Brain activation while forming memories of fearful and neutral faces in women and men.

Event-related functional MRI (fMRI) was used to assess brain activity during encoding of fearful and neutral faces in 12 women and 12 men. In a subsequent memory analysis, the authors separated successful from unsuccessful encoding of both types of faces, based on whether they were remembered or forgotten in a later recognition memory test. Overall, women and men recruited overlapping neural circuitries. Both sexes activated right-sided medial-temporal regions during successful encoding of fearful faces. Successful encoding of neutral faces was associated with left-sided lateral prefrontal and right-sided superior frontal activation in both sexes. In women, relatively greater encoding related activity for neutral faces was seen in the superior parietal and parahippocampal cortices. By contrast, men activated the left and right superior/middle frontal cortex more than women during successful encoding of the same neutral faces. These findings suggest that women and men use similar neural networks to encode facial information, with only subtle sex differences observed for neutral faces. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Effects of traumatic brain injury on working memory-related brain activation in adolescents.

Eight adolescents (ages 13-18 years) who sustained traumatic brain injury (TBI) and eight gender- and age-matched typically developing (TD) adolescents underwent event-related functional MRI (fMRI) while performing a Sternberg letter recognition task. Encoding, maintenance, and retrieval were examined with memory loads of one or four items during imaging. Both groups performed above a 70% accuracy criterion and did not differ in performance. TD adolescents showed greater increase in frontal and parietal activation during high-load relative to low-load maintenance than the TBI group. The TBI patients showed greater increase in activation during high-load relative to low-load encoding and retrieval than the TD group. Results from this preliminary study suggest that the capability to differentially allocate neural resources according to memory load is disrupted by TBI for the maintenance subcomponent of working memory. The overrecruitment of frontal and extrafrontal regions during encoding and retrieval following TBI may represent a compensatory process. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Memory encoding and retrieval in frontotemporal dementia and Alzheimer's disease.

Memory encoding and retrieval strategies were assessed in patients with behavior-executive variant frontotemporal dementia (FTD), language variant FTD, and Alzheimer's disease (AD) using verbal and visuospatial supraspan learning tests. FTD patients obtained higher free recall, cued recall, and recognition scores than AD patients. Comparison of free recall scores with cued recall and recognition scores was similar in the 3 dementia groups. Groups did not differ in semantic clustering strategies during learning, but serial-order recall was more common in FTD patients. These data do not support the idea that FTD patients' poor memory is due to a selective retrieval disorder, though FTD patients may fail to implement sophisticated organizational strategies during learning. FTD patients' retained capacity for encoding new information into long-term declarative memory is likely due to relatively spared medial temporal lobe involvement. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The psychobiological basis of posttraumatic stress disorder

Posttraumatic stress disorder is a disorder with an identifiable etiological factor (exposure to a traumatic event) and with a complex symptomatology (i.e. intrusive memories, avoidance, hyperarousal) that suggests dysfunction in multiple psychobiological systems. This review considers studies of the neurobiological consequences of acute and chronic stress showing that traumatic experiences can produce long-lasting alterations in multiple neurochemical systems. The role of the locus coeruleus noradrenergic system, prefrontal cortex dopaminergic system, endogenous opiates, hypothalamic-pituitary-adrenal axis, and cortico-releasing factors are reviewed. Several models of PTSD are highlighted, including fear conditioning, kindling, and sensitization. In particular, fear conditioning to explicit and contextual cues is proposed as a model for intrusive memories reactivated by trauma-related stimuli and hyperarousal, respectively. It is argued that the amygdala plays a crucial role in the encoding and retrieval of fear memories activated by specific stimuli that have been associated with aversive events. Association involving more complex environmental stimuli and aversive events may require the involvement of the hippocampus and the bed nucleus of the stria terminalis. Repeated activation of conditioned fear memories may produce a kindling-like process which results in spontaneous intrusive memories.     

Involvement of the supramammillary nucleus in aversive conditioning.

Mechanisms for the retention and retrieval of conditioned taste aversions (CTAs) have yet to be fully defined. The authors explored relevant subcortical forebrain regions by tracking the expression of immediate early genes, c-fos and zif268. The supramammillary nucleus (SuM) was activated following both viscerally based CTA and somatically based inhibitory avoidance (IA). Excitotoxic lesions of the SuM before conditioning caused no disruption of acquisition but accelerated the extinction of both the CTA and IA. In contrast, lesions after CTA conditioning did not impair retention or retrieval. The present study indicates that the SuM is activated by memory-elicited discomfort during retrieval, suggesting that it plays a role in resisting the extinction of a long-term aversive memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of bilingualism, aging, and semantic relatedness on memory under divided attention.

We examined how encoding and retrieval processes were affected by manipulations of attention, and whether the degree of semantic relatedness between words in the memory and distracting task modulated these effects. We also considered age and bilingual status as mediating factors. Monolingual and bilingual younger and older adults studied a list of words from a single semantic category presented auditorily, and later free recalled them aloud. During either study or retrieval, participants concurrently performed a distracting task requiring size decisions to words from either the same or a different semantic category as the words in the memory task. The greatest disruptions of memory from divided attention (DA) were for encoding rather than retrieval. The effect of semantic relatedness was significant only for DA at encoding. Older age and bilingualism were associated with lower recall scores in all conditions, but these factors did not influence the magnitude of memory interference. The results suggest that encoding is more sensitive to semantic similarity in a distracting task than is retrieval. The role of attention at encoding and retrieval is discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Comparative evaluation of two models for estimating sample sizes for tests on trends across repeated measurements

The emotional content of stimuli can enhance memory for those stimuli. This process may occur via an interaction with systems responsible for perception and memory or via the addition of distinct brain regions specialized for emotion which augment mnemonic processing. We performed an 15O PET study to identify neuroanatomical systems which encode visual stimuli with strong negative emotional valence compared to stimuli with neutral valence. Subjects also performed a recognition memory task for these same images, mixed with distracters of similar emotional valence. The experimental design permitted us to independently test effects of emotional content and recognition memory on regional activity. We found activity in the left amygdaloid complex associated with the encoding of emotional stimuli, although this activation appeared early in the scanning session and was not detectable during recognition memory. Visual recognition memory recruited the right middle frontal gyrus and the superior anterior cingulate cortex for both negative and neutral stimuli. An interaction occurred between emotional content and recognition in the lingual gyrus, where greater activation occurred during recognition of negative images compared to recognition of neutral images. Instead of distinct neuroanatomical systems for emotion augmenting memory, we found that emotionally salient stimuli appeared to enhance processing of early sensory input during visual recognition.     

The low-frequency encoding disadvantage: Word frequency affects processing demands.

Low-frequency words produce more hits and fewer false alarms than high-frequency words in a recognition task. The low-frequency hit rate advantage has sometimes been attributed to processes that operate during the recognition test (e.g., L. M. Reder et al., 2000). When tasks other than recognition, such as recall, cued recall, or associative recognition, are used, the effects seem to contradict a low-frequency advantage in memory. Four experiments are presented to support the claim that in addition to the advantage of low-frequency words at retrieval, there is a low-frequency disadvantage during encoding. That is, low-frequency words require more processing resources to be encoded episodically than high-frequency words. Under encoding conditions in which processing resources are limited, low-frequency words show a larger decrement in recognition than high-frequency words. Also, studying items (pictures and words of varying frequencies) along with low-frequency words reduces performance for those stimuli. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Implicit false memory in the DRM paradigm: Effects of amnesia, encoding instructions, and encoding duration.

Recent studies with the Deese/Roediger–McDermott (Deese 1959; Roediger & McDermott, 1995) paradigm have revealed that amnesic patients do not only show impaired veridical memory, but also diminished false memory for semantically related lure words. Due to the typically used explicit retrieval instructions, however, this finding may reflect problems at encoding, at recollection, or both. Therefore, the present experiments examined implicit as well as explicit false memory in patients suffering from Korsakoff’s syndrome and controls. In Experiment 1, encoding instructions either focused on remembering individual list words, or on discovering semantic relationships among the words. In Experiment 2, different presentation durations were used. Results emphasize the distinction between automatic and intentional retrieval: Korsakoff patients’ veridical and false memory scores were diminished when explicit recollection was required, but not when memory was tested implicitly. Encoding manipulations only significantly affected veridical memory: Priming was reduced with thematic encoding, and explicit retrieval was facilitated when given more study time. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Brain regions supporting intentional and incidental memory: a PET study

Regional brain activity associated with intentional and incidental memory retrieval was studied with PET. Previously studied and new words were presented in either an intentional or an incidental memory task. Type of task was crossed with an encoding manipulation ('deep' vs 'shallow') which varied the probability that studied items would be remembered. In both tasks, deeply encoded items were associated with greater activation in the left hippocampus than were items that had received shallow encoding, suggesting that the involvement of the hippocampus in memory retrieval is independent of whether remembering is intentional or incidental. Right prefrontal and bilateral parietal cortex were more activated during the international task than during the incidental task, irrespective of encoding condition. Thus, these regions play a more extensive role in memory retrieval when remembering is intentional.     

Effects of aging on the neural correlates of successful item and source memory encoding.

To investigate the neural basis of age-related source memory (SM) deficits, young and older adults were scanned with fMRI while encoding faces, scenes, and face-scene pairs. Successful encoding activity was identified by comparing encoding activity for subsequently remembered versus forgotten items or pairs. Age deficits in successful encoding activity in hippocampal and prefrontal regions were more pronounced for SM (pairs) as compared with item memory (faces and scenes). Age-related reductions were also found in regions specialized in processing faces (fusiform face area) and scenes (parahippocampal place area), but these reductions were similar for item and SM. Functional connectivity between the hippocampus and the rest of the brain was also affected by aging; whereas connections with posterior cortices were weaker in older adults, connections with anterior cortices, including prefrontal regions, were stronger in older adults. Taken together, the results provide a link between SM deficits in older adults and reduced recruitment of hippocampal and prefrontal regions during encoding. The functional connectivity findings are consistent with a posterior-anterior shift with aging previously reported in several cognitive domains and linked to functional compensation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Reconciling findings of emotion-induced memory enhancement and impairment of preceding items.

A large body of work has revealed that people remember emotionally arousing information better than neutral information. However, previous research has revealed contradictory effects of emotional events on memory for neutral events that precede or follow them: In some studies, emotionally arousing items have impaired memory for immediately preceding or following items, and in others arousing items enhanced memory for preceding items. By demonstrating both emotion-induced enhancement and impairment, Experiments 1 and 2 clarified the conditions under which these effects are likely to occur. The results suggest that emotion-induced enhancement is most likely to occur for neutral items that (a) precede (and so are poised to predict the onset of) emotionally arousing items, (b) have high attentional weights at encoding, and (c) are tested after a delay period of a week rather than within the same experimental session. In contrast, emotion-induced impairment is most likely to occur for neutral items near the onset of emotional arousal that are overshadowed by highly activated competing items during encoding. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

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. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Interactive Visual and Postvisual Processes and Their Roles in Form-Specific Memory.

Effects of depth of encoding on form-specific memory were examined. After viewing words (e.g., "bear") presented centrally during initial encoding, participants completed word stems (e.g., "BEA") presented laterally and pattern masked during subsequent test. When the encoding task was perceptual, letter-case specific memory was not observed, unlike in previous experiments without pattern masking. However, when the encoding task required both perceptual and conceptual processing, letter-case specific memory was observed in direct right-hemisphere, but not in direct left-hemisphere, test presentations, like in previous studies without pattern masking. Results were not influenced by whether stems were completed to form the first words that came to mind or words explicitly retrieved from encoding. Depth of encoding may influence form-specific memory through interactive processing of visual and postvisual information. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Dissociable effects of conscious emotion regulation strategies on explicit and implicit memory.

The authors manipulated emotion regulation strategies at encoding and administered explicit and implicit memory tests. In Experiment 1, participants used reappraisal to enhance and decrease the personal relevance of unpleasant and neutral pictures. In Experiment 2, decrease cues were replaced with suppress cues that directed participants to inhibit emotion-expressive behavior. Across experiments, using reappraisal to enhance the personal relevance of pictures improved free recall. By contrast, attempting to suppress emotional displays tended to impair recall, especially compared to the enhance condition. Using reappraisal to decrease the personal relevance of pictures had different effects depending on picture type. Paired with unpleasant pictures, the decrease cue tended to improve recall. Paired with neutral stimuli, the decrease cue tended to impair recall. Emotion regulation did not affect perceptual priming. Results highlight dissociable effects of emotion regulation on explicit and implicit memory, as well as dissociations between regulation strategies with respect to explicit memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The associative memory deficit of older adults: The role of strategy utilization.

Past research has established an associative deficit hypothesis (e.g., M. Naveh-Benjamin, 2000) that attributes part of older adults' poor episodic memory performance to their difficulty in creating and retrieving cohesive episodes. Here, the authors evaluate the degree to which this deficit can be reduced by older adults' use of associative strategies. Young and older adults learned word pairs under either intentional-learning instructions or instructions eliciting associative strategies either at encoding or both at encoding and at retrieval, and they then took tests on their memory for both the components and the associations. Results replicated the associative deficit of older adults under intentional-learning instructions. In addition, they showed that instructions to use appropriate associative strategies during either encoding or, even more so, during encoding and retrieval resulted in a significant decrease in the associative deficit. The theoretical and applied implications of these results are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Adult age differences in episodic memory: Further support for an associative-deficit hypothesis.

This study further tested an associative-deficit hypothesis (ADH; M. Naveh-Benjamin, 2000), which attributes a substantial part of older adults' deficient episodic memory performance to their difficulty in merging unrelated attributes-units of an episode into a cohesive unit. First, the results of 2 experiments replicate those observed by M. Naveh-Benjamin (2000) showing that older adults are particularly deficient in memory tests requiring associations. Second, the results extend the type of stimuli (pictures) under which older adults show this associative deficit. Third, the results support an ADH in that older adults show less of an associative deficit when the components of the episodes used are already connected in memory, thereby facilitating their encoding and retrieval. Finally, a group of younger adults who encoded the information under divided-attention conditions did not show this associative deficit. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Content-specific source encoding in the human medial temporal lobe.

Although the medial temporal lobe (MTL) is known to be essential for episodic encoding, the contributions of individual MTL subregions remain unclear. Data from recognition memory studies have provided evidence that the hippocampus supports relational encoding important for later episodic recollection, whereas the perirhinal cortex has been linked with encoding that supports later item familiarity. However, extant data also strongly implicate the perirhinal cortex in object processing and encoding, suggesting that perirhinal processes may contribute to later episodic recollection of object source details. To investigate this possibility, encoding activation in MTL subregions was analyzed on the basis of subsequent memory outcome while participants processed novel scenes paired with 1 of 6 repeating objects. Specifically, encoding activation correlating with later successful scene recognition memory was evaluated against that of source recollection for the object paired with the scene during encoding. In contrast to studies reporting a link between perirhinal cortex and item familiarity, it was found that encoding activation in the right perirhinal cortex correlates with successful recollection of the paired object. Furthermore, other MTL subregions also exhibited content-specific source encoding patterns of activation, suggesting that MTL subsequent memory effects are sensitive to stimulus category. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Using what we know: Consequences of intentionally retrieving gist versus item-specific information.

The effect of a prior gist-based versus item-specific retrieval orientation on recognition of objects and words was examined. Prior item-specific retrieval increased item-specific recognition of episodically related but not previously tested objects relative to both conceptual- and perceptual-gist retrieval. An item-specific retrieval advantage also was found when the stimuli were words (synonyms) rather than objects but not when participants overtly named objects during gist-based recognition testing, which suggests that they did not always label objects under general gist-retrieval instructions. Unlike verbal overshadowing, labeling objects during recognition attenuated (but did not eliminate) test- and interference-related forgetting. A full understanding of how retrieval affects subsequent memory, even for events or facts that are not themselves retrieved, must take into account the specificity with which that retrieval occurs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)