Sleep With Spatiotemporal Patterns of Transcranial Electrical Stimulation Can Boost Long-Term Metamemory of Individual Episodes in Humans.

Targeted memory reactivation (TMR) during slow-wave oscillations (SWOs) in sleep has been demonstrated with sensory cues to achieve about 5-12% improvement in post-nap memory performance on simple laboratory tasks. But prior work has not yet addressed the one-shot aspect of episodic memory acquisition, or dealt with the presence of interference from ambient environmental cues in real-world settings. Further, TMR with sensory cues may not be scalable to the multitude of experiences over one's lifetime. We designed a novel non-invasive non-sensory paradigm that tags one-shot experiences of minute-long naturalistic episodes in immersive virtual reality (VR) with unique spatiotemporal amplitude-modulated patterns (STAMPs) of transcranial electrical stimulation (tES). In particular, we demonstrated that these STAMPs can be re-applied as brief pulses during SWOs in sleep to achieve about 10-20% improvement in the metamemory of targeted episodes compared to the control episodes at 48 hours after initial viewing. We found that STAMPs can not only facilitate but also impair metamemory for the targeted episodes based on an interaction between pre-sleep metamemory and the number of STAMP applications during sleep. Overnight metamemory improvements were mediated by spectral power increases following the offset of STAMPs in the slow-spindle band (8-12 Hz) for left temporal areas in the scalp electroencephalography (EEG) during sleep. These results prescribe an optimal strategy to leverage STAMPs for boosting metamemory and suggest that real-world episodic memories can be modulated in a targeted manner even with coarser, non-invasive spatiotemporal stimulation.

Flexible conceptual combination: Electrophysiological correlates and consequences for associative memory.

When meaningful stimuli such as words are encountered in groups or pairs (e.g., "elephant-ferry"), they can be processed either separately or as an integrated concept ("an elephant ferry"). Prior research suggests that memory for integrated associations is supported by different mechanisms than is memory for nonintegrated associations. However, little is known about the neurocognitive mechanisms that support the integration of novel stimulus pairs. We recorded ERPs while participants memorized sequentially presented, unrelated noun pairs using a strategy that either did or did not involve attempting to construct coherent definitions. We varied the concreteness of the first noun in each pair to examine whether conceptual combination instructions would induce compositional concreteness effects, or differences in ERPs evoked by the second noun as a function of the concreteness of the first noun. We found that the conceptual combination task, but not the noncombinatory encoding task, produced compositional concreteness effects on a late frontal negativity previously linked to visual imagery. Moreover, word pairs studied under conceptual combination instructions showed evidence of more unitized or holistic memory representations on associative recognition and free recall tests. Finally, item analyses indicated that (a) items with higher normed imageability ratings were rated by participants as easier to conceptually combine, and (b) in the conceptual combination task, ease-of-combination ratings mediated an indirect relationship between imageability and subsequent associative memory. These data are suggestive of a role of compositional imagery in the online formation of novel concepts via conceptual combination.