The differential effects of emotional salience on direct associative and relational memory during a nap

Relational memories are formed from shared components between directly learned memory associations, flexibly linking learned information to better inform future judgments. Sleep has been found to facilitate both direct associative and relational memories. However, the impact of incorporating emotionally salient information into learned material and the interaction of emotional salience and sleep in facilitating both types of memory is unknown. Participants encoded two sets of picture pairs, with either emotionally negative or neutral objects paired with neutral faces. The same objects were present in both sets, paired with two different faces across the sets. Baseline memory for these directly paired associates was tested immediately after encoding, followed by either a 90-min nap opportunity or wakefulness. Five hours after learning, a surprise test assessed relational memory, the indirect association between two faces paired with the same object during encoding, followed by a retest of direct associative memory. Overall, negative information was remembered better than neutral for directly learned pairs. A nap facilitated both preservation of direct associative memories and formation of relational memories, compared to remaining awake. Interestingly, however, this sleep benefit was observed specifically for neutral directly paired associates, while both neutral and negative relational associations benefitted from a nap. Finally, REM sleep played opposing roles in neutral direct and relational associative memory formation, with more REM sleep leading to forgetting of direct associations but promoting relational associations, suggesting that, while not benefitting memory consolidation for directly learned details, REM sleep may foster the memory reorganization needed for relational memory.     

Residual effects of emotion are reflected in enhanced visual activity after sleep

Research has investigated how sleep affects emotional memory and how emotion enhances visual processing, but these questions are typically asked by re-presenting an emotional stimulus at retrieval. For the first time, we investigate whether sleep affects neural activity during retrieval when the memory cue is a neutral context that was previously presented with either emotional or nonemotional content during encoding. Participants encoded scenes composed of a negative or neutral object on a neutral background either in the morning (preceding 12 hours awake; wake group) or evening (preceding 12 hours including a night of sleep; sleep group). At retrieval, participants viewed the backgrounds without their objects, distinguishing new backgrounds from those previously studied. Occipital activity was greater within the sleep group than the wake group specifically during the successful retrieval of neutral backgrounds that had been studied with negative (but not neutral) objects. Moreover, there was enhanced connectivity between the middle occipital gyrus and hippocampus following sleep. Within the sleep group, the percentage of REM sleep obtained correlated with activity in the middle occipital gyrus, lingual gyrus, and cuneus during the successful retrieval of neutral backgrounds previously paired with negative objects. These results confirm that emotion affects neural activity during retrieval even when the cues themselves are neutral, and demonstrate, for the first time, that this residual effect of emotion on visual activity is greater after sleep and may be maximized by REM sleep.     

Culture impacts the magnitude of the emotion-induced memory trade-off effect

The present study assessed the extent to which culture impacts the emotion-induced memory trade-off effect. This trade-off effect occurs because emotional items are better remembered than neutral ones, but this advantage comes at the expense of memory for backgrounds such that neutral backgrounds are remembered worse when they occurred with an emotional item than with a neutral one. Cultures differ in their prioritisation of focal object versus contextual background information, with Westerners focusing more on objects and Easterners focusing more on backgrounds. Americans, a Western culture, and Turks, an Eastern-influenced culture, incidentally encoded positive, negative, and neutral items placed against neutral backgrounds, and then completed a surprise memory test with the items and backgrounds tested separately. Results revealed a reduced trade-off for Turks compared to Americans. Although both groups exhibited an emotional enhancement in item memory, Turks did not show a decrement in memory for backgrounds that had been paired with emotional items. These findings complement prior ones showing reductions in trade-off effects as a result of task instructions. Here, we suggest that a contextual-focus at the level of culture can mitigate trade-off effects in emotional memory.     

Effects of aging and encoding instructions on emotion-induced memory trade-offs

The effects of emotion on memory are often described in terms of trade-offs: People often remember central, emotional information at the expense of background details. The present experiment examined the effects of aging and encoding instructions on participants' ability to remember the details of central emotional objects and the backgrounds on which those objects were placed. When young and older adults passively viewed scenes, both age groups showed strong emotion-induced trade-offs. They were able to remember the visual details as well as the general theme of the emotional object, but they had difficulties remembering the visual specifics of the scene background. Age differences emerged, however, when participants were given encoding instructions that emphasized elaborative encoding of the entire scene. With these instructions, young adults overcame the trade-offs (i.e., they no longer showed impairing effects of emotion), whereas older adults continued to show good memory for the emotional object but poor memory for its background. These results suggest that aging impairs the ability to flexibly disengage attention from the negative arousing elements of scenes, preventing the successful encoding of nonemotional aspects of the environment.     

Binding neutral information to emotional contexts: Brain dynamics of long-term recognition memory

There is abundant evidence in memory research that emotional stimuli are better remembered than neutral stimuli. However, effects of an emotionally charged context on memory for associated neutral elements is also important, particularly in trauma and stress-related disorders, where strong memories are often activated by neutral cues due to their emotional associations. In the present study, we used event-related potentials (ERPs) to investigate long-term recognition memory (1-week delay) for neutral objects that had been paired with emotionally arousing or neutral scenes during encoding. Context effects were clearly evident in the ERPs: An early frontal ERP old/new difference (300–500 ms) was enhanced for objects encoded in unpleasant compared to pleasant and neutral contexts; and a late central-parietal old/new difference (400–700 ms) was observed for objects paired with both pleasant and unpleasant contexts but not for items paired with neutral backgrounds. Interestingly, objects encoded in emotional contexts (and novel objects) also prompted an enhanced frontal early (180–220 ms) positivity compared to objects paired with neutral scenes indicating early perceptual significance. The present data suggest that emotional—particularly unpleasant—backgrounds strengthen memory for items encountered within these contexts and engage automatic and explicit recognition processes. These results could help in understanding binding mechanisms involved in the activation of trauma-related memories by neutral cues.     

Mood-dependent retrieval in visual long-term memory: dissociable effects on retrieval probability and mnemonic precision

Although memories are more retrievable if observers’ emotional states are consistent between encoding and retrieval, it is unclear whether the consistency of emotional states increases the likelihood of successful memory retrieval, the precision of retrieved memories, or both. The present study tested visual long-term memory for everyday objects while consistent or inconsistent emotional contexts between encoding and retrieval were induced using background grey-scale images from the International Affective Picture System (IAPS). In the study phase, participants remembered colours of sequentially presented objects in a negative (Experiment 1a) or positive (Experiment 2a) context. In the test phase, participants estimated the colours of previously studied objects in either negative versus neutral (Experiment 1a) or positive versus neutral (Experiment 2a) contexts. Note, IAPS images in the test phase were always visually different from those initially paired with the studied objects. We found that reinstating negative context and positive context at retrieval resulted in better mnemonic precision and a higher probability of successful retrieval, respectively. Critically, these effects could not be attributed to a negative or positive context at retrieval alone (Experiments 1b and 2b). Together, these findings demonstrated dissociable effects of emotion on the quantitative and qualitative aspects of visual long-term memory retrieval.     

Emotion and vantage point in autobiographical

Autobiographical memories may be recalled from two different perspectives: Field memories in which the person seems to remember the scene from his/her original point of view and observer memories in which the rememberer sees him/herself in the memory image. Here, 122 undergraduates participated in an experiment examining the relation between field vs. observer perspective in memory for 10 different emotional states, including both positive and negative emotions and emotions associated with high vs. low intensity. Observer perspective was associated with reduced sensory and emotional reliving across all emotions. This effect was observed for naturally occurring memory perspective and when participants were instructed to change their perspective from field to observer, but not when participants were instructed to change perspective from observer to field.     

Emotional distress and positive and negative memories from military deployment: the influence of PTSD symptoms and time

During military deployment, soldiers are confronted with both negative and positive events. What is remembered and how it affects an individual is influenced by not only the perceived emotion of the event, but also the emotional state of the individual. Here we examined the most negative and most positive deployment memories from a company of 337 soldiers who were deployed together to Afghanistan. We examined how the level of emotional distress of the soldiers and the valence of the memory were related to the emotional intensity, experience of reliving, rehearsal and coherence of the memories, and how the perceived impact of these memories changed over time. We found that soldiers with higher levels of post-traumatic stress disorder (PTSD) symptoms were more affected by both their negative and positive memories, compared with soldiers with lower levels of PTSD symptoms. Emotional intensity of the most negative memory increased over time in the group with highest levels of PTSD symptoms, but dropped in the other groups. The present study adds to the literature on emotion and autobiographical memory and how this relationship interacts with an individual’s present level of emotional distress and the passage of time.     

Cross-cultural differences in item and background memory: examining the influence of emotional intensity and scene congruency

After viewing a scene, individuals differ in what they prioritise and remember. Culture may be one factor that influences scene memory, as Westerners have been shown to be more item-focused than Easterners (see Masuda, T., & Nisbett, R. E. (2001). Attending holistically versus analytically: Comparing the context sensitivity of Japanese and Americans. Journal of Personality and Social Psychology, 81, 922–934). However, cultures may differ in their sensitivity to scene incongruences and emotion processing, which may account for cross-cultural differences in scene memory. The current study uses hierarchical linear modeling (HLM) to examine scene memory while controlling for scene congruency and the perceived emotional intensity of the images. American and East Asian participants encoded pictures that included a positive, negative, or neutral item placed on a neutral background. After a 20-min delay, participants were shown the item and background separately along with similar and new items and backgrounds to assess memory specificity. Results indicated that even when congruency and emotional intensity were controlled, there was evidence that Americans had better item memory than East Asians. Incongruent scenes were better remembered than congruent scenes. However, this effect did not differ by culture. This suggests that Americans’ item focus may result in memory changes that are robust despite variations in scene congruency and perceived emotion.     

Memory enhancement by a semantically unrelated emotional arousal source induced after learning

It has been well established that moderate physiological or emotional arousal modulates memory. However, there is some controversy about whether the source of arousal must be semantically related to the information to be remembered. To test this idea, 35 healthy young adult participants learned a list of common nouns and afterward viewed a semantically unrelated, neutral or emotionally arousing videotape. The tape was shown after learning to prevent arousal effects on encoding or attention, instead influencing memory consolidation. Heart rate increase was significantly greater in the arousal group, and negative affect was significantly less reported in the non-arousal group after the video. The arousal group remembered significantly more words than the non-arousal group at both 30 min and 24 h delays, despite comparable group memory performance prior to the arousal manipulation. These results demonstrate that emotional arousal, even from an unrelated source, is capable of modulating memory consolidation. Potential reasons for contradictory findings in some previous studies, such as the timing of "delayed" memory tests, are discussed.     

The emotion-induced memory trade-off: More than an effect of overt attention?

Although it has been suggested that many effects of emotion on memory are attributable to attention, in the present study we addressed the hypothesis that such effects may relate to a number of different factors during encoding or postencoding. One way to look at the effects of emotion on memory is by examining the emotion-induced memory trade-off, whereby enhanced memory for emotional items often comes at the cost of memory for surrounding background information. We present evidence that this trade-off cannot be explained solely by overt attention (measured via eyetracking) directed to the emotional items during encoding. Participants did not devote more overt attention to emotional than to neutral items when those items were selectively remembered (at the expense of their backgrounds). Only when participants were asked to answer true/false questions about the items and the backgrounds—a manipulation designed to affect both overt attention and poststimulus elaboration—was there a reduction in selective emotional item memory due to an increase in background memory. These results indicate that the allocation of overt visual attention during encoding is not sufficient to predict the occurrence of selective item memory for emotional items.     

Closure of autobiographical memories: The effects of written recounting from first- or third-person visual perspective

Autobiographical memories are recalled with varying degrees of psychological closure. Closure is a subjective assessment of how far a remembered experience feels resolved, and it has been suggested that one predictor of closure is the amount of emotional detail in the memory. Study 1 examined which aspect of emotional detail is important for closure, and showed that open and closed negative memories were distinguished by ratings of emotion evoked during recall, not by remembered emotion from the time of the event. The recall of open memories was accompanied by more intense, more negative, and less positive emotion than the recall of closed memories. Biased retelling of memories has been shown to influence closure and on the basis of evidence that third-person recall serves a distancing function, Study 2 examined whether instructions to repeatedly recount an open memory from a third-person perspective would increase closure compared with a single or repeated recounting from a first-person perspective. While repeated third-person recounting had the greatest influence on closure, there were also increases in the first-person recounting groups. The results suggest that closure can be increased by reporting memories in written narrative form, particularly if repeatedly expressed from the third-person perspective.     

Closure of autobiographical memories: The effects of written recounting from first- or third-person visual perspective

Autobiographical memories are recalled with varying degrees of psychological closure. Closure is a subjective assessment of how far a remembered experience feels resolved, and it has been suggested that one predictor of closure is the amount of emotional detail in the memory. Study 1 examined which aspect of emotional detail is important for closure, and showed that open and closed negative memories were distinguished by ratings of emotion evoked during recall, not by remembered emotion from the time of the event. The recall of open memories was accompanied by more intense, more negative, and less positive emotion than the recall of closed memories. Biased retelling of memories has been shown to influence closure and on the basis of evidence that third-person recall serves a distancing function, Study 2 examined whether instructions to repeatedly recount an open memory from a third-person perspective would increase closure compared with a single or repeated recounting from a first-person perspective. While repeated third-person recounting had the greatest influence on closure, there were also increases in the first-person recounting groups. The results suggest that closure can be increased by reporting memories in written narrative form, particularly if repeatedly expressed from the third-person perspective.     

Time-limited effects of emotional arousal on item and source memory

Two experiments investigated the time-limited effects of emotional arousal on consolidation of item and source memory. In Experiment 1, participants memorized words (items) and the corresponding speakers (sources) and then took an immediate free recall test. Then they watched a neutral, positive, or negative video 5, 35, or 50?min after learning, and 24 hours later they took surprise memory tests. Experiment 2 was similar to Experiment 1 except that (a) a reality monitoring task was used; (b) elicitation delays of 5, 30, and 45?min were used; and (c) delayed memory tests were given 60?min after learning. Both experiments showed that, regardless of elicitation delay, emotional arousal did not enhance item recall memory. Second, both experiments showed that negative arousal enhanced delayed item recognition memory only at the medium elicitation delay, but not in the shorter or longer delays. Positive arousal enhanced performance only in Experiment 1. Third, regardless of elicitation delay, emotional arousal had little effect on source memory. These findings have implications for theories of emotion and memory, suggesting that emotion effects are contingent upon the nature of the memory task and elicitation delay.     

Emotion's influence on memory for spatial and temporal context

Individuals report remembering emotional items vividly. It is debated whether this report reflects enhanced memory accuracy or a bias to believe emotional memories are vivid. We hypothesised emotion would enhance memory accuracy, improving memory for contextual details. The hallmark of episodic memory is that items are remembered in a spatial and temporal context, so we examined whether an item's valence (positive, negative) or arousal (high, low) would influence its ability to be remembered with those contextual details. Across two experiments, high-arousal items were remembered with spatial and temporal context more often than low-arousal items. Item valence did not influence memory for those details, although positive high-arousal items were recognised or recalled more often than negative items. These data suggest that emotion does not just bias participants to believe they have a vivid memory; rather, the arousal elicited by an event can benefit memory for some types of contextual details.     

Only “efficient” emotional stimuli affect the content of working memory during free-recollection from natural scenes

Emotional events are thought to have privileged access to attention and memory, consuming resources needed to encode competing emotionally neutral stimuli. However, it is not clear whether this detrimental effect is automatic or depends on the successful maintenance of the specific emotional object within working memory. Here, participants viewed everyday scenes including an emotional object among other neutral objects followed by a free-recollection task. Results showed that emotional objects—irrespective of their perceptual saliency—were recollected more often than neutral objects. The probability of being recollected increased as a function of the arousal of the emotional objects, specifically for negative objects. Successful recollection of emotional objects (positive or negative) from a scene reduced the overall number of recollected neutral objects from the same scene. This indicates that only emotional stimuli that are efficient in grabbing (and then consuming) available attentional resources play a crucial role during the encoding of competing information, with a subsequent bias in the recollection of neutral representations.     

Emotion's influence on memory for spatial and temporal context

Individuals report remembering emotional items vividly. It is debated whether this report reflects enhanced memory accuracy or a bias to believe emotional memories are vivid. We hypothesized emotion would enhance memory accuracy, improving memory for contextual details. The hallmark of episodic memory is that items are remembered in a spatial and temporal context, so we examined whether an item's valence (positive, negative) or arousal (high, low) would influence its ability to be remembered with those contextual details. Across two experiments, high-arousal items were remembered with spatial and temporal context more often than low-arousal items. Item valence did not influence memory for those details, although positive high-arousal items were recognized or recalled more often than negative items. These data suggest that emotion does not just bias participants to believe they have a vivid memory; rather, the arousal elicited by an event can benefit memory for some types of contextual details.     

Memory consolidation during sleep: interactive effects of sleep stages and HPA regulation

Sleep is critically involved in the consolidation of previously acquired memory traces. However, nocturnal sleep is not uniform but is subject to distinct changes in electrophysiological and neuroendocrine activity. Specifically, the first half of the night is dominated by slow wave sleep (SWS), whereas rapid eye movement (REM) sleep prevails in the second half. Concomitantly, hypothalamo-pituitary-adrenal (HPA) activity as indicated by cortisol release is suppressed to a minimum during early sleep, while drastically increasing during late sleep. We have shown that the different sleep stages and the concomitant glucocorticoid release are interactively involved in the consolidation of different types of memories. SWS-rich early sleep has been demonstrated to benefit mainly the consolidation of hippocampus-dependent declarative memories (i.e. facts and episodes). In contrast, REM sleep-rich late sleep was shown to improve in particular emotional memories involving amygdalar function, as well as procedural memories (for skills) not depending on hippocampal or amygdalar function. Enhancing plasma glucocorticoid concentrations during SWS-rich early sleep counteracted hippocampus-dependent declarative memory consolidation, but did not affect hippocampus-independent procedural memory. Preventing the increase in cortisol during late REM sleep-rich sleep by administration of metyrapone impaired hippocampus-dependent declarative memory but enhanced amygdala-dependent emotional aspects of memory. The data underscore the importance of pituitary-adrenal inhibition during early SWS-rich sleep for efficient consolidation of declarative memory. The increase in cortisol release during late REM sleep-rich sleep may counteract an overshooting consolidation of emotional memories.     

Negative Emotion Enhances Memory Accuracy: Behavioral and Neuroimaging Evidence

ABSTRACT— There have been extensive discussions about whether emotional memories contain more accurate detail than nonemotional memories do, or whether individuals simply believe that they have remembered emotional experiences more accurately. I review evidence that negative emotion enhances not only the subjective vividness of a memory but also the likelihood of remembering some (but not all) event details. I then describe neuroimaging evidence suggesting that engagement of emotion-processing regions (particularly the amygdala and orbitofrontal cortex) relates to the encoding and retrieval of details intrinsically linked to negative items.     

Emotional Arousal and Memory Binding: An Object-Based Framework

ABSTRACT— Binding various features of an event together and maintaining these connections in memory is an essential component of episodic memories. Previous theories make contradictory predictions about the effects of emotional arousal on memory binding. In this article, I review evidence for both arousal-impaired and arousal-enhanced memory binding and explain these contradictory findings using an object-based framework. According to this framework, emotionally arousing objects attract attention that enhances binding of their constituent features. In contrast, the emotional arousal associated with one object either impairs or has no effect on the associations between that object and other distinct objects or background contextual information. After initial encoding, the attention-grabbing nature of emotionally arousing objects can lead to interference in working memory, making it more difficult to maintain other bound representations. These contrasting effects of arousal on memory binding should help predict which aspects of emotional memories are likely to be accurate and which aspects are likely to be misremembered.