Modulation of retrieval processing reflects accuracy of emotional source memory

There is considerable evidence that encoding and consolidation of memory are modulated by emotion, but the retrieval of emotional memories is not well characterized. Here we manipulated the emotional context with which affectively neutral stimuli were associated during encoding, allowing us to examine neural activity associated with retrieval of emotional memories without confounding the emotional attributes of cue items and the retrieved context. Using a source memory procedure we were also able to compare how retrieval processing was modulated when the emotional encoding context was recollected or not. An interaction between emotional encoding context and accuracy of source memory revealed that successful retrieval of emotional context was associated with activity in left amygdala, and a left frontotemporal network including anterior insula, prefrontal cortex and cingulate. In contrast, when contextual retrieval was unsuccessful, items encoded in emotional contexts elicited enhanced activity in right amygdala and a right-lateralized network that included extrastriate visual areas. These findings indicate distinct effects of emotion on successful and unsuccessful retrieval of source information, including lateralization of amygdala responses.     

Electrophysiological correlates of encoding and retrieving emotional events

This study examined the impact of emotional content on encoding and retrieval processes. Event-related potentials were recorded in a source recognition memory task. During encoding, a posterior positivity for positive and negative pictures (250-450 ms) that presumably reflects attentional capturing of emotionally valenced stimuli was found. Additionally, positive events, which were also rated as less arousing than negative events, gave rise to anterior and posterior slow wave activity as compared with neutral and negative events and also showed enhanced recognition memory. It is assumed that positive low-arousing events enter controlled and elaborated encoding processes that are beneficial for recognition memory performance. The high arousal of negative events may interfere with controlled encoding mechanisms and attenuate item recognition and the quality of remembering. Moreover, topographically distinct late posterior negativities were obtained for the retrieval of the context features location and time that support the view that this component reflects processes in service of reconstructing the study episode by binding together contextual details with an item and that varies with the kind of episodic detail to be retrieved.     

不同情绪背景下来源记忆的ERPs研究*

使用事件相关电位(ERP)技术和测试来源记忆的多键范式,探讨不同情绪效价背景下来源提取的认知神经机制。学习阶段,同时呈现汉字和3种效价情绪图片(重叠);测验阶段,只呈现汉字,要求被试进行四键判断：旧字且背景为中性,旧字且背景为正性,旧字且背景为负性,新字。结果发现：在提取阶段,刺激呈现后300~500 ms,3种情绪背景下来源判断正确项目和来源判断错误项目都比新项目诱发了更正的ERPs(即都存在新旧效应),这一结果反映出刺激呈现后300~500 ms 是一个早期的项目提取阶段,它独立于来源提取。并且,在中性背景下,两种新旧效应没有差别;而在正性和负性背景下,来源判断正确项目的新旧效应显著大于来源判断错误项目。说明相比于中性背景,情绪背景下被试可能更早对来源信息产生熟悉感。在500~650 ms,3种情绪背景下都存在来源判断正确项目的新旧效应,在来源判断错误项目和新项目之间没有显著差异,来源正确与来源错误有显著的新旧效应差异,但二者在头皮分布上是类似的,这反映了晚期的来源提取过程。同时来源正确的新旧效应在这两个时段有显著不同的头皮分布,表明这两个时段有不同加工过程。另外,在500~650 ms,存在显著的情绪效应,正性背景下正确判断来源诱发的ERPs比中性和负性背景下的更正,而且来源判断错误条件下没有情绪效应。综上所述,来源正确和来源错误可能仅仅反映了大脑激活在量上的不同,并不能推断两者存在质的差异;大脑神经活动的早期不仅反映了对记忆项目的熟悉性,而且也受情绪效应的影响,500 ms之后大脑神经活动反映的是对记忆项目回想的过程,这一回想过程也与情绪效应有关,受到情绪效应的调节。     

Attention during memory retrieval enhances future remembering

Memory retrieval is a powerful learning event that influences whether an experience will be remembered in the future. Although retrieval can succeed in the presence of distraction, dividing attention during retrieval may reduce the power of remembering as an encoding event. In the present experiments, participants studied pictures of objects under full attention and then engaged in item recognition and source memory retrieval under full or divided attention. Two days later, a second recognition and source recollection test assessed the impact of attention during initial retrieval on long-term retention. On this latter test, performance was superior for items that had been tested initially under full versus divided attention. More importantly, even when items were correctly recognized on the first test, divided attention reduced the likelihood of subsequent recognition on the second test. The same held true for source recollection. Additionally, foils presented during the first test were also less likely to be later recognized if they had been encountered initially under divided attention. These findings demonstrate that attentive retrieval is critical for learning through remembering.     

The self-choice effect from a multiple-cue perspective

The self-choice effect refers to the fact that self-chosen items are remembered better than experimenter-assigned items (Takahashi, 1991). The present study investigated the hypothesis that (a) response choice involves relational processing as activation of both target and context items, and (b) such activated context items are effective as potential retrieval cues for recall of target items. In the experiment, participants chose (choice condition) or were assigned (force condition) a target to remember for each trial. Prior to free recall of the target items, context words, related new words, or unrelated words were presented in a recognition task as potential retrieval cues. The results of a subsequent free recall test indicated that the incidental cues were more effective in the choice condition than in the force condition. Also, recognition resulted in a greater rate of successfully recognized context words at the cost of increasing falsely recognized related new words in the choice condition in comparison with the force condition. These results indicated that response choice activates context items at encoding, which operate as potential retrieval cues for recall of target items. Such cuing mechanisms operative in the self-choice effect are consistent with the multiple-cue theory proposed by Soraci et al. (1994; see also Soraci et al., 1999) for generative processing.     

Memory retrieval as a self-propagating process

Retrieval of a subset of studied items and the presentation of those items as retrieval cues typically impair retrieval of the other items. Previous research on this self-limiting property of memory retrieval has relied heavily on short retention intervals and similar context between encoding and test. Here, we examined retrieval dynamics also after a prolonged retention interval with different spatial and social context between encoding and test, conditions that mimic people’s remembering in many situations of daily life. For both unrelated word lists and more integrated prose material, we found retrieval and cuing to impair recall of other studied items after a short retention interval, but to improve recall in the prolonged retention interval condition. The results demonstrate that retrieval dynamics depend critically on situation, indicating that quite often in daily life, retrieval may be a self-propagating, rather than a self-limiting process.     

Variations in constrained retrieval

Mentally reinstating encoding operations at retrieval might improve access to memories; however, such constrained retrieval is an effortful process that may not always be used. The memory-for-foils procedure (Jacoby, Shimizu, Daniels, & Rhodes, Psychonomic Bulletin & Review 12, 852-857, 2005) infers participant-initiated mental reinstatement of encoding operations during attempts at recognition from the differential memory that accrues to foils during a test of deeply processed items versus during a test of shallowly processed items, as indicated by performance on a final recognition memory test for the foils. Experiment 1 tested whether differential memory for foils is due to the evocation of task context during recollection of neighboring old items. Experiment 2 tested whether inducing a set to respond without much effort on a prior recognition test affects the likelihood of constrained retrieval on later tests. Experiment 3 tested whether constrained retrieval is less likely to occur when the deep versus shallow source of test items is intermixed, rather than blocked in separate tests. These experiments provide evidence that people query memory by mentally reinstating encoding operations and identify conditions that affect the probability of constrained retrieval.     

Fearful expressions enhance recognition memory: electrophysiological evidence

Facial expressions play a key role in affective and social behavior. However, the temporal dynamics of the brain responses to emotional faces remain still unclear, in particular an open question is at what stage of face processing expressions might influence encoding and recognition memory. To try and answer this question we recorded the event-related potentials (ERPs) elicited in an old/new recognition task. A novel aspect of the present design was that whereas faces were presented during the study phase with either a happy, fearful or neutral expression, they were always neutral during the memory retrieval task. The ERP results showed three main findings: An enhanced early fronto-central positivity for faces encoded as fearful, both during the study and the retrieval phase. During encoding subsequent memory (Dm effect) was influenced by emotion. At retrieval the early components P100 and N170 were modulated by the emotional expression of the face at the encoding phase. Finally, the later ERP components related to recognition memory were modulated by the previously encoded facial expressions. Overall, these results suggest that face recognition is modulated by top-down influences from brain areas associated with emotional memory, enhancing encoding and retrieval in particular for fearful emotional expressions.     

Hippocampal Activations during Repetitive Learning and Recall of Geometric Patterns

Hippocampal activation is required for episodic memory. Encoding and retrieval of novel and memorable items have been related to different locations in the hippocampus; however, the data remain ambiguous. The application of a newly designed keyboard allowed investigation of brain activation during encoding and free immediate and delayed recall with functional magnetic resonance imaging (fMRI) in young healthy controls (n=12). Because of the repetitive learning and recall conditions, an individual learning gradient was used to contrast neural activity at different individual levels of novelty. During learning, subjects were asked to memorize 10 geometric patterns requiring the establishment of intra-item associations for memorization. After learning, subjects were asked to recall the items actively via the keyboard. Learning and recall were alternated five times. Delayed recall was scanned about 15 min after the fifth immediate recall condition without subjects having seen the items again. Left-sided anterior hippocampal activity was observed during conditions of initial learning as well as maximum recall. Neural activity during delayed recall did not reveal hippocampal responses and was characterized by a transition of neural activity from occipitoparietal regions to bilateral temporal cortices. We conclude that both lateralization and segregation depend on the specific relational characteristics of the stimuli requiring establishment of intra-item associations for encoding as well as retrieval. The absence of hippocampal activation during delayed recall together with the increase of lateral temporal involvement possibly corresponds with an emerging transition from episodic to long-term memory.     

编码任务对来源记忆的影响:一项事件相关电位研究

本研究采用ERPs技术,探讨编码任务对来源记忆的影响及其神经机制。在学习阶段,被试进行两种编码操作：匹配判断任务和大小判断任务;在测验阶段,让被试进行来源判断任务。行为结果发现：在编码阶段,匹配判断任务比大小判断任务需要更长的时间;在测验阶段,匹配判断任务在“项目旧/背景旧”和“部分旧”条件的正确率显著高于大小判断任务,匹配判断任务在“项目旧/背景旧”条件的反应时显著短于大小判断任务。测验阶段ERPs结果发现：两种编码任务的新旧效应在200~700ms均有较广泛的头皮分布,而在700~1100ms呈前额皮层最大化分布;匹配判断任务的新旧效应比大小判断任务更大,表现在600~700ms的FPz电极以及700~800ms的Pz、P4电极。由此可见,在来源记忆中,编码任务影响来源记忆,主要表现在后部顶区位置;晚期前额皮层新旧效应不受任务难度因素的影响。     

Cholinergic modulation of the hippocampus during encoding and retrieval of tone/shock-induced fear conditioning

We investigated the role of acetylcholine (ACh) during encoding and retrieval of tone/shock-induced fear conditioning with the aim of testing Hasselmo's cholinergic modulation model of encoding and retrieval using a task sensitive to hippocampal disruption. Lesions of the hippocampus impair acquisition and retention of contextual conditioning with no effect on tone conditioning. Cholinergic antagonists also impair acquisition of contextual conditioning. Saline, scopolamine, or physostigmine was administered directly into the CA3 subregion of the hippocampus 10 min before rats were trained on a tone/shock-induced fear conditioning paradigm. Freezing behavior was used as the measure of learning. The scopolamine group froze significantly less during acquisition to the context relative to controls. The scopolamine group also froze less to the context test administered 24 h posttraining. A finer analysis of the data revealed that scopolamine disrupted encoding but not retrieval. The physostigmine group initially froze less during acquisition to the context, although this was not significantly different from controls. During the context test, the physostigmine group froze less initially but quickly matched the freezing levels of controls. A finer analysis of the data indicated that physostigmine disrupted retrieval but not encoding. These results suggest that increased ACh levels are necessary for encoding new spatial contexts, whereas decreased ACh levels are necessary for retrieving previously learned spatial contexts.     

Emotional valence influences the neural correlates associated with remembering and knowing

In the present study, we examined whether emotional valence modulates the neural processes that are engaged during the encoding of information that is later vividly remembered versus that which is only known to be familiar. Participants underwent an fMRI scan while viewing positive, negative, and neutral stimuli. Later, recognized items were labeled as either remembered or known. Negative items that were later vividly remembered recruited temporo-occipital regions associated with sensory processing more than did positive or neutral items that were vividly remembered. The encoding of positive information later known recruited the cingulate gyrus and bilateral frontal and parietal areas—regions associated with episodic and semantic retrieval and self-referential processing—more than did the encoding of negative or neutral items that were later known. These results suggest that memories for negative items may be vividly recollected due to increased sensory processing during encoding, whereas enhanced gist-based processing of positive information may lead to increased feelings of familiarity.     

Retrieval during learning facilitates subsequent memory encoding

In multiple-list learning, retrieval during learning has been suggested to improve recall of the single lists by enhancing list discrimination and, at test, reducing interference. Using electrophysiological, oscillatory measures of brain activity, we examined to what extent retrieval during learning facilitates list encoding. Subjects studied 5 lists of items in anticipation of a final cumulative recall test and did either a retrieval or a no-retrieval task between study of the lists. Retrieval was from episodic memory (recall of the previous list), semantic memory (generation of exemplars from an unrelated category), or short-term memory (2-back task). Behaviorally, all 3 forms of retrieval enhanced recall of both previously and subsequently studied lists. Physiologically, the results showed an increase of alpha power (8-14 Hz) from List 1 to List 5 encoding when no retrieval activities were interpolated but no such increase when any of the 3 retrieval activities occurred. Brain-behavior correlations showed that alpha-power dynamics from List 1 to List 5 encoding predicted subsequent recall performance. The results suggest that, without intermittent retrieval, encoding becomes ineffective across lists. In contrast, with intermittent retrieval, there is a reset of the encoding process for each single list that makes encoding of later lists as effective as encoding of early lists.     

注意促进效应的ERP研究

近期发现,采用目标探测任务作为二级任务时,目标探测条件不仅不会削弱,甚至会提高记忆成绩,产生注意促进效应（ABE）。为进一步探究目标探测的作用机制,研究采用事件相关电位（ERP）技术,对比目标与分心条件在编码和提取过程中所诱发的ERP差异。结果表明,两类条件在再认过程中所诱发的ERP差异不仅表现在早期与知觉加工相关的ERP成份上,并且一直持续到了后期与细节提取相关的ERP成分上。由此推测,ABE不仅体现在编码早期的知觉加工阶段,更会延续到后期的复述与精加工。     

“Wanted!” The effects of reward on face recognition: electrophysiological correlates

The aim of the present study was to uncover the temporal dynamics of face recognition as a function of reward. Event-related potentials (ERPs) were recorded during the encoding and the subsequent old/new memory test in response to faces that could be associated with a monetary reward. The behavioral results showed that faces associated with reward at both encoding and retrieval were recognized better than the unrewarded ones. ERP responses highlighted that successful encoding predictive of subsequent memory was greater for faces associated with reward than for faces without reward-driven motivational learning. At retrieval, an early positive-going component was elicited for potentially rewarded faces on frontal regions, while the occipito-temporal N170 component showed priming effects as a function of reward. Later on, larger centro-parietal ERP components, related to recognition memory, were found selectively for reward-associated faces. Remarkably, electrophysiological responses varied in a graded manner, with the largest amplitude yielded by faces with double reward, followed by faces associated with reward only at encoding. Taken together, the present data show that the processing of outcome expectations affects face structural encoding and increases memory efficiency, yielding a robust and sustained modulation over frontal and temporal areas where reward and memory mechanisms operate in conjunction.     

Functional asymmetry of human prefrontal cortex: encoding and retrieval of verbally and nonverbally coded information

There are several views about the organization of memory functions in the human prefrontal cortex. One view assumes a process-specific brain lateralization according to different memory subprocesses, that is, encoding and retrieval. An alternative view emphasizes content-specific lateralization of brain systems involved in memory processes. This study addresses this apparent inconsistency between process- and content-specific lateralization of brain activity by investigating the effects of verbal and nonverbal encoding on prefrontal activations during encoding and retrieval of environmental novel sounds using fMRI. An intentional memory task was applied in which subjects were required either to judge the sounds' loudness (nonverbal encoding task) or to indicate whether or not a sound can be verbally described (verbal encoding task). Retrieval processes were examined in a subsequent yes/no recognition test. In the study phase the right posterior dorsolateral prefrontal cortex (PFC) was activated in both tasks. During verbal encoding additional activation of the left dorsolateral PFC was obtained. Retrieval-related fMRI activity varied as a function of encoding task: For the nonverbal task we detected an activation focus in the right posterior dorsolateral PFC whereas an activation in the left dorsolateral PFC was observed for the verbal task. These findings indicate that the right dorsolateral PFC is engaged in encoding of auditory information irrespective of encoding task. The lateralization of PFC activity during retrieval was shown to depend on the availability of verbal codes, with left hemispheric involvement for verbally and right hemispheric activation for nonverbally coded information.     

故事情境下真实记忆与虚假记忆的活动差异

以故事情境为实验材料,采用学习再认实验范式,通过操纵再认靶词与故事情境的关联性考察了情境故事对关联虚假记忆和真实再认神经活动的影响。结果发现：在记忆再认的熟悉性加工和回想性加工阶段,虚假记忆与真实记忆所诱发FN400和LPC成分走向一致;在额叶后提取阶段,虚假再认所诱发的晚慢成分波幅大于真实再认。结果说明：情境性关联虚假记忆与真实记忆表现出相似的熟悉性和回想性加工,但在后期需耗费额外的加工资源进行成功的提取。     

The effect of context variability on source memory

High-context-variability (HCV) items are experienced in many more preexperimental semantic contexts than are low-context-variability (LCV) items. LCV confers an advantage to recognition memory (Steyvers & Malmberg, 2003). In the present study, we tested whether or not that advantage could be causedby enhanced source memory. Both context variability and word frequency were manipulated, and both factors generally affected source monitoring. Accuracy was better for LCV items than for HCV items and better for low- than for high-word-frequency items. We also considered whether context variability exerts its influence at encoding or at retrieval. We concluded that better recognition memory for LCV items was due, in part, to better retrieval of contextual details that specify how an item was originally studied.     

The Relation of ERP Components to Complex Memory Processing

The relation between various ERP components generated during encoding of a word and its subsequent recall were investigated using a “rote” serial-order and an “elaborative” category memory task. Words (flashed separately) were time-locked to EEG recordings from 21 cortical sites. ERP components from the five subjects having the highest recall scores were compared to the five lowest scoring subjects. Results based on the P200 peak amplitude data as well as the N400 and late positive component peak amplitude and latency data suggest that anterior and posterior distributional differences are elicited during encoding of words for rote and elaborative memory tasks. Furthermore, strong individual differences in these patterns were found as a function of task. A tentative argument was made that the obtained anterior and posterior differences may index different word feature selection and encoding processes, which are differentially utilized by high and low recallers.     

Episodic encoding and recognition of pictures and words: role of the human medial temporal lobes

In the present PET study, we examined brain activity related to processing of pictures and printed words in episodic memory. Our goal was to determine how the perceptual format of objects (verbal versus pictorial) is reflected in the neural organization of episodic memory for common objects. We investigated this issue in relation to encoding and recognition with a particular focus on medial temporal-lobe (MTL) structures. At encoding, participants saw pictures of objects or their written names and were asked to make semantic judgments. At recognition, participants made yes-no recognition judgments in four different conditions. In two conditions, target items were pictures of objects; these objects had originally been encoded either in picture or in word format. In two other conditions, target items were words; they also denoted objects originally encoded either as pictures or as words. Our data show that right MTL structures are differentially involved in picture processing during encoding and recognition. A posterior MTL region showed higher activation in response to the presentation of pictures than of words across all conditions. During encoding, this region may be involved in setting up a representation of the perceptual information that comprises the picture. At recognition, it may play a role in guiding retrieval processes based on the perceptual input, i.e. the retrieval cue. Another more anterior right MTL region was found to be differentially involved in recognition of objects that had been encoded as pictures, irrespective of whether the retrieval cue provided was pictorial or verbal in nature; this region may be involved in accessing stored pictorial representations. Our results suggest that left MTL structures contribute to picture processing only during encoding. Some regions in the left MTL showed an involvement in semantic encoding that was picture specific; others showed a task-specific involvement across pictures and words. Together, our results provide evidence that the involvement of some but not all MTL regions in episodic encoding and recognition is format specific.