Curiosity Satisfaction Increases Event-related Potentials Sensitive to Reward.

Successful learning depends on various factors such as depth of processing, motivation, or curiosity about information. A strong drive to learn something or the expectation of receiving a reward can be crucial to enhance learning. However, the influence of curiosity on the processing of new information and its similarity with reward processing is not well understood. This study examined whether states of curiosity influence specific ERPs associated with reward processing and whether these ERPs are related with later memory benefits. In an initial screening phase, participants indicated their curiosity and confidence in prior knowledge about answers to various trivia questions. In a subsequent study phase, we targeted different time windows related to reward processing during the presentation of trivia answers containing the reward positivity (RewP; 250-350 msec), the P3 (250-500 msec), and the late-positive-potential (LPP; 600-1000 msec). In a following surprise memory test, we found that participants recalled more high- than low-curiosity answers. The RewP, P3, and LPP showed greater positive mean amplitudes for high compared with low curiosity, reflecting increased reward processing. In addition, we found that the RewP and the P3 showed more positive mean amplitudes for later recalled compared with later forgotten answers, but curiosity did not modulate this encoding-related results. These findings support the view that the satisfaction of curiosity resembles reward processing, indicated by ERPs.

ORCA: A picture database of object-scene arrangements for cross-cultural and aging research.

In recent years, cross-cultural research on the modulation of basic cognitive processes by culture has intensified - also from an aging perspective. Despite this increased research interest, only a few cross-culturally normed non-verbal stimulus sets are available to support cross-cultural cognitive research in younger and older adults. Here we present the ORCA (Official Rating of Complex Arrangements) picture database, which includes a total of 720 object-scene compositions sorted into 180 quadruples (e.g., two different helmets placed in two different deserts). Each quadruple contains visually and semantically matched pairs of objects and pairs of scenes with varying degrees of semantic fit between objects and scenes. A total of 95 younger and older German and Chinese adults rated every object-scene pair on object familiarity and semantic fit between object and scene. While the ratings were significantly correlated between cultures and age groups, small but significant culture and age differences emerged. Object familiarity was higher for older adults than younger adults and for German participants than for Chinese participants. Semantic fit was rated lower by German older adults and Chinese younger adults as compared to German younger adults and Chinese older adults. Due to the large number of stimuli, our database is particularly well suited for cognitive and neuroscientific research on cross-cultural and age-related differences in perception, attention, and memory.

Electrophysiological evidence for context reinstatement effects on object recognition memory.

Reinstating the context present at encoding during the test phase generally enhances recognition memory compared with changing the context when specific item-context associations are established during encoding. However, it remains unclear whether context reinstatement improves the performance in differentiating between old and similar items in recognition memory tests and what underlying cognitive processes are involved. Using the context reinstatement paradigm together with event-related potentials (ERP), we examined the context-dependent effects of background scenes on recognition discrimination among similar objects. Participants were instructed to associate intentionally specific objects with background scenes during the encoding phase and subsequently complete an object recognition memory task, during which old and similar new objects were presented superimposed over the studied old or similar new background scenes. Electroencephalogram was recorded to measure the electrophysiological manifestations of cognitive processes associated with episodic retrieval. Behavioral results revealed enhanced performance in differentiating old from similar objects in the old context, as opposed to the similar context condition. Importantly, ERP results indicated a more pronounced recollection-related parietal object old/new effect in the old context compared to the similar context condition. This suggests that the ability to distinguish between old and similar objects in recognition memory is primarily driven by recollection rather than familiarity, particularly when the encoding context is reinstated during the test phase. Our findings are in line with the account that the impact of context reinstatement on object recognition memory is attributable to the enhanced recollection of specific item-context associations during retrieval and provides evidence for the specificity of episodic associative representations.

Effects of emotional study context on immediate and delayed recognition memory: Evidence from event-related potentials.

Whilst research has largely focused on the recognition of emotional items, emotion may be a more subtle part of our surroundings and conveyed by context rather than by items. Using ERPs, we investigated which effects an arousing context during encoding may have for item-context binding and subsequent familiarity-based and recollection-based item-memory. It has been suggested that arousal could facilitate item-context bindings and by this enhance the contribution of recollection to subsequent memory judgements. Alternatively, arousal could shift attention onto central features of a scene and by this foster unitisation during encoding. This could boost the contribution of familiarity to remembering. Participants learnt neutral objects paired with ecologically highly valid emotional faces whose names later served as neutral cues during an immediate and delayed test phase. Participants identified objects faster when they had originally been studied together with emotional context faces. Items with both neutral and emotional context elicited an early frontal ERP old/new difference (200-400 ms). Neither the neurophysiological correlate for familiarity nor recollection were specific to emotionality. For the ERP correlate of recollection, we found an interaction between stimulus type and day, suggesting that this measure decreased to a larger extend on Day 2 compared with Day 1. However, we did not find direct evidence for delayed forgetting of items encoded in emotional contexts at Day 2. Emotion at encoding might make retrieval of items with emotional context more readily accessible, but we found no significant evidence that emotional context either facilitated familiarity-based or recollection-based item-memory after a delay of 24 h.

Can the elderly take the action? - The influence of unitization induced by action relationships on the associative memory deficit.

Healthy aging is associated with intact familiarity, whereas recollection, usually supporting associative memory, is attenuated. Accordingly, associative memory shows a stronger age-related decline than item memory. One approach to alleviate age-related associative memory deficits is to increase the contribution of familiarity to associative memory by creating encoding conditions that allow to integrate separate stimuli to an entity (unitization). The current study investigated whether bottom-up unitization can reduce age-related differences in associative memory. Younger (YA) and older adults (OA) studied associations between semantically unrelated objects, spatially arranged in a way that an action between these two objects is possible (unitized, e.g., emptying a bottle into a sneaker) or not (non-unitized). At test, participants distinguished intact from recombined and new object pairs. As expected, we found larger age differences for associative memory than for item memory. Additionally, the presence of action relationships supports memory performance in both age groups. In the event-related potentials (ERP) of the test phase, we observed an age-related attenuation of recollection and preserved familiarity independent of the action relationship condition. Considering comparisons including the recombined pairs, the ERP correlate of associative familiarity (i.e., intact vs. recombined) was present in OA for action-related pairs, whereas for YA, there was no evidence for enhanced familiarity for action-related pairs. In the late time window, ERP evidence for recollection for intact action-related object pairs was obtained independent of age group. In conclusion, both age groups benefited from unitization by action relationships but by different mechanisms. While YA show no associative familiarity for action-related object pairs but a general reliance on recollection for associations in action-related and -unrelated pairs, OA seem to rely more on familiarity for the specific arrangement of action-related pairs.

The more you know: Schema-congruency supports associative encoding of novel compound words. Evidence from event-related potentials.

We aimed to investigate the neurocognitive mechanisms of event congruency with prior (schema) knowledge for the learning of novel compound words. Event-related potentials (ERPs) were recorded during an incidental learning task, in which novel noun-noun compounds were presented in a semantically congruent context, enabling schema-supported processing, or in a neutral context. As expected, associative memory performance was better for compounds preceded by a congruent context. Although the N400 was attenuated in the congruent condition, subsequent memory effects (SMEs) in the N400 time interval did not differ across conditions, suggesting that the processes reflected in the N400 cannot account for the memory advantage in the congruent condition. However, a parietal SME was obtained for compounds preceded by a congruent context, only, which we interpret as reflecting the schema-supported formation of a conceptual compound representation. A late frontal SME was obtained in both conditions, presumably reflecting the more general inter-item associative encoding of compound constituents.

Improving episodic memory: Frontal-midline theta neurofeedback training increases source memory performance.

Cognitive and neurofeedback training (NFT) studies have demonstrated that training-induced alterations of frontal-midline (FM) theta activity (4-8 Hz) transfer to cognitive control processes. Given that FM theta oscillations are assumed to provide top-down control for episodic memory retrieval, especially for source retrieval, that is, accurate recollection of contextual details of prior episodes, the present study investigated whether FM theta NFT transfers to memory control processes. It was assessed (1) whether FM theta NFT improves source retrieval and modulates its underlying EEG characteristics and (2) whether this transfer extends over two posttests. Over seven NFT sessions, the training group who trained individual FM theta activity showed greater FM theta increase than an active control group who trained randomly chosen frequency bands. The training group showed better source retrieval in a posttraining session performed 13 days after NFT and their performance increases from pre- to both posttraining sessions were predicted by NFT theta increases. Thus, training-induced enhancement of memory control processes seems to protect newly formed memories from proactive interference of previously learned information. EEG analyses revealed that during pretest both groups showed source memory specific theta activity at frontal and parietal sites. Surprisingly, training-induced improvements in source retrieval tended to be accompanied by less prestimulus FM theta activity, which was predicted by NFT theta change for the training but not the control group, suggesting a more efficient use of memory control processes after training. The present findings provide unique evidence for the enhancement of memory control processes by FM theta NFT.

Feedback timing modulates interactions between feedback processing and memory encoding: Evidence from event-related potentials.

Feedback-based learning relies on a procedural learning system driven by reward prediction errors (RPEs). The processing of temporally delayed feedback is supported by brain structures associated with declarative memory processes, but it is still unknown how delayed feedback processing and memory encoding interact. In this study, a subsequent memory paradigm was employed to investigate how the incidental encoding of feedback pictures presented with a short (SD, 500 ms) or long (LD, 6500 ms) delay in a probabilistic learning task affects the event-related potential (ERP) correlate of RPEs (i.e., the feedback-related negativity; FRN). In an ensuing test phase, a surprise recognition memory test for the feedback pictures was conducted. FRN amplitudes measured in the feedback-locked ERPs recorded during the learning phase (FRNpeak) and in the negative minus positive feedback difference wave (FRNdiff) were compared for subsequently remembered and forgotten feedback pictures. Feedback processing as reflected in the FRNpeak was diminished for remembered LD feedback pictures, indicating that delayed feedback processing and memory encoding competed for similar neural processing resources. As evidenced by large FRNdiff amplitudes in the SD condition, the evaluation of shortly delayed feedback strongly relied on the procedural learning system. A complementary model-based single trial analysis was conducted to validate models of the functional significance of the FRN. Consistent with previous studies, feedback-locked N170 and P300 amplitudes were sensitive to feedback delay. In the test phase, memory for LD feedback pictures was better than for SD pictures and accompanied by a late old-new effect, presumably reflecting extended recollective processing.

The effect of shared distinctiveness on source memory: An event-related potential study.

An illusory correlation (IC) is the erroneous perception that two actually uncorrelated categories are correlated. The Shared Distinctiveness Approach (SDA) explains ICs with heightened accessibility of distinctive category combinations in episodic memory. However, empirical evidence for this approach is heterogeneous. In the present event-related potential (ERP) study, we exploited the fact that more distinctive items elicit larger P300 responses than less distinctive items, which potentially predict subsequent memory performance differences for such items. Distinctiveness at encoding was created by presenting words that differed from frequently presented, positive words in valence, font color, or both. We hypothesized that shared distinctiveness (deviation in both color and valence) would lead to an enhanced P300 subsequent memory effect (SME), better source memory performance, and an overestimation of the frequency of shared distinctive items. Behavioral results indicated the presence of shared distinctiveness effects on source memory and frequency estimation. Unexpectedly, memory also was enhanced for positive items in the frequent color. This pattern also was reflected in the P300 for highly positive and negative items. However, shared distinctiveness did not modulate the P300 SME, indicating that the processing of distinctive features might only indirectly contribute to better encoding. This study shows that shared distinctiveness indeed is associated with better source memory and ICs. Because effects were observed for the most frequent and the least frequent category combination, our results imply that the processing of distinctiveness might involve attention allocation to diametrical category combinations, thereby accentuating the differences between the categories.

May the source be with you! Electrophysiological correlates of retrieval orientation are associated with source memory performance.

Successful source memory retrieval is assumed to rely on intact preretrieval processes, such as retrieval orientation (RO). RO is the specialized processing of retrieval cues, depending on the type of information, memory is searched for. In a previous study, a positive frontal slow wave RO ERP effect was interpreted as reflecting memory search for self-relevant information. However, such a functional interpretation is hampered by the use of retrieval strategies as a consequence of which target source information can be indirectly inferred from the correct classification of non-target source information. To overcome this limitation, the present study compared two types of source information (i.e. color or character information) by asking participants to remember details within each source type and thus enforcing the selective retrieval of target information. Consistent with previous research, a positive frontal ERP component (600-800 ms post-stimulus) differentiated between correct rejections in both tasks, probably reflecting memory search for self-relevant information. Moreover, the RO ERP effect was associated with better source memory performance, providing evidence for the beneficial effect of ROs for memory retrieval. This relationship might be covered in memory exclusion tasks due to non-target retrieval.

The neural mechanism of fluency-based memory illusions: the role of fluency context.

Recognition memory judgments can be influenced by a variety of signals including fluency. Here, we investigated whether the neural correlates of memory illusions (i.e., misattribution of fluency to prior study) can be modulated by fluency context. Using a masked priming/recognition memory paradigm, we found memory illusions for low confidence decisions. When fluency varied randomly across trials, we found reductions in perirhinal cortex (PrC) activity for primed trials, as well as a (pre)cuneus-PrC (BA 35) connectivity. When the fluency context was unchanging, there was increased PrC activity for primed trials, with the (pre)cuneus showing greater connectivity with PrC (BA 36). Thus, our results tentatively suggest two neural mechanisms via which fluency can lead to memory illusions.

Effects of acute psychosocial stress on the neural correlates of episodic encoding: Item versus associative memory.

Acute stress is known to modulate episodic memory, but little is known about the extent to, and the circumstances under, which stress affects encoding of item vs. inter-item associative information for words of different valences. Furthermore, the precise neuro-cognitive mechanisms underlying stress effects on episodic encoding in humans are largely unknown. To address these questions, in the present study we recorded EEG activity while male participants encoded neutral, negative and positive words, each paired with another word that was always neutral. Immediately before encoding, half of the participants experienced a psychosocial stressor, the Trier Social Stress Test, while the other half underwent a control procedure. Twenty-four hours later, participants completed separate item and associative recognition tests. Pre-learning stress enhanced item recognition accuracy for the positive, but not for the negative words. By contrast, there was no evidence for stress effects on associative recognition. The increase in item recognition was accompanied by a higher familiarity-, but not recollection-, based item retrieval of positive and neutral, but not negative words. Crucially, in the event-related potential (ERP) stress affected the amplitude of the frontal slow wave in general, and the frontal slow wave subsequent memory effect for positive words in specific, and the subsequent memory effect was correlated with cortisol levels after the stress manipulation. Our results suggest that positive words are encoded more elaboratively under stress, leading to a higher likelihood of subsequent item retrieval. An interaction of cortisol with frontal-lobe dependent control processes as well as a shift in attentional biases may contribute to this stress-induced modulation of episodic encoding.

Topographical differences of frontal-midline theta activity reflect functional differences in cognitive control abilities.

Electrophysiological oscillations are assumed to be the core mechanism for large-scale network communication. The specific role of frontal-midline theta oscillations as cognitive control mechanism is under debate. According to the dual mechanisms of control framework, cognitive control processes can be divided into proactive and reactive control. The present study aimed at investigating the role of frontal-midline theta activity by assessing oscillations in two tasks varying in the type of cognitive control needed. More specifically, a delayed match to sample (DMTS) task requiring proactive control and a color Stroop task recruiting reactive control processes were conducted within the same group of participants. Moreover, both tasks contained conditions with low and high need for cognitive control. As expected larger frontal-midline theta activity was found in conditions with high need for cognitive control. However, theta activity was focally activated at frontal sites in the DMTS task whereas it had a broader topographical distribution in the Stroop task, indicating that both proactive and reactive control are reflected in frontal-midline theta activity but reactive control is additionally characterized by a broader theta activation. These findings support the conclusion that frontal-midline theta acts functionally different depending on task requirements.

Illusory correlations despite equated category frequencies: A test of the information loss account.

Illusory correlations (IC) are the perception of covariation, where none exists. For example, people associate majorities with frequent behavior and minorities with infrequent behavior even in the absence of such an association. According to the information loss account, ICs result from greater fading of infrequent group-behavior combinations in memory. We conducted computer simulations based on this account which showed that ICs are expected under standard conditions with skewed category frequencies (i.e. 2:1 ratio for positive and negative descriptions), but not under conditions with equated category frequencies (i.e. 1:1 ratio for positive and negative descriptions). Contrary to these simulations, our behavioral experiments revealed an IC under both conditions, which did not decrease over time. Thus, information loss alone is not sufficient as an explanation for the formation of ICs. These results imply that negative items contribute to ICs not only due to their infrequency, but also due to their emotional salience.

Separating Event-related Potential Effects for Conceptual Fluency and Episodic Familiarity.

ERP old/new effects have been associated with different subprocesses of episodic recognition memory. The notion that recollection is reflected in the left parietal old/new effect seems to be uncontested. However, an association between episodic familiarity and the mid-frontal old/new effect is not uncontroversial. It has been argued that the mid-frontal old/new effect is functionally equivalent to the N400 and hence merely reflects differences in conceptual fluency between old and new items. Therefore, it is related to episodic familiarity only in situations in which conceptual fluency covaries with familiarity. Alternatively, the old/new effect in this time window reflects an interaction of episodic familiarity and conceptual processing with each making a unique functional contribution. To test this latter account, we manipulated conceptual fluency and episodic familiarity orthogonally in an incidental recognition test: Visually presented old and new words were preceded by either conceptually related or unrelated auditory prime words. If the mid-frontal old/new effect is functionally distinguishable from conceptual priming effects, an ERP contrast reflecting pure priming (correct rejections in the related vs. unrelated condition) and a contrast reflecting priming plus familiarity (hits in the related vs. correct rejections in the unrelated condition) should differ in scalp distribution. As predicted, the pure priming contrast had a right-parietal distribution, as typically observed for the N400 effect, whereas the priming plus familiarity contrast was significantly more frontally accentuated. These findings implicate that old/new effects in this time window are driven by unique functional contributions of episodic familiarity and conceptual processing.

Remembering specific features of emotional events across time: The role of REM sleep and prefrontal theta oscillations.

When an episode of emotional significance is encountered, it often results in the formation of a highly resistant memory representation that is easily retrieved for many succeeding years. Recent research shows that beyond generic consolidation processes, rapid eye movement (REM) sleep importantly contributes to this effect. However, the boundary conditions of consolidation processes during REM sleep, specifically whether these extend to source memory, have not been examined extensively. The current study tested the effects of putative consolidation processes emerging during REM sleep and slow wave sleep (SWS) on item and source memory of negative and neutral images, respectively. Results demonstrate superior emotional relative to neutral item memory retention after both late night REM sleep and early night SWS. Emotional source memory, on the other hand, exhibited an attenuated decline following late night REM sleep, whereas neutral source memory was selectively preserved across early night SWS. This pattern of results suggests a selective preservation of emotional source memory during REM sleep that is functionally dissociable from SWS-dependent reprocessing of neutral source memory. This was further substantiated by a neurophysiological dissociation: Postsleep emotional source memory was selectively correlated with frontal theta lateralization (REM sleep), whereas postsleep neutral item memory was correlated with SWS spindle power. As such, the present results contribute to a more comprehensive characterization of sleep-related consolidation mechanisms underlying emotional and neutral memory retention. Subsidiary analysis of emotional reactivity to previously encoded material revealed an enhancing rather than attenuating effect of late night REM sleep on emotional responses.

How does testing affect retrieval-related processes? An event-related potential (ERP) study on the short-term effects of repeated retrieval.

The testing effect is conceptualized as the benefit for remembering items that were studied and tested rather than just studied. Thus far, little is known about the underlying neurocognitive mechanisms. In an event-related potential (ERP) study, we investigated the immediate consequences of testing on recollection processes. During an initial study phase, participants encountered object names together with pictures of the denoted objects ("perceived items") or with the instruction to mentally visualize them ("imagined items"). Directly afterward, they had to differentiate between perceived, imagined, and new items, in two consecutive source memory tests. Half of the studied items were presented in the first run, and all items in the second. Behaviorally, repeated testing led to improved item and source memory, as well as faster reaction times, relative to items that had been tested once. In accordance with these behavioral changes, the left parietal old-new effect (500-900 ms) as the putative correlate of recollection was strongly enhanced by previous testing. An enhancement after testing was also observed for the early portion of the late right frontal old-new effect (700-900 ms). In contrast, old-new effects after 900 ms were not modulated by previous testing. The finding of a stronger left parietal old-new effect for previously tested items suggests that testing leads to an elaboration of memory traces, whereas the faster reaction times are more likely explained in terms of transfer-appropriate processing. The combination of more elaborated memory traces and transfer-appropriate processing provides a tentative explanation for the effectiveness of testing in enhancing retrieval performance.

Nap sleep preserves associative but not item memory performance.

Many studies have shown that sleep improves memory performance, and that even short naps during the day are beneficial. Certain physiological components of sleep such as spindles and slow-wave-sleep are thought to be particularly important for memory consolidation. The aim of this experiment was to reveal the role of naps for hippocampus-dependent associative memory (AM) and hippocampus-independent item memory (IM) alongside their corresponding ERP old/new effects. Participants learnt single words and word-pairs before performing an IM- and an AM-test (baseline). One group was subsequently allowed to nap (∼90min) while the other watched DVDs (control group). Afterwards, both groups performed a final IM- and AM-test for the learned stimuli (posttest). IM performance decreased for both groups, while AM performance decreased for the control group but remained constant for the nap group, consistent with predictions concerning the selective impact of napping on hippocampus-dependent recognition. Putative ERP correlates of familiarity and recollection were observed in the IM posttest, whereas only the later recollection-related effect was present in the AM test. Notably, none of these effects varied with group. Positive correlations were observed between spindle density during slow-wave-sleep and AM posttest performance as well as between spindle density during non-REM sleep and AM baseline performance, showing that successful learning and retrieval both before and after sleep relates to spindle density during nap sleep. Together, these results speak for a selective beneficial impact of naps on hippocampus-dependent memories.

Modifications of recognition memory processes in preterm children: an event-related potential study.

Prematurity may cause hippocampal compromise. Therefore, hippocampus-dependent memory processes (recollection-based retrieval) may be more impaired than hippocampus-independent processes (familiarity-based retrieval). The memory of 18 children born preterm with reduced hippocampal volumes, without neonatal complications (weeks of gestation < 34, weight < 1,600 g), and 15 controls (8-10 years) was tested using an item recognition task. While groups were equal in memory performance, dissociation was found: The event-related potential (ERP) correlate of familiarity was intact in the preterm group, whereas the correlate of recollection was attenuated. A follow-up experiment ruled out that this was due to general cognitive deficits. Furthermore, gestational age correlated with the ERP index of recollection. Thus, recognition memory in preterm children may be characterized by a compensation of attenuated recollection by familiarity.

Fine neural tuning for orthographic properties of words emerges early in children reading alphabetic script.

The left-lateralized N170 component of ERPs for words compared with various control stimuli is considered as an electrophysiological manifestation of visual expertise for written words. To understand the information sensitivity of the effect, researchers distinguish between coarse tuning for words (the N170 amplitude difference between words and symbol strings) and fine tuning for words (the N170 amplitude difference between words and consonant strings). Earlier developmental ERP studies demonstrated that the coarse tuning for words occurred early in children (8 years old), whereas the fine tuning for words emerged much later (10 years old). Given that there are large individual differences in reading ability in young children, these tuning effects may emerge earlier than expected in some children. This study measured N170 responses to words and control stimuli in a large group of 7-year-olds that varied widely in reading ability. In both low and high reading ability groups, we observed the coarse neural tuning for words. More interestingly, we found that a stronger N170 for words than consonant strings emerged in children with high but not low reading ability. Our study demonstrates for the first time that fine neural tuning for orthographic properties of words can be observed in young children with high reading ability, suggesting that the emergent age of this effect is much earlier than previously assumed. The modulation of this effect by reading ability suggests that fine tuning is flexible and highly related to experience. Moreover, we found a correlation between this tuning effect at left occipitotemporal electrodes and children's reading ability, suggesting that the fine tuning might be a biomarker of reading skills at the very beginning of learning to read.