Judgments of Learning Reactively Improve Memory by Enhancing Learning Engagement and Inducing Elaborative Processing: Evidence from an EEG Study.

Making judgments of learning (JOLs) can reactively alter memory itself, a phenomenon termed the reactivity effect. The current study recorded electroencephalography (EEG) signals during the encoding phase of a word list learning task to explore the neurocognitive features associated with JOL reactivity. The behavioral results show that making JOLs reactively enhances recognition performance. The EEG results reveal that, compared with not making JOLs, making JOLs increases P200 and LPC amplitudes and decreases alpha and beta power. Additionally, the signals of event-related potentials (ERPs) and event-related desynchronizations (ERDs) partially mediate the reactivity effect. These findings support the enhanced learning engagement theory and the elaborative processing explanation to account for the JOL reactivity effect.

Anodal tDCS of the left inferior parietal cortex enhances memory for correct information without affecting recall of misinformation.

False memories during testimony are an enormous challenge for criminal trials. Exposure to post-event misinformation can lead to inadvertent creation of false memories, known as the misinformation effect. We investigated anodal transcranial direct current stimulation (tDCS) on the left inferior parietal lobe (IPL) during recall testing to enhance accurate recall while addressing the misinformation effect. Participants (N = 60) watched a television series depicting a fictional terrorist attack, then received an audio recording with misinformation, consistent information, and control information. During cued recall testing, participants received anodal or sham tDCS. Results revealed a robust misinformation effect in both groups, with participants falsely recalling on average 26.6% of the misinformed items. Bayesian statistics indicated substantial evidence in favour of the null hypothesis that there was no difference between groups in the misinformation effect. Regarding correct recall however, the anodal group exhibited significantly improved recall for items from the original video. Together, these results demonstrate that anodal tDCS of the left IPL enhances correct recall of the episodes from the original event without affecting false recall of misinformation. The findings support the IPL's role in recollection and source attribution of episodic memories.

The relation between learning and stimulus-response binding.

Perception and action rely on integrating or binding different features of stimuli and responses. Such bindings are short-lived, but they can be retrieved for a limited amount of time if any of their features is reactivated. This is particularly true for stimulus-response bindings, allowing for flexible recycling of previous action plans. A relation to learning of stimulus-response associations suggests itself, and previous accounts have proposed binding as an initial step of forging associations in long-term memory. The evidence for this claim is surprisingly mixed, however. Here we propose a framework that explains previous failures to detect meaningful relations of binding and learning by highlighting the joint contribution of three variables: (a) decay, (b) the number of repetitions, and (c) the time elapsing between repetitions. Accounting for the interplay of these variables provides a promising blueprint for innovative experimental designs that bridge the gap between immediate bindings on the one hand and lasting associations in memory on the other hand. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Prestimulus alpha power signals attention to retrieval.

The human brain is in distinct processing modes at different times. Specifically, a distinction can be made between encoding and retrieval modes, which refer to the brain's state when it is storing new information or searching for old information, respectively. Recent research proposed the idea of a "ready-to-encode" mode, which describes a prestimulus effect in brain activity that signals (external) attention to encoding and predicts subsequent memory performance. Whether there is also a corresponding "ready-to-retrieve" mode in human brain activity is currently unclear. In this study, we examined whether prestimulus oscillations can be linked to (internal) attention to retrieval. We show that task cues to prepare for retrieval (or testing) in comparison with restudy of previously studied vocabulary word pairs led to a significant decrease of prestimulus alpha power just before the onset of word stimuli. Beamformer analysis localized this effect in the right secondary visual cortex (Brodmann area 18). Correlation analysis showed that the task cue-induced, prestimulus alpha power effect is positively related to stimulus-induced alpha/beta power, which in turn predicted participants' memory performance. The results are consistent with the idea that prestimulus alpha power signals internal attention to retrieval, which promotes the elaborative processing of episodic memories. Future research on brain-computer interfaces may find the findings interesting regarding the potential of using online measures of fluctuating alpha oscillations to trigger the presentation and sequencing of restudy and testing trials, ultimately enhancing instructional learning strategies.

The oscillatory fingerprints of self-prioritization: Novel markers in spectral EEG for self-relevant processing.

Self-prioritization is a very influential modulator of human information processing. Still, little is known about the time-frequency dynamics of the self-prioritization network. In this EEG study, we used the familiarity-confound free matching task to investigate the spectral dynamics of self-prioritization and their underlying cognitive functions in a drift-diffusion model. Participants (N = 40) repeatedly associated arbitrary geometric shapes with either "the self" or "a stranger." Behavioral results demonstrated prominent self-prioritization effects (SPEs) in reaction time and accuracy. Remarkably, EEG cluster analysis also revealed two significant SPEs, one in delta/theta power (2-7 Hz) and one in beta power (19-29 Hz). Drift-diffusion modeling indicated that beta activity was associated with evidence accumulation, whereas delta/theta activity was associated with response selection. The decreased beta suppression of the SPE might indicate more efficient sensorimotor processing of self-associated stimulus-response features, whereas the increased delta/theta SPE might refer to the facilitated retrieval of self-relevant features across a widely distributed associative self-network. These novel oscillatory biomarkers of self-prioritization indicate their function as an associative glue for the self-concept.

Perception-Action Integration Is Altered in Functional Movement Disorders.

BACKGROUND: Although functional neurological movement disorders (FMD) are characterized by motor symptoms, sensory processing has also been shown to be disturbed. However, how the integration of perception and motor processes, essential for the control of goal-directed behavior, is altered in patients with FMD is less clear. A detailed investigation of these processes is crucial to foster a better understanding of the pathophysiology of FMD and can systematically be achieved in the framework of the theory of event coding (TEC). OBJECTIVE: The aim was to investigate perception-action integration processes on a behavioral and neurophysiological level in patients with FMD. METHODS: A total of 21 patients and 21 controls were investigated with a TEC-related task, including concomitant electroencephalogram (EEG) recording. We focused on EEG correlates established to reflect perception-action integration processes. Temporal decomposition allowed to distinguish between EEG codes reflecting sensory (S-cluster), motor (R-cluster), and integrated sensory-motor processing (C-cluster). We also applied source localization analyses. RESULTS: Behaviorally, patients revealed stronger binding between perception and action, as evidenced by difficulties in reconfiguring previously established stimulus-response associations. Such hyperbinding was paralleled by a modulation of neuronal activity clusters, including reduced C-cluster modulations of the inferior parietal cortex and altered R-cluster modulations in the inferior frontal gyrus. Correlations of these modulations with symptom severity were also evident. CONCLUSIONS: Our study shows that FMD is characterized by altered integration of sensory information with motor processes. Relations between clinical severity and both behavioral performance and neurophysiological abnormalities indicate that perception-action integration processes are central and a promising concept for the understanding of FMD. © 2023 The Authors. Movement Disorders published by Wiley Periodicals LLC on behalf of International Parkinson and Movement Disorder Society.

The Forward Testing Effect Is Resistant to Acute Psychosocial Retrieval Stress.

The forward testing effect refers to the finding that testing of previously studied information improves memory for subsequently studied newer information. Recent research showed that the effect is immune to acute psychosocial encoding/retrieval stress, i.e., stress that is induced before initial encoding. The present study investigated whether the forward testing effect is also robust to acute psychosocial retrieval stress, i.e., stress that is induced after encoding but before retrieval of the critical item list. Participants (N = 128) studied three lists of words in anticipation of a final cumulative recall test. Participants were tested immediately on Lists 1 and 2 (testing condition) or restudied the two lists after initial study (restudy condition). After study of the critical List 3, psychosocial stress was induced in half of the participants (stress group), whereas no stress was induced in the other half (control group). The Trier Social Stress Test for Groups (TSST-G) was used for stress induction. Salivary cortisol, alpha amylase, and subjective stress were repeatedly measured. The results of the criterion test showed a generally detrimental effect of psychosocial retrieval stress on List 3 recall. Importantly, the forward testing effect was unaffected by retrieval stress. The findings are discussed with respect to current theories of the forward testing effect.

Posterior delta/theta EEG activity as an early signal of Stroop conflict detection.

The conflict monitoring theory postulates that conflict detection is initiated in the anterior cingulate cortex (ACC), indexed by midfrontal theta oscillations in the electroencephalogram (EEG). Recent research suggested that distractor detection (in the Eriksen flanker task) can be initiated relatively early by attentional control processes in the occipital lobe. Whether attentional control is also involved in the detection of stimulus-response overlapping conflict in the Stroop task is yet unclear. In the present study, we analyzed EEG time-frequency data (N = 47) to investigate the contribution of early attentional control processes to the detection of response conflict and semantic conflict in a lateralized version of the color-word Stroop task. The behavioral results showed significant conflict effects in response times (RT). The EEG results showed a prominent midfrontal response conflict effect in total theta power (4-8 Hz). Importantly, detection of response conflict and semantic conflict was observed in posterior delta/theta power (2-8 Hz), which was lateralized depending on the presentation side of the irrelevant Stroop words. In explorative regression analysis, both the midfrontal and the posterior response conflict effects predicted the size of response conflict errors. These results suggest that attentional control processes in posterior areas contribute to the initiation of response-conflict detection in the Stroop task. The findings are consistent with the idea of a representational link between stimulus and response features, known as the common coding principle.

Stimulus decay functions in action control.

When facing particular combinations of stimuli and responses, people create temporary event-files integrating the corresponding stimulus and response features. Subsequent repetition of one or more of these features retrieves the entire event-file, which impairs performance if not all features are repeated (partial-repetition costs). In the literature, different decay functions have been reported presumably dependent on the type of feature that is repeated (e.g. target vs. distractor features). Here, we use a variant of the S1R1-S2R2 and distractor-response binding task and analyze for the first time target-based and distractor-based event-file decay functions within the same task and sample. While we found evidence for decay functions and also stronger retrieval due to target than distractor repetitions, slopes of the decay functions were comparable suggesting that the decay process itself is equal irrespective of the type of stimulus feature that is repeated. Our study thereby confirms overarching approaches that summarize paradigm specific findings with the same set of core processes.

Retrieval Practice Enhances New Learning but does Not Affect Performance in Subsequent Arithmetic Tasks.

The forward testing effect is an indirect benefit of retrieval practice. It refers to the finding that retrieval practice of previously studied information enhances learning and retention of subsequently studied other information in episodic memory tasks. Here, two experiments were conducted that investigated whether retrieval practice influences participants' performance in other tasks, i.e., arithmetic tasks. Participants studied three lists of words in anticipation of a final recall test. In the testing condition, participants were immediately tested on lists 1 and 2 after study of each list, whereas in the restudy condition, they restudied lists 1 and 2 after initial study. Before and after study of list 3, participants did an arithmetic task. Finally, participants were tested on list 3, list 2, and list 1. Different arithmetic tasks were used in the two experiments. Participants did a modular arithmetic task in Experiment 1a and a single-digit multiplication task in Experiment 1b. The results of both experiments showed a forward testing effect with interim testing of lists 1 and 2 enhancing list 3 recall in the list 3 recall test, but no effects of recall testing of lists 1 and 2 for participants' performance in the arithmetic tasks. The findings are discussed with respect to cognitive load theory and current theories of the forward testing effect.

Can People Intentionally and Selectively Forget Prose Material?

List-method directed forgetting (LMDF) is the demonstration that people can intentionally forget previously studied information when they are asked to forget what they have previously learned and remember new information instead. In addition, recent research demonstrated that people can selectively forget when cued to forget only a subset of the previously studied information. Both forms of forgetting are typically observed in recall tests, in which the to-be-forgotten and to-be-remembered information is tested independent of original cuing. Thereby, both LMDF and selective directed forgetting (SDF) have been studied mostly with unrelated item materials (e.g., word lists). The present study examined whether LMDF and SDF generalize to prose material. Participants learned three prose passages, which they were cued to remember or forget after the study of each passage. At the time of testing, participants were asked to recall the three prose passages regardless of original cuing. The results showed no significant differences in recall of the three lists as a function of cuing condition. The findings suggest that LMDF and SDF do not occur with prose material. Future research is needed to replicate and extend these findings with (other) complex and meaningful materials before drawing firm conclusions. If the null effect proves to be robust, this would have implications regarding the ecological validity and generalizability of current LMDF and SDF findings.

Watching the Brain as It (Un)Binds: Beta Synchronization Relates to Distractor-Response Binding.

Human action control relies on event files, that is, short-term stimulus-response bindings that result from the integration of perception and action. The present EEG study examined oscillatory brain activities related to the integration and disintegration of event files in the distractor-response binding (DRB) task, which relies on a sequential prime-probe structure with orthogonal variation of distractor and response relations between prime and probe. Behavioral results indicated a DRB effect in RTs, which was moderated by the duration of the response-stimulus interval (RSI) between prime response and probe stimulus onset. Indeed, a DRB effect was observed for a short RSI of 500 msec but not for a longer RSI of 2000 msec, indicating disintegration of event files over time. EEG results revealed a positive correlation between individual DRB in the RSI-2000 condition and postmovement beta synchronization after both prime and probe responses. Beamformer analysis localized this correlation effect to the middle occipital gyrus, which also showed highest coherency with precentral and inferior parietal brain regions. Together, these findings suggest that postmovement beta synchronization is a marker of event file disintegration, with the left middle occipital gyrus being a hub region for stimulus-response bindings in the visual DRB task.

Oscillatory Correlates of Selective Restudy.

Prior behavioral work has shown that selective restudy of some studied items leaves recall of the other studied items unaffected when lag between study and restudy is short, but improves recall of the other items when lag is prolonged. The beneficial effect has been attributed to context retrieval, assuming that selective restudy reactivates the context at study and thus provides a retrieval cue for the other items (Bäuml, 2019). Here the results of two experiments are reported, in each of which subjects studied a list of items and then, after a short 2-min or a prolonged 10-min lag, restudied some of the list items. Participants' electroencephalography (EEG) was recorded during both the study and restudy phases. In Experiment 2, but not in Experiment 1, subjects engaged in a mental context reinstatement task immediately before the restudy phase started, trying to mentally reinstate the study context. Results of Experiment 1 revealed a theta/alpha power increase from study to restudy after short lag and an alpha/beta power decrease after long lag. Engagement in the mental context reinstatement task in Experiment 2 eliminated the decrease in alpha/beta power. The results are consistent with the view that the observed alpha/beta decrease reflects context retrieval, which became obsolete when there was preceding mental context reinstatement.

Electrophysiological correlates of saving-enhanced memory: Exploring similarities to list-method directed forgetting.

People regularly outsource parts of their memory onto external memory stores like computers or smartphones. Such cognitive offloading can enhance subsequent memory performance, as referred to the saving-enhanced memory effect (Storm & Stone, 2015). The cognitive mechanisms of this effect are not clear to date, however similarities to list-method directed forgetting (LMDF) have been stated. Here, we examined in 52 participants the electrophysiological (EEG) correlates of the saving-enhanced memory effect and compared our results to earlier LMDF findings (Hanslmayr et al., 2012). For this purpose, EEG alpha power and alpha phase synchrony during the encoding of two word lists were compared as a function of saving or no-saving. We hypothesised that if saving-enhanced memory was related to LMDF, saving in comparison to no-saving between lists should reduce alpha power and alpha phase synchrony during List 2 encoding, two effects that have been related to List 2 encoding benefits and List 1 inhibition in the earlier LMDF work. The results showed no statistically significant saving-enhanced memory effect and no significant effects in EEG alpha power or alpha phase synchrony. Possible explanations for and implications of these non-significant findings are discussed.

Transcranial direct current stimulation over the left anterior temporal lobe during memory retrieval differentially affects true and false recognition in the DRM task.

Transcranial direct current stimulation (tDCS) is a form of non-invasive brain stimulation that has been used to modulate human brain activity and cognition. One area which has not yet been extensively explored using tDCS is the generation of false memories. In this study, we combined the Deese-Roediger-McDermott (DRM) task with stimulation of the left anterior temporal lobe (ATL) during retrieval. This area has been shown to be involved in semantic processing in general and retrieval of false memories in the DRM paradigm in particular. During stimulation, 0.7 mA were applied via a 9 cm² electrode over the left ATL, with the 35 cm² return electrode placed over the left deltoid. We contrasted the effects of cathodal, anodal, and sham stimulation, which were applied in the recognition phase of the experiment on a sample of 78 volunteers. Results showed impaired recognition of true memories after both anodal and cathodal stimulation in comparison to sham stimulation, suggesting a reduced signal-to-noise ratio. In addition, the results revealed enhanced false recognition of concept lure items during cathodal stimulation compared to anodal stimulation, indicating a polarity-dependent impact of tDCS on false memories in the DRM task. The pathway by which tDCS modulated false recognition remains unclear: stimulation may have changed the activation of irrelevant lures or affected the weighting and monitoring of lure activations. Nevertheless, these results are a first step towards using brain stimulation to decrease false memories. Practical implications of the findings for real-life settings, for example, in the courtroom, need to be addressed in future work.

Target Amplification and Distractor Inhibition: Theta Oscillatory Dynamics of Selective Attention in a Flanker Task.

Selective attention is a key mechanism to monitor conflict-related processing and behaviour, by amplifying task-relevant processing and inhibiting task-irrelevant information. Conflict monitoring and resolution is typically associated with brain oscillatory power increase in the theta frequency range (3-8 Hz), as indexed by increased midfrontal theta power. We expand previous findings of theta power increase related to conflict processing and distractor inhibition by considering attentional target amplification to be represented in theta frequency as well. The present study (N = 41) examined EEG oscillatory activities associated with stimulus and response conflict in a lateralized flanker task. Depending on the perceptual (in)congruency and response (in)compatibility of distractor-target associations, resulting stimulus and response conflicts were examined in behavioural and electrophysiological data analyses. Both response and stimulus conflict emerged in RT analysis. Regarding EEG data, response-locked cluster analysis showed an increase of midfrontal theta power related to response conflict. In addition, stimulus-locked cluster analysis revealed early clusters with increased parietal theta power for nonconflicting compared to conflicting trials, followed by increased midfrontal theta power for both stimulus and response conflict. Our results suggest that conflict resolution in the flanker task relies on a combination of target amplification, depicted by parietal theta power increase, and distractor inhibition, indexed by midfrontal theta power increase, for both stimulus and response conflicts. Attentional amplification of sensory target features is discussed with regard to a domain-general conflict monitoring account.

Failure to modulate reward prediction errors in declarative learning with theta (6 Hz) frequency transcranial alternating current stimulation.

Recent evidence suggests that reward prediction errors (RPEs) play an important role in declarative learning, but its neurophysiological mechanism remains unclear. Here, we tested the hypothesis that RPEs modulate declarative learning via theta-frequency oscillations, which have been related to memory encoding in prior work. For that purpose, we examined the interaction between RPE and transcranial Alternating Current Stimulation (tACS) in declarative learning. Using a between-subject (real versus sham stimulation group), single-blind stimulation design, 76 participants learned 60 Dutch-Swahili word pairs, while theta-frequency (6 Hz) tACS was administered over the medial frontal cortex (MFC). Previous studies have implicated MFC in memory encoding. We replicated our previous finding of signed RPEs (SRPEs) boosting declarative learning; with larger and more positive RPEs enhancing memory performance. However, tACS failed to modulate the SRPE effect in declarative learning and did not affect memory performance. Bayesian statistics supported evidence for an absence of effect. Our study confirms a role of RPE in declarative learning, but also calls for standardized procedures in transcranial electrical stimulation.

Does Amount of Pre-cue Encoding Modulate Selective List Method Directed Forgetting?

Prior work reported evidence that when people are presented with both a relatively short list of relevant information and a relatively short list of irrelevant information, a subsequent cue to forget the irrelevant list can induce successful selective directed forgetting of the irrelevant list without any forgetting of the relevant list. The goal of the present study is to determine whether this selectivity effect is restricted to short lists of information (six items per list), or if the effect generalizes to longer lists (12 items per list). In Experiment 1, we replicate the finding that selective directed forgetting can occur when short lists of relevant and irrelevant information are involved. Going beyond this replication, we show in Experiment 2 that such selectivity can arise both when shorter and when relatively long lists of items are used. The results are consistent with the view that selective directed forgetting can result from the action of a flexible inhibitory mechanism. They are less well in line with the view that selective cues to forget pre-cue information induce a change in participants' mental context.

The Forward Testing Effect is Immune to Acute Psychosocial Encoding/Retrieval Stress.

The forward testing effect (FTE) refers to the finding that testing of previously studied information enhances memory for subsequently studied other information. Previous research demonstrated that the FTE is a robust phenomenon that generalizes across different materials and populations. The present study examined whether the FTE is robust under acute psychosocial encoding/retrieval stress. In each of two experimental conditions, participants studied three item lists in anticipation of final cumulative recall testing. In the testing condition, participants were tested immediately on lists 1 and 2, whereas in the restudy condition, they restudied lists 1 and 2. In both conditions, participants were tested immediately on list 3. Acute psychosocial stress was induced in participants prior to the encoding of item lists using the Trier social stress test for groups protocol. No stress was induced in a control group. Salivary cortisol, alpha amylase, and subjective stress were measured repeatedly to capture the biopsychological stress response. The results showed a significant FTE on list 3 recall, that is, testing of lists 1 and 2 enhanced the recall of list 3. No significant effect of stress on the FTE was observed, suggesting that the FTE is robust under acute psychosocial encoding/retrieval stress. The discussion provides suggestions for future research directions.

EEG beta power increase indicates inhibition in motor memory.

Beta power increase has been suggested to be an electrophysiological marker of response inhibition during voluntary action stopping. We examined whether beta power increase accompanies inhibition in human motor memory, focusing the phenomenon of retrieval-induced forgetting that has been assumed to be the consequence of inhibition in memory. Whereas most studies on this effect comprise word materials, a variant of the retrieval-induced-forgetting paradigm exists for examining motor memory. In the present study, we recorded scalp EEG during this motor-specific variant and examined EEG oscillatory correlates of retrieval-induced slowing of motor actions. Here, forgetting occurred in the form of significant slowing when executing motor sequences in a final recall test. Participants first learned to associate memory cues to sequential finger movements, half of which were performed with the left hand and half with the right hand. They then selectively retrieval-practiced half of the items of one hand before finally memory for all items was tested. On a behavioral level, selective retrieval practice induced slower execution of the non-retrieved items of the retrieval-practiced hand in a final memory test. On a physiological level, this retrieval-induced-slowing effect was predicted by an increase of EEG beta power during retrieval practice. The results suggest that motor retrieval-induced forgetting is a consequence of inhibition in human memory.