Emotionally arousing pictures increase blood glucose levels and enhance recall

Arousal enhances memory in human participants and this enhancing effect is likely due to the release of peripheral epinephrine. As epinephrine does not readily enter the brain, one way that peripheral epinephrine may enhance memory is by increasing circulating blood glucose levels. The present study investigated the possibility that emotionally arousing color pictures would improve memory and elevate blood glucose levels in human participants. Blood glucose levels were measured before, 15 min, and 30 min after male university students viewed 60 emotionally arousing or relatively neutral pictures. Participants viewed each picture for 6 s and then had 10 s to rate the arousal (emotional intensity) and valence (pleasantness) of each picture. A free-recall memory test was given 30 min after the last picture was viewed. Although the emotionally arousing and neutral picture sets were given comparable valence ratings, participants who viewed the emotionally arousing pictures rated the pictures as being more arousing, recalled more pictures, and had higher blood glucose levels after viewing the pictures than did participants who viewed the neutral pictures. These findings indicate that emotionally arousing pictures increase blood glucose levels and enhance memory, and that this effect is not due to differences in the degree of pleasantness of the stimuli. These findings support the possibility that increases in circulating blood glucose levels in response to emotional arousal may be part of the biological mechanism that allows emotional arousal to enhance memory.     

Memory bias for negative emotional words in recognition memory is driven by effects of category membership

Recognition memory studies often find that emotional items are more likely than neutral items to be labelled as studied. Previous work suggests this bias is driven by increased memory strength/familiarity for emotional items. We explored strength and bias interpretations of this effect with the conjecture that emotional stimuli might seem more familiar because they share features with studied items from the same category. Categorical effects were manipulated in a recognition task by presenting lists with a small, medium or large proportion of emotional words. The liberal memory bias for emotional words was only observed when a medium or large proportion of categorised words were presented in the lists. Similar, though weaker, effects were observed with categorised words that were not emotional (animal names). These results suggest that liberal memory bias for emotional items may be largely driven by effects of category membership.     

Effects of emotional content on working memory capacity

Emotional events tend to be remembered better than neutral events, but emotional states and stimuli may also interfere with cognitive processes that underlie memory performance. The current study investigated the effects of emotional content on working memory capacity (WMC), which involves both short term storage and executive attention control. We tested competing hypotheses in a preregistered experiment (N?=?297). The emotional enhancement hypothesis predicts that emotional stimuli attract attention and additional processing resources relative to neutral stimuli, thereby making it easier to encode and store emotional information in WMC. The emotional impairment hypothesis, by contrast, predicts that emotional stimuli interfere with attention control and the active maintenance of information in working memory. Participants completed a common measure of WMC (the operation span task; Turner, M. L., &; Engle, R. W. [1989]. Is working memory capacity task dependent? Journal of Memory and Language, 28, 127–154) that included either emotional or neutral words. Results revealed that WMC was reduced for emotional words relative to neutral words, consistent with the emotional impairment hypothesis.     

无关工作记忆表征的负性情绪信息能否捕获视觉注意?一项眼动研究

研究通过分析视觉搜索任务的首次注视点和行为反应时, 探讨无关工作记忆表征的负性情绪信息对视觉注意选择的影响。实验1发现在反映早期注意选择的首次注视点百分率指标上, 不管工作记忆表征的情绪效价如何, 均出现了显著的注意捕获效应; 实验2发现当采用中性情绪靶子刺激时, 首次注视点百分率指标上仍表现出了稳健的注意捕获效应; 在首次注视点持续时间指标上, 实验1和实验2均发现记忆匹配条件的干扰刺激显著小于控制条件的干扰刺激, 表现出注意的快速脱离; 而在行为反应时指标上, 早期的注意捕获效应消失(实验1), 甚至被反转为注意抑制效应(实验2)。这些结果表明在早期注意选择阶段, 记忆驱动的注意捕获效应不受工作记忆表征情绪效价的影响, 但认知控制会在早期注意捕获之后促使注意快速脱离记忆匹配的干扰刺激, 其作用效果受靶子刺激情绪效价的调节。     

Acute effects of alcohol on memory: Impact of emotional context and serial position

Although the amnestic effects of alcohol in humans are well known, its effects on emotional memory are unclear. In this study, using a randomized double-blind placebo-controlled design, we examine narrative emotional episodic memory in healthy human female volunteers (n = 32) who received either a single dose of alcohol (0.6 g/kg), or a placebo and then viewed neutral story elements presented in either a neutral or emotional context. Memory was tested for gist and detail of the neutral elements 3 days later in a surprise recognition test. Since alcohol modulates GABAergic neurotransmission and may exert its effects on emotion through the limbic system, we predicted that acute alcohol treatment would reduce the expected emotional memory-advantage for gist, leaving detail memory relatively unaffected. Furthermore, given previous findings showing that ‘primacy’ memory is enhanced by physiological arousal, we predicted that reduced arousal produced by alcohol would have the opposite effect and impair primacy memory relative to the middle or ‘recency’ sections of the narrative. Emotional arousal was expected to oppose this effect, so impaired primacy memory following alcohol was only expected in the neutral version of the narrative. Although there was a main effect of story phase (though not of story version), contrary to expectations, alcohol impaired primacy memory for emotionally encoded neutral material. The results suggest that under certain circumstances emotional context or physiological arousal make memories labile and susceptible to disruption through pharmacological manipulation during encoding.     

Immediate and prolonged effects of cortisol, but not propranolol, on memory retrieval in healthy young men

BackgroundWhile acute cortisol administration has been found to impair retrieval of emotional memories in healthy subjects, the duration of this memory impairment is still unknown. Propranolol, on the other hand, may impair the reconsolidation of emotional memories during reactivation, although human studies examining such effects are scarce. The present investigation was therefore undertaken to examine the immediate and prolonged effects of a single administered dose of cortisol or propranolol on memory retrieval in a double-blind placebo controlled design.MethodsEighty-five healthy male participants were asked to retrieve previously learned emotional and neutral information after ingestion of 35 mg cortisol, 80 mg propranolol or placebo. After a washout period of 1 week, recall was again tested.ResultsMemory retrieval of neutral and emotional information was impaired by a single dose of cortisol compared to placebo. The memory impairment due to cortisol remained, even after a washout period of 1 week. No immediate or prolonged effects of propranolol on memory retrieval were found, despite significant reductions in sympathetic arousal.ConclusionsThese results lend support to the hypothesis that cortisol is able to attenuate (emotional) memory recall in men over longer time spans and may therefore augment the treatment of disorders like post-traumatic stress disorder and phobias, but do not clarify the mechanism(s) through which propranolol exerts its therapeutic effects.     

Effect of negative emotional content on working memory and long-term memory

In long-term memory, negative information is better remembered than neutral information. Differences in processes important to working memory may contribute to this emotional memory enhancement. To examine the effect that the emotional content of stimuli has on working memory performance, the authors asked participants to perform working memory tasks with negative and neutral stimuli. Task accuracy was unaffected by the emotional content of the stimuli. Reaction times also did not differ for negative relative to neutral words, but on an n-back task using faces, participants were slower to respond to fearful faces than to neutral faces. These results suggest that although emotional content does not have a robust effect on working memory, in some instances emotional salience can impede working memory performance.     

Working memory in social anxiety disorder: better manipulation of emotional versus neutral material in working memory

Individuals with social anxiety disorder (SAD) engage in post-event processing, a form of perseverative thinking. Given that deficits in working memory might underlie perseverative thinking, we examined working memory in SAD with a particular focus on the effects of stimulus valence. SAD (n?=?31) and healthy control (n?=?20) participants either maintained (forward trials) or reversed (backward trials) in working memory the order of four emotional or four neutral pictures, and we examined sorting costs, which reflect the extent to which performance deteriorated on the backward trials compared to the forward trials. Emotionality of stimuli affected performance of the two groups differently. Whereas control participants exhibited higher sorting costs for emotional stimuli compared to neutral stimuli, SAD participants exhibited the opposite pattern. Greater attention to emotional stimuli in SAD might facilitate the processing of emotional (vs. neutral) stimuli in working memory.     

Emotional distraction unbalances visual processing

Brain mechanisms used to control nonemotional aspects of cognition may be distinct from those regulating responses to emotional stimuli, with activity of the latter being detrimental to the former. Previous studies have shown that suppression of irrelevant emotional stimuli produces a largely right-lateralized pattern of frontal brain activation, thus predicting that emotional stimuli may invoke temporary, lateralized costs to performance on nonemotional cognitive tasks. To test this, we briefly (85 ms) presented a central, irrelevant, expressive (angry, happy, sad, or fearful) or neutral face 100 ms prior to a letter search task. The presentation of emotional versus neutral faces slowed subsequent search for targets appearing in the left, but not the right, hemifield, supporting the notion of a right-lateralized, emotional response mechanism that competes for control with nonemotional cognitive processes. Presentation of neutral, scrambled, or inverted neutral faces produced no such laterality effects on visual search response times.     

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.     

The role of attention in the associative binding of emotionally arousing words

In the present study, we examined the role of attention in modulating the memory benefit of emotional arousal for same-valence word pair associations. To assess the role of attention either at encoding or at retrieval, participants studied lists of positive, neutral, and negative words pairs under full attention, divided attention at encoding, or divided attention at retrieval, and then were tested on the single words and on the associations between words. Consistent with past studies, memory accuracy was higher for emotional items than for neutral items, and no memory difference was observed across emotional arousal conditions for associations when encoding occurred under full attention. In contrast, memory accuracy was higher for emotionally arousing items and associations relative to neutral items when encoding occurred under divided attention. Finally, dividing attention at retrieval revealed similar effects across emotion conditions, suggesting that retrieval of emotional stimuli relative to neutral stimuli, unlike encoding, does not benefit from automatic processing. The discussion emphasizes the role of automatic processing during encoding in producing the benefit of emotionally enhanced memory, as well as the extent to which controlled attention is responsible for eliminating or reversing (relative to neutral materials) emotionally enhanced memory for associations. Additionally, the implications of the divided-attention-at-retrieval manipulation include consideration of the way in which emotional items may be consciously processed during encoding.     

Event-related potentials of emotional memory: encoding pleasant,unpleasant, and neutral pictures

Emotional events tend to be remembered better than nonemotional events. We investigated this phenomenon by measuring two event-related potential (ERP) effects: the emotion effect (more positive ERPs for pleasant or unpleasant stimuli than for neutral stimuli) and the subsequent memory effect (more positive ERPs for subsequently remembered items than for subsequently forgotten items). ERPs were measured while subjects rated the emotional content of pleasant, unpleasant, and neutral pictures. As was expected, subsequent recall was better for pleasant and unpleasant pictures than for neutral pictures. The emotion effect was sensitive to arousal in parietal electrodes and to both arousal and valence in frontocentral electrodes. The subsequent memory effect at centroparietal electrodes was greater for emotional pictures than for neutral pictures during an early epoch (400-600 msec). This result suggests that emotional information has privileged access to processing resources, possibly leading to better memory formation.     

双加工再认提取中的情绪记忆增强效应

根据双加工理论, 再认提取可以划分为两种不同的加工：熟悉性与回想。以往的研究普遍认为情绪对再认提取具有增强效应, 但是情绪对再认记忆中两种不同加工方式(熟悉性与回想)的增强效应的神经机制尚存争议。本研究以ERP作为测量手段, 采用修正的“记得/知道”范式对情绪图片进行再认测验, 旨在探究在学习?测验间隔较长的条件下, 情绪信息如何对熟悉性与回想产生增强效应。在行为结果中, 对于知道判断, 情绪图片的记忆表现显著高于中性图片, 情绪效价间没有差异; 对于记得判断, 负性情绪图片的记忆表现要显著高于正性与中性图片。在ERP数据上, 对于知道判断, 情绪图片的FN400新旧效应有增强作用。对于记得判断, 负性图片的LPC的新旧效应波幅显著高于中性与负性图片; 而正性图片的LPC新旧效应增强作用不显著。以上证据表明：学习?测验间隔较长的情况下, 熟悉性再认提取中, 情绪记忆增强效应主要体现在情绪唤醒对记忆强度的提高, 而不仅仅体现在情绪效价对熟悉判断反应倾向的影响。在回想的再认提取中, 情绪效价的调节占据主导地位, 负性情绪图片有显著的情绪记忆增强效应; 正性情绪图片则不具有增强效应。     

Enhanced selective memory consolidation following post-learning pleasant and aversive arousal

It is well established that emotions modulate memory, typically enhancing consolidation through post-learning arousal. However, many aspects of this phenomenon have yet to be delineated. For example, it remains unclear whether or not the type of arousal is relevant (pleasant vs. aversive), whether arousal enhances memory selectively for some stimuli but not others (emotional vs. neutral), which specific aspects of the stimulus representation (gist vs. detail) are affected, and whether these mechanisms are sexually dimorphic. In order to explore these issues, 178 undergraduate participants viewed a series of negative, positive and neutral pictures. They were then subjected to a post-learning arousal manipulation in the form of a pleasantly arousing-, aversively arousing-, or neutral video. Free recall tests one week later indicated that both pleasant and aversive post-learning arousal enhanced memory consolidation for positive and negative but not neutral stimuli, independent of the participants' sex. Further analysis for gist and detail aspects suggests that post-learning arousal enhances memory for the gist of the stimuli. The study has implications for the understanding of healthy and pathological cognitive-affective processes in humans.     

Shuttle-box avoidance learning in mice: improvement by glucose combined with stimulant drugs

Glucose was tested alone or in combination with two stimulant drugs, amphetamine and nicotine, in mice of the CD-1 strain subjected to five daily shuttle-box training sessions. Pretraining intraperitoneal administration of glucose (50 or 100 mg/kg) had no effect, while amphetamine and nicotine, given alone, significantly improved avoidance acquisition at a dose of 0.5 mg/kg, but not 0.025 mg/kg. Significant improvement of avoidance learning was also produced by a combination of glucose with the lower dose of amphetamine or nicotine. This enhancing action, produced by a combination of glucose and stimulant drugs, at doses ineffective by themselves, might be due to a concomitant cholinergic and dopaminergic activation, induced by glucose and stimulant drugs, respectively.     

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.     

The emotional carryover effect in memory for words

Emotional material rarely occurs in isolation; rather it is experienced in the spatial and temporal proximity of less emotional items. Some previous researchers have found that emotional stimuli impair memory for surrounding information, whereas others have reported evidence for memory facilitation. Researchers have not determined which types of emotional items or memory tests produce effects that carry over to surrounding items. Six experiments are reported that measured carryover from emotional words varying in arousal to temporally adjacent neutral words. Taboo, non-taboo emotional, and neutral words were compared using different stimulus onset asynchronies (SOAs), recognition and recall tests, and intentional and incidental memory instructions. Strong emotional memory effects were obtained in all six experiments. However, emotional items influenced memory for temporally adjacent words under limited conditions. Words following taboo words were more poorly remembered than words following neutral words when relatively short SOAs were employed. Words preceding taboo words were affected only when recall tests and relatively short retention intervals were used. These results suggest that increased attention to the emotional items sometimes produces emotional carryover effects; however, retrieval processes also contribute to retrograde amnesia and may extend the conditions under which anterograde amnesia is observed.     

Reduced memory for the spatial and temporal context of unpleasant words

Emotional stimuli are consistently better remembered than neutral stimuli. However, the reported effects of emotional stimuli on source memory are less consistent. In four experiments, we examined spatial and temporal source memory and free recall for emotional words previously studied in an fMRI experiment. In the fMRI experiment, the unpleasant but not the pleasant words were shown to activate the amygdala. In the experiments reported here, spatial and temporal source memory were reduced for the unpleasant words compared to pleasant and neutral words. Reduced spatial memory for unpleasant words was observed across studies that varied the specific stimuli, task conditions, and incidental versus intentional encoding instructions. Free recall was enhanced for the pleasant and unpleasant words compared to neutral words. The results demonstrate reduced source memory for unpleasant words even when item recall is enhanced. Our findings are consistent with prior reports of amygdala-mediated amnestic effects evoked by unpleasant stimuli.     

The enhancing effects of hippocampal infusions of glucose are not restricted to spatial working memory

Extensive evidence shows that hippocampal infusions of glucose enhance spontaneous alternation (SA) performance or reverse deficits in this task. The current experiments determined whether the enhancing effects of hippocampal infusions of glucose are restricted to spatial working memory. Specifically we tested whether hippocampal infusions of glucose would reverse deficits in an emotional reference memory task (continuous multiple trial inhibitory avoidance [CMIA]) produced by septal infusions of the gamma-aminobutyric acid agonist muscimol. Male Sprague-Dawley rats were given septal infusions of vehicle or muscimol (0.15 nmol: SA; 5 nmol: CMIA) combined with hippocampal infusions of vehicle or glucose (50 nmol) 15 min prior to assessing SA or CMIA training. CMIA retention was tested 48 h later. Muscimol infusions decreased percent alternation scores and avoidance retention latencies. Importantly, hippocampal infusions of glucose reversed the deficits produced by the septal muscimol infusions on both tasks. These findings show for the first time that hippocampal glucose infusions also influence emotional memory, indicating that the enhancing effects of glucose generalize to memory tasks that vary in motivational and cognitive demand.     

Cannabinoid receptors activation and glucocorticoid receptors deactivation in the amygdala prevent the stress-induced enhancement of a negative learning experience

The enhancement of emotional memory is clearly important as emotional stimuli are generally more significant than neutral stimuli for surviving and reproduction purposes. Yet, the enhancement of a negative emotional memory following exposure to stress may result in dysfunctional or intrusive memory that underlies several psychiatric disorders. Here we examined the effects of stress exposure on a negative emotional learning experience as measured by a decrease in the magnitude of the expected quantity of reinforcements in an alley maze. In contrast to other fear-related negative experiences, reward reduction is more associated with frustration and is assessed by measuring the latency to run the length of the alley to consume the reduced quantity of reward. We also examined whether the cannabinoid receptors agonist WIN55,212-2 (5 μg/side) and the glucocorticoid receptors (GRs) antagonist RU-486 (10 ng/side) administered into the rat basolateral amygdala (BLA) could prevent the stress-induced enhancement. We found that intra-BLA RU-486 or WIN55,212 before stress exposure prevented the stress-induced enhancement of memory consolidation for reduction in reward magnitude. These findings suggest that cannabinoid receptors and GRs in the BLA are important modulators of stress-induced enhancement of emotional memory.