Event-related Potentials of Emotional Memory: Encoding Pleasant, Unpleasant, and Neutral Pictures

Cognitive, Affective & Behavioral Neuroscience. Sep, 2002  |  Pubmed ID: 12775189

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.

Hemispheric Asymmetry Reduction in Older Adults: the HAROLD Model

Psychology and Aging. Mar, 2002  |  Pubmed ID: 11931290

A model of the effects of aging on brain activity during cognitive performance is introduced. The model is called HAROLD (hemispheric asymmetry reduction in older adults), and it states that, under similar circumstances, prefrontal activity during cognitive performances tends to be less lateralized in older adults than in younger adults. The model is supported by functional neuroimaging and other evidence in the domains of episodic memory, semantic memory, working memory, perception, and inhibitory control. Age-related hemispheric asymmetry reductions may have a compensatory function or they may reflect a dedifferentiation process. They may have a cognitive or neural origin, and they may reflect regional or network mechanisms. The HAROLD model is a cognitive neuroscience model that integrates ideas and findings from psychology and neuroscience of aging.

Brain Imaging of Human Memory Systems: Between-systems Similarities and Within-system Differences

Brain Research. Cognitive Brain Research. Apr, 2002  |  Pubmed ID: 11958972

There is much evidence for the existence of multiple memory systems. However, it has been argued that tasks assumed to reflect different memory systems share basic processing components and are mediated by overlapping neural systems. Here we used multivariate analysis of PET-data to analyze similarities and differences in brain activity for multiple tests of working memory, semantic memory, and episodic memory. The results from two experiments revealed between-systems differences, but also between-systems similarities and within-system differences. Specifically, support was obtained for a task-general working-memory network that may underlie active maintenance. Premotor and parietal regions were salient components of this network. A common network was also identified for two episodic tasks, cued recall and recognition, but not for a test of autobiographical memory. This network involved regions in right inferior and polar frontal cortex, and lateral and medial parietal cortex. Several of these regions were also engaged during the working-memory tasks, indicating shared processing for episodic and working memory. Fact retrieval and synonym generation were associated with increased activity in left inferior frontal and middle temporal regions and right cerebellum. This network was also associated with the autobiographical task, but not with living/non-living classification, and may reflect elaborate retrieval of semantic information. Implications of the present results for the classification of memory tasks with respect to systems and/or processes are discussed.

Similarities and Differences in the Neural Correlates of Episodic Memory Retrieval and Working Memory

NeuroImage. Jun, 2002  |  Pubmed ID: 12030819

Functional neuroimaging studies have shown that different cognitive functions activate overlapping brain regions. An activation overlap may occur because a region is involved in operations tapped by different cognitive functions or because the activated area comprises subregions differentially involved in each of the functions. To investigate these issues, we directly compared brain activity during episodic retrieval (ER) and working memory (WM) using event-related functional MRI (fMRI). ER was investigated with a word recognition test, and WM was investigated with a word delayed-response test. Two-phase trials distinguished between retrieval mode and cue-specific aspects of ER, as well as between encoding/maintenance and retrieval aspects of WM. The results revealed a common fronto-parieto-cerebellar network for ER and WM, as well as subregions differentially involved in each function. Specifically, there were two main findings. First, the results differentiated common and specific subregions within the prefrontal cortex: (i) left dorsolateral areas were recruited by both functions, possibly reflecting monitoring operations; (ii) bilateral anterior and ventrolateral areas were more activated during ER than during WM, possibly reflecting retrieval mode and cue-specific ER operations, respectively; and (iii) left posterior/ventral (Broca's area) and bilateral posterior/dorsal areas were more activated during WM than during ER, possibly reflecting phonological and generic WM operations, respectively. Second, hippocampal and parahippocampal regions were activated not only for ER but also for WM. This result suggests that indexing operations mediated by the medial temporal lobes apply to both long-term and short-term memory traces. Overall, our results show that direct cross-function comparisons are critical to understand the role of different brain regions in various cognitive functions.

Aging Gracefully: Compensatory Brain Activity in High-performing Older Adults

NeuroImage. Nov, 2002  |  Pubmed ID: 12414279

Whereas some older adults show significant cognitive deficits, others perform as well as young adults. We investigated the neural basis of these different aging patterns using positron emission tomography (PET). In PET and functional MRI (fMRI) studies, prefrontal cortex (PFC) activity tends to be less asymmetric in older than in younger adults (Hemispheric Asymmetry Reduction in Old Adults or HAROLD). This change may help counteract age-related neurocognitive decline (compensation hypothesis) or it may reflect an age-related difficulty in recruiting specialized neural mechanisms (dedifferentiation hypothesis). To compare these two hypotheses, we measured PFC activity in younger adults, low-performing older adults, and high-performing older adults during recall and source memory of recently studied words. Compared to recall, source memory was associated with right PFC activations in younger adults. Low-performing older adults recruited similar right PFC regions as young adults, but high-performing older adults engaged PFC regions bilaterally. Thus, consistent with the compensation hypothesis and inconsistent with the dedifferentiation hypothesis, a hemispheric asymmetry reduction was found in high-performing but not in low-performing older adults. The results suggest that low-performing older adults recruited a similar network as young adults but used it inefficiently, whereas high-performing older adults counteracted age-related neural decline through a plastic reorganization of neurocognitive networks.

Hemispheric Asymmetry and Aging: Right Hemisphere Decline or Asymmetry Reduction

Neuroscience and Biobehavioral Reviews. Nov, 2002  |  Pubmed ID: 12470693

We review evidence for two models of hemispheric asymmetry and aging: the right hemi-aging model, which proposes that the right hemisphere shows greater age-related decline than the left hemisphere, and the hemispheric asymmetry reduction in old adults (HAROLD) model, which proposes that frontal activity during cognitive performance tends to be less lateralized in older than in younger adults. The right hemi-aging model is supported by behavioral studies in the domains of cognitive, affective, and sensorimotor processing, but the evidence has been mixed. In contrast, available evidence is generally consistent with the HAROLD model, which is supported primarily by functional neuroimaging evidence in the domains of episodic memory encoding and retrieval, semantic memory retrieval, working memory, perception, and inhibitory control. Age-related asymmetry reductions may reflect functional compensation or dedifferentiation, and the evidence, although scarce, tends to support the compensation hypothesis. The right hemi-aging and the HAROLD models are not incompatible. For example, the latter may apply to prefrontal regions and the former to other brain regions.

Lateralization of Prefrontal Activity During Episodic Memory Retrieval: Evidence for the Production-monitoring Hypothesis

Journal of Cognitive Neuroscience. Feb, 2003  |  Pubmed ID: 12676062

We propose a new hypothesis concerning the lateralization of prefrontal cortex (PFC) activity during verbal episodic memory retrieval. The hypothesis states that the left PFC is differentially more involved in semantically guided information production than is the right PFC, and that the right PFC is differentially more involved in monitoring and verification than is the left PFC. This "production-monitoring hypothesis" differs from the existing "systematic-heuristic hypothesis," which proposes that the left PFC is primarily involved in systematic retrieval operations, and the right PFC in heuristic retrieval operations. To compare the two hypotheses, we measured PFC activity using positron emission tomography (PET) during the performance of four episodic retrieval tasks: stem cued recall, associative cued recall, context recognition (source memory), and item recognition. Recall tasks emphasized production processes, whereas recognition tasks emphasized monitoring processes. Stem cued recall and context-recognition tasks underscored systematic operations, whereas associative cued recall and item-recognition tasks underscored heuristic operations. Consistent with the production-monitoring hypothesis, the left PFC was more activated for recall than for recognition tasks and the right PFC was more activated for recognition than for recall tasks. Inconsistent with the systematic-heuristic hypothesis, the left PFC was more activated for heuristic than for systematic tasks and the right PFC showed the converse result. Additionally, the study yielded activation differences outside the PFC. In agreement with a previous recall/recognition PET study, anterior cingulate, cerebellar, and striatal regions were more activated for recall than for recognition tasks, and the converse occurred for posterior parietal regions. A right medial temporal lobe region was more activated for stem cued recall and context recognition than for associative cued recall and item recognition, possibly reflecting perceptual integration. In sum, the results provide evidence for the production-monitoring hypothesis and clarify the role of different brain regions typically activated in PET and functional magnetic resonance imaging (fMRI) studies of episodic retrieval.

Functional Neuroimaging of Memory

Neuropsychologia. 2003  |  Pubmed ID: 12457749

Common Prefrontal Activations During Working Memory, Episodic Memory, and Semantic Memory

Neuropsychologia. 2003  |  Pubmed ID: 12457761

Regions of the prefrontal cortex (PFC) are typically activated in many different cognitive functions. In most studies, the focus has been on the role of specific PFC regions in specific cognitive domains, but more recently similarities in PFC activations across cognitive domains have been stressed. Such similarities may suggest that a region mediates a common function across a variety of cognitive tasks. In this study, we compared the activation patterns associated with tests of working memory, semantic memory and episodic memory. The results converged on a general involvement of four regions across memory tests. These were located in left frontopolar cortex, left mid-ventrolateral PFC, left mid-dorsolateral PFC and dorsal anterior cingulate cortex. These findings provide evidence that some PFC regions are engaged during many different memory tests. The findings are discussed in relation to theories about the functional contribution of the PFC regions and the architecture of memory.

Attention-related Activity During Episodic Memory Retrieval: a Cross-function FMRI Study

Neuropsychologia. 2003  |  Pubmed ID: 12457763

In functional neuroimaging studies of episodic retrieval (ER), activations in prefrontal, parietal, anterior cingulate, and thalamic regions are typically attributed to episodic retrieval processes. However, these activations are also frequent during visual attention (VA) tasks, suggesting that their role in ER may reflect attentional rather than mnemonic processes. To investigate this possibility, we directly compared brain activity during ER and VA tasks using event-related fMRI. The ER task was a word recognition test with a retrieval mode component, and the VA task was a target detection task with a sustained attention component. The study yielded three main findings. First, a common fronto-parietal-cingulate-thalamic network was found for ER and VA, suggesting that the involvement of these regions during ER reflects general attentional processes. This idea is compatible with some of the interpretations proposed in the ER literature (e.g. postretrieval monitoring), which may be rephrased in terms of attentional processes. Second, several subregions were differentially involved in ER versus VA. For example, the frontopolar cortex and the precuneus were more activated for ER than for VA, possibly reflecting retrieval mode and processing of internally generated stimuli, respectively. Finally, the study yielded an unexpected finding: some medial temporal lobe regions were similarly activated for ER and VA. This finding suggests that the medial temporal lobes may be involved in indexing representations within the focus of consciousness, regardless of whether they are mnemonic or perceptual. Overall, the present results suggest that many of the activations attributed to specific cognitive processes, such as episodic memory, may actually reflect more general cognitive operations.

Task-independent and Task-specific Age Effects on Brain Activity During Working Memory, Visual Attention and Episodic Retrieval

Cerebral Cortex (New York, N.Y. : 1991). Apr, 2004  |  Pubmed ID: 15028641

It is controversial whether the effects of aging on various cognitive functions have the same common cause or several different causes. To investigate this issue, we scanned younger and older adults with functional magnetic resonance imaging (fMRI) while performing three different tasks: working memory, visual attention and episodic retrieval. There were three main results. First, in all three tasks, older adults showed weaker occipital activity and stronger prefrontal and parietal activity than younger adults. The occipital reduction is consistent with the view that sensory processing decline is a common cause in cognitive aging, and the prefrontal increase may reflect functional compensation. Secondly, older adults showed more bilateral patterns of prefrontal activity than younger adults during working memory and visual attention tasks. These findings are consistent with the Hemispheric Asymmetry Reduction in Older Adults (HAROLD) model. Finally, compared to younger adults, older adults showed weaker hippocampal formation activity in all three tasks but stronger parahippocampal activity in the episodic retrieval task. The former finding suggests that age-related hippocampal deficits may have a global effect in cognition, and the latter is consistent with an age-related increase in familiarity-based recognition. Taken together, the results indicate that both common and specific factors play an important role in cognitive aging.

Interaction Between the Amygdala and the Medial Temporal Lobe Memory System Predicts Better Memory for Emotional Events

Neuron. Jun, 2004  |  Pubmed ID: 15182723

Emotional events are remembered better than neutral events possibly because the amygdala enhances the function of medial temporal lobe (MTL) memory system (modulation hypothesis). Although this hypothesis has been supported by much animal research, evidence from humans has been scarce and indirect. We investigated this issue using event-related fMRI during encoding of emotional and neutral pictures. Memory performance after scanning showed a retention advantage for emotional pictures. Successful encoding activity in the amygdala and MTL memory structures was greater and more strongly correlated for emotional than for neutral pictures. Moreover, a double dissociation was found along the longitudinal axis of the MTL memory system: activity in anterior regions predicted memory for emotional items, whereas activity in posterior regions predicted memory for neutral items. These results provide direct evidence for the modulation hypothesis in humans and reveal a functional specialization within the MTL regarding the effects of emotion on memory formation.

Age-related Preservation of Top-down Attentional Guidance During Visual Search

Psychology and Aging. Jun, 2004  |  Pubmed ID: 15222823

Younger (19-27 years of age) and older (60-82 years of age) adults performed a letter search task in which a color singleton was either noninformative (baseline condition) or highly informative (guided condition) regarding target location. In the guided condition, both age groups exhibited a substantial decrease in response time (RT) to singleton targets, relative to the baseline condition, as well as an increase in RT to nonsingleton targets. The authors conclude that under conditions that equate the physical structure of individual displays, top-down attentional guidance can be at least as effective for older adults as for younger adults.

Dissociable Effects of Arousal and Valence on Prefrontal Activity Indexing Emotional Evaluation and Subsequent Memory: an Event-related FMRI Study

NeuroImage. Sep, 2004  |  Pubmed ID: 15325353

Prefrontal cortex (PFC) activity associated with emotional evaluation and subsequent memory was investigated with event-related functional MRI (fMRI). Participants were scanned while rating the pleasantness of emotionally positive, negative, and neutral pictures, and memory for the pictures was tested after scanning. Emotional evaluation was measured by comparing activity during the picture rating task relative to baseline, and successful encoding was measured by comparing activity for subsequently remembered versus forgotten pictures (Dm effect). The effect of arousal on these measures was indicated by greater activity for both positive and negative pictures than for neutral ones, and the effect of valence was indicated by differences in activity between positive and negative pictures. The study yielded three main results. First, consistent with the valence hypothesis, specific regions in left dorsolateral PFC were more activated for positive than for negative picture evaluation, whereas regions in right ventrolateral PFC showed the converse pattern. Second, dorsomedial PFC activity was sensitive to emotional arousal, whereas ventromedial PFC activity was sensitive to positive valence, consistent with evidence linking these regions, respectively, to emotional processing and self-awareness or appetitive behavior. Finally, successful encoding (Dm) activity in left ventrolateral and dorsolateral PFC was greater for arousing than for neutral pictures. This finding suggests that the enhancing effect of emotion on memory formation is partly due to an augmentation of PFC-mediated strategic, semantic, and working memory operations. These results underscore the critical role of PFC in emotional evaluation and memory, and disentangle the effects of arousal and valence across PFC regions associated with different cognitive functions.

Brain Activity During Episodic Retrieval of Autobiographical and Laboratory Events: an FMRI Study Using a Novel Photo Paradigm

Journal of Cognitive Neuroscience. Nov, 2004  |  Pubmed ID: 15622612

Functional neuroimaging studies of episodic memory retrieval generally measure brain activity while participants remember items encountered in the laboratory ("controlled laboratory condition") or events from their own life ("open autobiographical condition"). Differences in activation between these conditions may reflect differences in retrieval processes, memory remoteness, emotional content, retrieval success, self-referential processing, visual/spatial memory, and recollection. To clarify the nature of these differences, a functional MRI study was conducted using a novel "photo paradigm," which allows greater control over the autobiographical condition, including a measure of retrieval accuracy. Undergraduate students took photos in specified campus locations ("controlled autobiographical condition"), viewed in the laboratory similar photos taken by other participants (controlled laboratory condition), and were then scanned while recognizing the two kinds of photos. Both conditions activated a common episodic memory network that included medial temporal and prefrontal regions. Compared with the controlled laboratory condition, the controlled autobiographical condition elicited greater activity in regions associated with self-referential processing (medial prefrontal cortex), visual/spatial memory (visual and parahippocampal regions), and recollection (hippocampus). The photo paradigm provides a way of investigating the functional neuroanatomy of real-life episodic memory under rigorous experimental control.

Neural Correlates of Relational Memory: Successful Encoding and Retrieval of Semantic and Perceptual Associations

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. Feb, 2005  |  Pubmed ID: 15689557

Using event-related functional magnetic resonance imaging, we identified brain regions involved in successful relational memory (RM) during encoding and retrieval for semantic and perceptual associations or in general, independent of phase and content. Participants were scanned while encoding and later retrieving associations between pairs of words (semantic RM) or associations between words and fonts (perceptual RM). Encoding success activity (ESA) was identified by comparing study-phase activity for items subsequently remembered (hits) versus forgotten (misses) and retrieval success activity (RSA) by comparing test-phase activity for hits versus misses. The study yielded three main sets of findings. First, ESA-RSA differences were found within the medial temporal lobes (MTLs) and within the prefrontal cortex (PFC). Within the left MTL, ESA was greater in the anterior hippocampus, and RSA was greater in the posterior parahippocampal cortex/hippocampus. This finding is consistent with the notion of an encoding-retrieval gradient along the longitudinal MTL axis. Within the left PFC, ESA was greater in ventrolateral PFC, and RSA was greater in dorsolateral and anterior PFC. This is the first evidence of a dissociation in successful encoding and retrieval activity within left PFC. Second, consistent with the transfer-appropriate processing principle, some ESA regions were reactivated during RSA in a content-specific manner. For semantic RM, these regions included the left ventrolateral PFC, whereas for perceptual RM, they included occipitoparietal and right parahippocampal regions. Finally, only one region in the entire brain was associated with RM in general (i.e., for both semantic and perceptual ESA and RSA): the left hippocampus. This finding highlights the fundamental role of the hippocampus in RM.

Remembering One Year Later: Role of the Amygdala and the Medial Temporal Lobe Memory System in Retrieving Emotional Memories

Proceedings of the National Academy of Sciences of the United States of America. Feb, 2005  |  Pubmed ID: 15703295

The memory-enhancing effect of emotion can be powerful and long-lasting. Most studies investigating the neural bases of this phenomenon have focused on encoding and early consolidation processes, and hence little is known regarding the contribution of retrieval processes, particularly after lengthy retention intervals. To address this issue, we used event-related functional MRI to measure neural activity during the retrieval of emotional and neutral pictures after a retention interval of 1 yr. Retrieval activity for emotional and neutral pictures was separately analyzed for successfully (hits) vs. unsuccessfully (misses) retrieved items and for responses based on recollection vs. familiarity. Recognition performance was better for emotional than for neutral pictures, and this effect was found only for recollection-based responses. Successful retrieval of emotional pictures elicited greater activity than successful retrieval of neutral pictures in the amygdala, entorhinal cortex, and hippocampus. Moreover, in the amygdala and hippocampus, the emotion effect was greater for recollection than for familiarity, whereas in the entorhinal cortex, it was similar for both forms of retrieval. These findings clarify the role of the amygdala and the medial temporal lobe memory regions in recollection and familiarity of emotional memory after lengthy retention intervals.

Co-activation of the Amygdala, Hippocampus and Inferior Frontal Gyrus During Autobiographical Memory Retrieval

Neuropsychologia. 2005  |  Pubmed ID: 15721179

Functional MRI was used to investigate the role of medial temporal lobe and inferior frontal lobe regions in autobiographical recall. Prior to scanning, participants generated cue words for 50 autobiographical memories and rated their phenomenological properties using our autobiographical memory questionnaire (AMQ). During scanning, the cue words were presented and participants pressed a button when they retrieved the associated memory. The autobiographical retrieval task was interleaved in an event-related design with a semantic retrieval task (category generation). Region-of-interest analyses showed greater activation of the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval relative to semantic retrieval. In addition, the left inferior frontal gyrus showed a more prolonged duration of activation in the semantic retrieval condition. A targeted correlational analysis revealed pronounced functional connectivity among the amygdala, hippocampus, and right inferior frontal gyrus during autobiographical retrieval but not during semantic retrieval. These results support theories of autobiographical memory that hypothesize co-activation of frontotemporal areas during recollection of episodes from the personal past.

False Memory Across Languages: Implicit Associative Response Vs Fuzzy Trace Views

Memory (Hove, England). Jan, 2005  |  Pubmed ID: 15724903

We investigated false recognition across languages using the Deese-Roediger-McDermott (DRM) paradigm. A group of English-French bilinguals studied lists of converging associates, some lists in English and some in French, and then performed a recognition test containing studied list items and nonstudied critical lures whose language matched or mismatched the language at study. Participants were instructed to answer old only if the test cue was in the same language as the studied word. The results yielded a robust false memory rate both within-language and across-languages. The effect of the study-test language shift was much larger for list items than for critical lures. This finding suggests that memory representations for critical lures contain primarily semantic gist traces and little surface information, and hence is more consistent with the fuzzy trace view than with the implicit associative response view. In sum, the study demonstrates the existence of false memory across languages, and provides information about the memory traces underlying veridical and illusory recognition.

Cognitive Neuroscience of Emotional Memory

Nature Reviews. Neuroscience. Jan, 2006  |  Pubmed ID: 16371950

Emotional events often attain a privileged status in memory. Cognitive neuroscientists have begun to elucidate the psychological and neural mechanisms underlying emotional retention advantages in the human brain. The amygdala is a brain structure that directly mediates aspects of emotional learning and facilitates memory operations in other regions, including the hippocampus and prefrontal cortex. Emotion-memory interactions occur at various stages of information processing, from the initial encoding and consolidation of memory traces to their long-term retrieval. Recent advances are revealing new insights into the reactivation of latent emotional associations and the recollection of personal episodes from the remote past.

Role of Prefrontal and Anterior Cingulate Regions in Decision-making Processes Shared by Memory and Nonmemory Tasks

Cerebral Cortex (New York, N.Y. : 1991). Nov, 2006  |  Pubmed ID: 16400154

In the episodic retrieval (ER) domain, activations in right dorsolateral prefrontal cortex (DLPFC) are often attributed to postretrieval monitoring. Yet, right DLPFC activations are also frequently found during nonmemory tasks. To investigate the role of this region across different cognitive functions, we directly compared brain activity during ER and visual perception (VP) using event-related functional magnetic resonance imaging. In the ER task, participants decided whether words were old or new, whereas in the VP task, they decided which of the two colored screen areas was larger. In both tasks, each decision was followed by a confidence rating. The main finding was that right DLPFC (Brodmann area 46/10) activity was greater for low- than for high-confidence decisions in both tasks, demonstrating a general role in decision making. Even when reaction times (RTs) were included in the model, confidence remained the significant predictor of activity, suggesting that right DLPFC is involved in discontinuous evaluation rather than in continuous monitoring. In contrast, activity in anterior cingulate cortex was not only greater for low-confidence decisions but also increased with RT, reflecting a role in continuous conflict monitoring. Overall, the results demonstrate how direct cross-function comparisons clarify the generality and specificity of the functions of various brain regions.

Effects of Healthy Aging on Hippocampal and Rhinal Memory Functions: an Event-related FMRI Study

Cerebral Cortex (New York, N.Y. : 1991). Dec, 2006  |  Pubmed ID: 16421332

Event-related functional magnetic resonance imaging was used to study the effects of healthy aging on hippocampal and rhinal memory functions. Memory for past events can be based on retrieval accompanied by specific contextual details (recollection) or on the feeling that an event is old or new without the recovery of contextual details (familiarity). There is evidence that recollection is more dependent on hippocampus, whereas familiarity is more dependent on the rhinal cortex, and that healthy aging has greater effects on recollection than on familiarity. However, little evidence is available about the neural correlates of these effects. Here, we isolated activity associated with recollection and familiarity by distinguishing between linear and quasi-exponential "perceived oldness" functions derived from recognition confidence levels. The main finding was a double dissociation within the medial temporal lobes between recollection-related activity in hippocampus, which was reduced by aging, and familiarity-related activity in rhinal cortex, which was increased by aging. In addition, age dissociations were found within parietal and posterior midline regions. Finally, aging reduced functional connectivity within a hippocampal-retrosplenial/parietotemporal network but increased connectivity within a rhinal-frontal network. These findings indicate that older adults compensate for hippocampal deficits by relying more on rhinal cortex, possibly through a top-down frontal modulation. This finding has important clinical implications because early Alzheimer's disease impairs both hippocampus and rhinal cortex.

The Medial Temporal Lobe Distinguishes Old from New Independently of Consciousness

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. May, 2006  |  Pubmed ID: 16723542

Although it is widely accepted that the medial temporal lobes (MTLs) are critical for becoming aware that something happened in the past, there is virtually no evidence whether MTL sensitivity to event oldness also depends on conscious awareness. Using event-related functional magnetic resonance imaging, we show that activity in posterior MTL tracks whether an item is actually old (true oldness), regardless of participants' awareness of oldness (perceived oldness). Confirming its sensitivity to the objective nature of the stimulus, activity in this region was strongly correlated with individual memory performance (r = 0.74). At the same time, we found that memory errors (misses) were associated with activity in an anterior MTL region, which signaled whether an item was consciously experienced as new (perceived novelty). Logistic regression analyses based on individual trial activity indicated that the two MTL regions showed opposing relationships with behavior, and that memory performance was determined by their joint activity. Furthermore, functional connectivity analyses showed that perceived novelty activity in the ANTERIOR [corrected] MTL inhibited true oldness activity in the POSTERIOR [corrected] MTL. These findings indicate that participants' behavior reflected the combined effects of multiple MTL regions. More generally, our results show that parts of MTL can distinguish old from new independently of consciousness.

Effects of Aging on Transient and Sustained Successful Memory Encoding Activity

Neurobiology of Aging. Nov, 2007  |  Pubmed ID: 16919850

Event-related fMRI studies have investigated age-related changes in encoding by identifying greater activity for items that are later remembered than for those that are forgotten (difference in memory, or Dm). The present study used hybrid blocked/event-related analyses to distinguish between transient Dm versus sustained Dm. Dm was identified as parametric increases in encoding activity as a function of a combined subsequent memory/confidence scale. Dm was measured in each trial (transient activity) and in blocks of eight trials (sustained activity). Transient Dm analyses showed age-related reductions in the left hippocampus but increases in left prefrontal cortex (PFC). Sustained Dm analyses showed age-related reductions in right PFC, but no region showing increased activity in older adults. These findings suggests that during semantic classification older adults show less spontaneous hippocampal-mediated encoding processes, but greater PFC-mediated semantic processes. Additionally, the decline in sustained Dm in PFC may involve age-related deficits in sustained attention that impact encoding processes. The results underscore the importance of investigating aging effects on both transient and sustained neural activity.

Differential Contributions of Prefrontal, Medial Temporal, and Sensory-perceptual Regions to True and False Memory Formation

Cerebral Cortex (New York, N.Y. : 1991). Sep, 2007  |  Pubmed ID: 17110592

The neural correlates of true memory formation (TMF) and false memory formation (FMF) were investigated using functional magnetic resonance imaging (fMRI). Using a parametric subsequent memory paradigm, encoding activity was analyzed as a function of whether it predicted subsequent hits to targets (TMF activity) or subsequent false alarms to critical lures (FMF activity). The fMRI analyses yielded 3 main findings. First, the left prefrontal cortex (PFC) was involved in both TMF and FMF activities. This finding is consistent with the evidence that semantic elaboration, which has been associated with left PFC, tends to enhance both true and false remembering. Second, the left posterior medial temporal lobes (MTLs) contributed to TMF but not to FMF activity. This finding is consistent with the notion that MTL is involved in the storage of a consciously, but not unconsciously, processed event. Third, late visual regions were engaged in both TMF and FMF activities, whereas early visual areas were involved primarily in TMF activity. This dissociation indicates that elaborative perceptual processing, but not basic sensory processing, contributes to false remembering. Taken together, the results suggest that FMF is an unintended consequence, or by-product, of elaborative semantic and visual encoding processes.

Sustained and Transient Neural Modulations in Prefrontal Cortex Related to Declarative Long-term Memory, Working Memory, and Attention

Cortex; a Journal Devoted to the Study of the Nervous System and Behavior. Jan, 2007  |  Pubmed ID: 17334205

Common activations in prefrontal cortex (PFC) during episodic and semantic long-term memory (LTM) tasks have been hypothesized to reflect functional overlap in terms of working memory (WM) and cognitive control. To evaluate a WM account of LTM-general activations, the present study took into consideration that cognitive task performance depends on the dynamic operation of multiple component processes, some of which are stimulus-synchronous and transient in nature; and some that are engaged throughout a task in a sustained fashion. PFC and WM may be implicated in both of these temporally independent components. To elucidate these possibilities we employed mixed blocked/event-related functional magnetic resonance imaging (fMRI) procedures to assess the extent to which sustained or transient activation patterns overlapped across tasks indexing episodic and semantic LTM, attention (ATT), and WM. Within PFC, ventrolateral and medial areas exhibited sustained activity across all tasks, whereas more anterior regions including right frontopolar cortex were commonly engaged in sustained processing during the three memory tasks. These findings do not support a WM account of sustained frontal responses during LTM tasks, but instead suggest that the pattern that was common to all tasks reflects general attentional set/vigilance, and that the shared WM-LTM pattern mediates control processes related to upholding task set. Transient responses during the three memory tasks were assessed relative to ATT to isolate item-specific mnemonic processes and were found to be largely distinct from sustained effects. Task-specific effects were observed for each memory task. In addition, a common item response for all memory tasks involved left dorsolateral PFC (DLPFC). The latter response might be seen as reflecting WM processes during LTM retrieval. Thus, our findings suggest that a WM account of shared PFC recruitment in LTM tasks holds for common transient item-related responses rather than sustained state-related responses that are better seen as reflecting more general attentional/control processes.

Functional Neuroimaging of Autobiographical Memory

Trends in Cognitive Sciences. May, 2007  |  Pubmed ID: 17382578

Functional neuroimaging studies of autobiographical memory have grown dramatically in recent years. These studies are important because they can investigate the neural correlates of processes that are difficult to study using laboratory stimuli, including: (i) complex constructive processes, (ii) recollective qualities of emotion and vividness, and (iii) remote memory retrieval. Constructing autobiographical memories involves search, monitoring and self-referential processes that are associated with activity in separable prefrontal regions. The contributions of emotion and vividness have been linked to the amygdala and visual cortex respectively. Finally, there is evidence that recent and remote autobiographical memories might activate the hippocampus equally, which has implications for memory-consolidation theories. The rapid development of innovative methods for eliciting personal memories in the scanner provides the opportunity to delve into the functional neuroanatomy of our personal past.

Adult Age Differences in the Functional Neuroanatomy of Visual Attention: a Combined FMRI and DTI Study

Neurobiology of Aging. Mar, 2007  |  Pubmed ID: 16500004

We combined measures from event-related functional magnetic resonance imaging (fMRI), diffusion tensor imaging (DTI), and cognitive performance (visual search response time) to test the hypotheses that differences between younger and older adults in top-down (goal-directed) attention would be related to cortical activation, and that white matter integrity as measured by DTI (fractional anisotropy, FA) would be a mediator of this age-related effect. Activation in frontal and parietal cortical regions was overall greater for older adults than for younger adults. The relation between activation and search performance supported the hypothesis of age differences in top-down attention. When the task involved top-down control (increased target predictability), performance was associated with frontoparietal activation for older adults, but with occipital (fusiform) activation for younger adults. White matter integrity (FA) exhibited an age-related decline that was more pronounced for anterior brain regions than for posterior regions, but white matter integrity did not specifically mediate the age-related increase in activation of the frontoparietal attentional network.

Trusting Our Memories: Dissociating the Neural Correlates of Confidence in Veridical Versus Illusory Memories

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. Nov, 2007  |  Pubmed ID: 17989285

Although memory confidence and accuracy tend to be positively correlated, people sometimes remember with high confidence events that never happened. How can confidence correlate with accuracy but apply also to illusory memories? One possible explanation is that high confidence in veridical versus illusory memories depends on different neural mechanisms. The present study investigated this possibility using functional magnetic resonance imaging and a modified version of the Deese-Roediger-McDermott false-memory paradigm. Participants read short lists of categorized words, and brain activity was measured while they performed a recognition test with confidence rating. The study yielded three main findings. First, compared with low-confidence responses, high-confidence responses were associated with medial temporal lobe (MTL) activity in the case of true recognition but with frontoparietal activity in the case of false recognition. Second, these regions showed significant confidence-by-veridicality interactions. Finally, only MTL regions showed greater activity for high-confidence true recognition than for high-confidence false recognition, and only frontoparietal regions showed greater activity for high-confidence false recognition than for high-confidence true recognition. These findings indicate that confidence in true recognition is mediated primarily by a recollection-related MTL mechanism, whereas confidence in false recognition reflects mainly a familiarity-related frontoparietal mechanism. This account is consistent with the fuzzy trace theory of false recognition. Correlation analyses revealed that MTL and frontoparietal regions play complementary roles during episodic retrieval. In sum, the present study shows that when one focuses exclusively on high-confidence responses, the neural correlates of true and false memory are clearly different.

Parietal Lobe and Episodic Memory: Bilateral Damage Causes Impaired Free Recall of Autobiographical Memory

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. Dec, 2007  |  Pubmed ID: 18160649

Does the parietal lobe have a critical role in memory? The neuroimaging literature indicates that it has an important role, especially in episodic memory. However, the neuropsychological literature suggests that its role is more limited to attentional, spatial, or imagery aspects of memory. Here, we present data to adjudicate this disagreement. Two patients with bilateral parietal lobe damage received detailed assessments of their autobiographical memories. The results show that although both patients easily recalled various memories, their freely recalled memories were relatively impoverished, lacking in detail. This deficit was ubiquitous, and not limited to spatial or perceptual aspects of memory. The memory deficit disappeared when memory was specifically probed by asking pointed questions. Additional tests show that it is unlikely that their free recall deficit can be explained by general mental imagery problems. In sum, the parietal lobe appears to have a critical role in recollection aspects of episodic memory.

Role of Aerobic Fitness and Aging on Cerebral White Matter Integrity

Annals of the New York Academy of Sciences. Feb, 2007  |  Pubmed ID: 17413020

Neuroimaging research suggests that cerebral white matter (WM) integrity, as reflected in fractional anisotropy (FA) via diffusion tensor imaging (DTI), is decreased in older adults, especially in the prefrontal regions of the brain. Behavioral investigations of cognitive functioning suggest that some aspects of cognition may be better preserved in older adults who possess higher levels of aerobic fitness. There are only a few studies, however, investigating potential mechanisms for the improvements in aerobic fitness. Our study suggests that greater aerobic fitness may be related to greater WM integrity in select brain regions.

Distinguishing the Neural Correlates of Episodic Memory Encoding and Semantic Memory Retrieval

Psychological Science. Feb, 2007  |  Pubmed ID: 17425535

Episodic memory and semantic memory interact very closely. In particular, episodic memory encoding (EE) tends to elicit semantic memory retrieval (SR), and vice versa. Thus, similar activations for EE and SR in functional neuroimaging studies may reflect shared memory processes, or they may reflect the fact that EE and SR are usually confounded. To address this issue, we used a factorial functional magnetic resonance imaging approach to disentangle the neural correlates of EE and SR. Within the left temporal lobe, the hippocampus was associated with successful EE, whereas a posterior lateral region was associated with successful SR. Within the left inferior prefrontal cortex, a posterior region was involved in SR, a mid region was involved in both SR and EE, and an anterior region was involved in EE, but only when SR was also high. Thus, the neural correlates of EE and SR are dissociable but interact in specific brain regions.

Neural Correlates of Promotion and Prevention Goal Activation: an FMRI Study Using an Idiographic Approach

Journal of Cognitive Neuroscience. Jul, 2007  |  Pubmed ID: 17583991

Regulatory focus theory [Higgins, E. T. Beyond pleasure and pain. American Psychologist, 52, 1280-1300, 1997] postulates two social-cognitive motivational systems, the promotion and prevention systems, for self-regulation of goal pursuit. However, the neural substrates of promotion and prevention goal activation remain unclear. Drawing on several literatures, we hypothesized that priming promotion versus prevention goals would activate areas in the left versus right prefrontal cortex (PFC), respectively, and that activation in these areas would be correlated with individual differences in chronic regulatory focus. Sixteen participants underwent functional magnetic resonance imaging while engaged in a depth-of-processing task, during which they were exposed incidentally to their own promotion and prevention goals. Task-related cortical activation was consistent with previous studies. At the same time, incidental priming of promotion goals was associated with left orbital PFC activation, and activation in this area was stronger for individuals with a chronic promotion focus. Findings regarding prevention goal priming were not consistent with predictions. The data illustrate the centrality of self-regulation and personal goal pursuit within the multilayered process of social cognition.

Effects of Aging on True and False Memory Formation: an FMRI Study

Neuropsychologia. Nov, 2007  |  Pubmed ID: 17716696

Compared to young, older adults are more likely to forget events that occurred in the past as well as remember events that never happened. Previous studies examining false memories and aging have shown that these memories are more likely to occur when new items share perceptual or semantic similarities with those presented during encoding. It is theorized that decreased item-specific encoding and increased gist encoding contribute to these age differences in memory performance. The current study used a modified version of the Deese-Roediger-McDermott (DRM) paradigm to investigate the neural correlates of true and false memory encoding. Results indicated that, compared to young, older adults showed reduced activity in medial temporal lobes (MTL), left ventrolateral prefrontal cortex (VLPFC), and visual cortices associated with subsequent true memories. Despite these decreases older adults showed increased activity in right VLPFC and left superior temporal gyrus (STG) for subsequent true memories. Age-related increases in STG were also associated with subsequent false memories. Results support the theory that older adults engage in less item-specific encoding and greater gist encoding, and that these increases in gist encoding support both subsequent true and false memories. Furthermore, results extend findings of reduced frontal asymmetry in aging, often found in block designs, to the subsequent memory paradigm. Results suggest that greater bilateral frontal activity during encoding in aging are not just task-related, but may be associated with subsequent successful memory performance.

Que PASA? The Posterior-anterior Shift in Aging

Cerebral Cortex (New York, N.Y. : 1991). May, 2008  |  Pubmed ID: 17925295

A consistent finding from functional neuroimaging studies of cognitive aging is an age-related reduction in occipital activity coupled with increased frontal activity. This posterior-anterior shift in aging (PASA) has been typically attributed to functional compensation. The present functional magnetic resonance imaging sought to 1) confirm that PASA reflects the effects of aging rather than differences in task difficulty; 2) test the compensation hypothesis; and 3) investigate whether PASA generalizes to deactivations. Young and older participants were scanned during episodic retrieval and visual perceptual tasks, and age-related changes in brain activity common to both tasks were identified. The study yielded 3 main findings. First, inconsistent with a difficulty account, the PASA pattern was found across task and confidence levels when matching performance among groups. Second, supporting the compensatory hypothesis, age-related increases in frontal activity were positively correlated with performance and negatively correlated with the age-related occipital decreases. Age-related increases and correlations with parietal activity were also found. Finally, supporting the generalizability of the PASA pattern to deactivations, aging reduced deactivations in posterior midline cortex but increased deactivations in medial frontal cortex. Taken together, these findings demonstrate the validity, function, and generalizability of PASA, as well as its importance for the cognitive neuroscience of aging.

The Spatiotemporal Dynamics of Autobiographical Memory: Neural Correlates of Recall, Emotional Intensity, and Reliving

Cerebral Cortex (New York, N.Y. : 1991). Jan, 2008  |  Pubmed ID: 17548799

We sought to map the time course of autobiographical memory retrieval, including brain regions that mediate phenomenological experiences of reliving and emotional intensity. Participants recalled personal memories to auditory word cues during event-related functional magnetic resonance imaging (fMRI). Participants pressed a button when a memory was accessed, maintained and elaborated the memory, and then gave subjective ratings of emotion and reliving. A novel fMRI approach based on timing differences capitalized on the protracted reconstructive process of autobiographical memory to segregate brain areas contributing to initial access and later elaboration and maintenance of episodic memories. The initial period engaged hippocampal, retrosplenial, and medial and right prefrontal activity, whereas the later period recruited visual, precuneus, and left prefrontal activity. Emotional intensity ratings were correlated with activity in several regions, including the amygdala and the hippocampus during the initial period. Reliving ratings were correlated with activity in visual cortex and ventromedial and inferior prefrontal regions during the later period. Frontopolar cortex was the only brain region sensitive to emotional intensity across both periods. Results were confirmed by time-locked averages of the fMRI signal. The findings indicate dynamic recruitment of emotion-, memory-, and sensory-related brain regions during remembering and their dissociable contributions to phenomenological features of the memories.

The Short and Long of It: Neural Correlates of Temporal-order Memory for Autobiographical Events

Journal of Cognitive Neuroscience. Jul, 2008  |  Pubmed ID: 18284345

Previous functional neuroimaging studies of temporal-order memory have investigated memory for laboratory stimuli that are causally unrelated and poor in sensory detail. In contrast, the present functional magnetic resonance imaging (fMRI) study investigated temporal-order memory for autobiographical events that were causally interconnected and rich in sensory detail. Participants took photographs at many campus locations over a period of several hours, and the following day they were scanned while making temporal-order judgments to pairs of photographs from different locations. By manipulating the temporal lag between the two locations in each trial, we compared the neural correlates associated with reconstruction processes, which we hypothesized depended on recollection and contribute mainly to short lags, and distance processes, which we hypothesized to depend on familiarity and contribute mainly to longer lags. Consistent with our hypotheses, parametric fMRI analyses linked shorter lags to activations in regions previously associated with recollection (left prefrontal, parahippocampal, precuneus, and visual cortices), and longer lags with regions previously associated with familiarity (right prefrontal cortex). The hemispheric asymmetry in prefrontal cortex activity fits very well with evidence and theories regarding the contributions of the left versus right prefrontal cortex to memory (recollection vs. familiarity processes) and cognition (systematic vs. heuristic processes). In sum, using a novel photo-paradigm, this study provided the first evidence regarding the neural correlates of temporal-order for autobiographical events.

Age-related Differences in Brain Activity During True and False Memory Retrieval

Journal of Cognitive Neuroscience. Aug, 2008  |  Pubmed ID: 18303982

Compared to young adults, older adults show not only a reduction in true memories but also an increase in false memories. We investigated the neural bases of these age effects using functional magnetic resonance imaging and a false memory task that resembles the Deese-Roediger-McDermott (DRM) paradigm. Young and older participants were scanned during a word recognition task that included studied words and new words that were strongly associated with studied words (critical lures). During correct recognition of studied words (true memory), older adults showed weaker activity than young adults in the hippocampus but stronger activity than young adults in the retrosplenial cortex. The hippocampal reduction is consistent with age-related deficits in recollection, whereas the retrosplenial increase suggests compensatory recruitment of alternative recollection-related regions. During incorrect recognition of critical lures (false memory), older adults displayed stronger activity than young adults in the left lateral temporal cortex, a region involved in semantic processing and semantic gist. Taken together, the results suggest that older adults' deficits in true memories reflect a decline in recollection processes mediated by the hippocampus, whereas their increased tendency to have false memories reflects their reliance on semantic gist mediated by the lateral temporal cortex.

Recollection- and Familiarity-based Memory in Healthy Aging and Amnestic Mild Cognitive Impairment

Neuropsychology. Mar, 2008  |  Pubmed ID: 18331160

Little is known about the cognitive mechanisms of the memory impairment associated with amnestic mild cognitive impairment (aMCI). We explored recollection and familiarity in 27 healthy young adults, 45 healthy older adults, and 17 individuals with aMCI. Relative to the younger adults, recollection was reduced in the older adults, especially among those with aMCI. Familiarity did not differ among groups. In the healthy younger and older adults, better performance on a set of clinical memory measures that are sensitive to medial temporal lobe functioning was associated with greater recollection. In addition, among the healthy older adults better executive functioning was also associated with greater recollection. These results are consistent with the notion that recollection is a product of strategic processes mediated by the prefrontal cortex that suppport the retrieval of context-dependent memories from the hippocampus. Hippocampal atrophy associated with aMCI may disrupt this brain network, and thereby interfere with recollection.

Role of Amygdala Connectivity in the Persistence of Emotional Memories over Time: an Event-related FMRI Investigation

Cerebral Cortex (New York, N.Y. : 1991). Nov, 2008  |  Pubmed ID: 18375529

According to the consolidation hypothesis, enhanced memory for emotional information reflects the modulatory effect of the amygdala on the medial temporal lobe (MTL) memory system during consolidation. Although there is evidence that amygdala-MTL connectivity enhances memory for emotional stimuli, it remains unclear whether this enhancement increases over time, as consolidation processes unfold. To investigate this, we used functional magnetic resonance imaging to measure encoding activity predicting memory for emotionally negative and neutral pictures after short (20-min) versus long (1-week) delays. Memory measures distinguished between vivid remembering (recollection) and feelings of knowing (familiarity). Consistent with the consolidation hypothesis, the persistence of recollection over time (long divided by short) was greater for emotional than neutral pictures. Activity in the amygdala predicted subsequent memory to a greater extent for emotional than neutral pictures. Although this advantage did not vary with delay, the contribution of amygdala-MTL connectivity to subsequent memory for emotional items increased over time. Moreover, both this increase in connectivity and amygdala activity itself were correlated with individual differences in recollection persistence for emotional but not neutral pictures. These results suggest that the amygdala and its connectivity with the MTL are critical to sustaining emotional memories over time, consistent with the consolidation hypothesis.

Role of Parietal Regions in Episodic Memory Retrieval: the Dual Attentional Processes Hypothesis

Neuropsychologia. 2008  |  Pubmed ID: 18439631

Although parietal cortex is frequently activated during episodic memory retrieval, damage to this region does not markedly impair episodic memory. To account for these and other findings, a new dual attentional processes (DAP) hypothesis is proposed. According to this hypothesis, dorsal parietal cortex (DPC) contributes top-down attentional processes guided by retrieval goals, whereas ventral parietal cortex (VPC) contributes bottom-up attentional processes captured by the retrieval output. Consistent with this hypothesis, DPC activity increases with retrieval effort whereas VPC activity increases with confidence in old and new responses. The DAP hypothesis can also account for the overlap of parietal activations across different cognitive domains and for opposing effects of parietal activity on encoding vs. retrieval. Finally, the DAP hypothesis explains why VPC lesions yield a memory neglect syndrome: a deficit in spontaneously reporting relevant memory details but not in accessing the same details when guided by specific questions.

Orbitofrontal and Hippocampal Contributions to Memory for Face-name Associations: the Rewarding Power of a Smile

Neuropsychologia. 2008  |  Pubmed ID: 18455740

Memory processes can be enhanced by reward, and social signals such a smiling face can be rewarding to humans. Using event-related functional MRI (fMRI), we investigated the rewarding effect of a simple smile during the encoding and retrieval of face-name associations. During encoding, participants viewed smiling or neutral faces, each paired with a name, and during retrieval, only names were presented, and participants retrieved the associated facial expressions. Successful memory activity of face-name associations was identified by comparing remembered vs. forgotten trials during both encoding and retrieval, and the effect of a smile was identified by comparing successful memory trials for smiling vs. neutral faces. The study yielded three main findings. First, behavioral results showed that the retrieval of face-name associations was more accurate and faster for smiling than neutral faces. Second, the orbitofrontal cortex and the hippocampus showed successful encoding and retrieval activations, which were greater for smiling than neutral faces. Third, functional connectivity between the orbitofrontal cortex and the hippocampus during successful encoding and retrieval was stronger for smiling than neutral faces. As a part of the reward system, the orbitofrontal cortex may modulate memory processes of face-name associations mediated by the hippocampus. Interestingly, the effect of a smile during retrieval was found even though only names were presented as retrieval cues, suggesting that the effect was mediated by face imagery. Taken together, the results demonstrate how rewarding social signals from a smiling face can enhance relational memory for face-name associations.

Effects of Aging on the Neural Correlates of Successful Item and Source Memory Encoding

Journal of Experimental Psychology. Learning, Memory, and Cognition. Jul, 2008  |  Pubmed ID: 18605869

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.

The Parietal Cortex and Episodic Memory: an Attentional Account

Nature Reviews. Neuroscience. Aug, 2008  |  Pubmed ID: 18641668

The contribution of the parietal cortex to episodic memory is a fascinating scientific puzzle. On the one hand, parietal lesions do not normally yield severe episodic-memory deficits; on the other hand, parietal activations are seen frequently in functional-neuroimaging studies of episodic memory. A review of these two categories of evidence suggests that the answer to the puzzle requires us to distinguish between the contributions of dorsal and ventral parietal regions and between the influence of top-down and bottom-up attention on memory.

Age-related Slowing of Memory Retrieval: Contributions of Perceptual Speed and Cerebral White Matter Integrity

Neurobiology of Aging. Jul, 2008  |  Pubmed ID: 17383774

Previous research suggests that, in reaction time (RT) measures of episodic memory retrieval, the unique effects of adult age are relatively small compared to the effects aging shares with more elementary abilities such as perceptual speed. Little is known, however, regarding the mechanisms of perceptual speed. We used diffusion tensor imaging (DTI) to test the hypothesis that white matter integrity, as indexed by fractional anisotropy (FA), serves as one mechanism of perceptual slowing in episodic memory retrieval. Results indicated that declines in FA in the pericallosal frontal region and in the genu of the corpus callosum, but not in other regions, mediated the relationship between perceptual speed and episodic retrieval RT. This relation held, though to a different degree, for both hits and correct rejections. These findings suggest that white matter integrity in prefrontal regions is one mechanism underlying the relation between individual differences in perceptual speed and episodic retrieval.

When Learning and Remembering Compete: a Functional MRI Study

PLoS Biology. Jan, 2009  |  Pubmed ID: 19143473

Recent functional neuroimaging evidence suggests a bottleneck between learning new information and remembering old information. In two behavioral experiments and one functional MRI (fMRI) experiment, we tested the hypothesis that learning and remembering compete when both processes happen within a brief period of time. In the first behavioral experiment, participants intentionally remembered old words displayed in the foreground, while incidentally learning new scenes displayed in the background. In line with a memory competition, we found that remembering old information was associated with impaired learning of new information. We replicated this finding in a subsequent fMRI experiment, which showed that this behavioral effect was coupled with a suppression of learning-related activity in visual and medial temporal areas. Moreover, the fMRI experiment provided evidence that left mid-ventrolateral prefrontal cortex is involved in resolving the memory competition, possibly by facilitating rapid switching between learning and remembering. Critically, a follow-up behavioral experiment in which the background scenes were replaced with a visual target detection task provided indications that the competition between learning and remembering was not merely due to attention. This study not only provides novel insight into our capacity to learn and remember, but also clarifies the neural mechanisms underlying flexible behavior.

Effects of Aging on Functional Connectivity of the Amygdala for Subsequent Memory of Negative Pictures: a Network Analysis of Functional Magnetic Resonance Imaging Data

Psychological Science. Jan, 2009  |  Pubmed ID: 19152542

Aging is associated with preserved enhancement of emotional memory, as well as with age-related reductions in memory for negative stimuli, but the neural networks underlying such alterations are not clear. We used a subsequent-memory paradigm to identify brain activity predicting enhanced emotional memory in young and older adults. Activity in the amygdala predicted enhanced emotional memory, with subsequent-memory activity greater for negative stimuli than for neutral stimuli, across age groups, a finding consistent with an overall enhancement of emotional memory. However, older adults recruited greater activity in anterior regions and less activity in posterior regions in general for negative stimuli that were subsequently remembered. Functional connectivity of the amygdala with the rest of the brain was consistent with age-related reductions in memory for negative stimuli: Older adults showed decreased functional connectivity between the amygdala and the hippocampus, but increased functional connectivity between the amygdala and dorsolateral prefrontal cortices. These findings suggest that age-related differences in the enhancement of emotional memory might reflect decreased connectivity between the amygdala and typical subsequent-memory regions, as well as the engagement of regulatory processes that inhibit emotional responses.

Assessing the Effects of Age on Long White Matter Tracts Using Diffusion Tensor Tractography

NeuroImage. Jun, 2009  |  Pubmed ID: 19385018

Aging is associated with significant white matter deterioration and this deterioration is assumed to be at least partly a consequence of myelin degeneration. The present study investigated specific predictions of the myelodegeneration hypothesis using diffusion tensor tractography. This technique has several advantages over other methods of assessing white matter architecture, including the possibility of isolating individual white matter tracts and measuring effects along the whole extent of each tract. The study yielded three main findings. First, age-related white matter deficits increased gradually from posterior to anterior segments within specific fiber tracts traversing frontal and parietal, but not temporal cortex. This pattern inverts the sequence of myelination during childhood and early development observed in previous studies and lends support to a "last-in-first-out" theory of the white matter health across the lifespan. Second, both the effects of aging on white matter and their impact on cognitive performance were stronger for radial diffusivity (RD) than for axial diffusivity (AD). Given that RD has previously been shown to be more sensitive to myelin integrity than AD, this second finding is also consistent with the myelodegeneration hypothesis. Finally, the effects of aging on select white matter tracts were associated with age difference in specific cognitive functions. Specifically, FA in anterior tracts was shown to be primarily associated with executive tasks and FA in posterior tracts mainly associated with visual memory tasks. Furthermore, these correlations were mirrored in RD, but not AD, suggesting that RD is more sensitive to age-related changes in cognition. Taken together, the results help to clarify how age-related white matter decline impairs cognitive performance.

Neural Correlates of Idiographic Goal Priming in Depression: Goal-specific Dysfunctions in the Orbitofrontal Cortex

Social Cognitive and Affective Neuroscience. Sep, 2009  |  Pubmed ID: 19433416

We used functional magnetic resonance imaging (fMRI) to determine whether depressed (vs non-depressed) adults showed differences in cortical activation in response to stimuli representing personal goals. Drawing upon regulatory focus theory as well as previous research, we predicted that depressed patients would manifest attenuated left orbitofrontal cortex (OFC) activation in response to their own promotion goals as well as exaggerated right OFC activation in response to their own prevention goals. Unmedicated adults with major depression (n = 22) and adults with no history of affective disorder (n = 14) completed questionnaires and a personal goal interview. Several weeks later, they were scanned during a judgment task which (unknown to them) included stimuli representing their promotion and prevention goals. Both groups showed similar patterns of task-related activation. Consistent with predictions, patients showed significantly decreased left OFC and increased right OFC activation compared to controls on trials in which they were exposed incidentally to their promotion and prevention goals, respectively. The results suggest that depression involves dysfunction in processing two important types of personal goals. The findings extend models of the etiology of depression to incorporate cognitive and motivational processes underlying higher order goal representation and ultimately may provide an empirical basis for treatment matching.

Common and Specific Brain Regions in High- Versus Low-confidence Recognition Memory

Brain Research. Jul, 2009  |  Pubmed ID: 19501072

The goal of the present functional magnetic resonance imaging (fMRI) study was to investigate whether and to what extent brain regions involved in high-confidence recognition (HCR) versus low-confidence recognition (LCR) overlap or separate from each other. To this end, we performed conjunction analyses involving activations elicited during high-confidence hit, low-confidence hit, and high-confidence correct rejection responses. The analyses yielded 3 main findings. First, sensory/perceptual and associated posterior regions were common to HCR and LCR, indicating contribution of these regions to both HCR and LCR activity. This finding may help explain why these regions are among the most common in functional neuroimaging studies of episodic retrieval. Second, medial temporal lobe (MTL) and associated midline regions were associated with HCR, possibly reflecting recollection-related processes, whereas specific prefrontal cortex (PFC) regions were associated with LCR, possibly reflecting executive control processes. This finding is consistent with the notion that the MTL and PFC networks play complementary roles during episodic retrieval. Finally, within posterior parietal cortex, a dorsal region was associated with LCR, possibly reflecting top-down attentional processes, whereas a ventral region was associated with HCR, possibly reflecting bottom-up attentional processes. This finding may help explain why functional neuroimaging studies have found diverse parietal effects during episodic retrieval. Taken together, our findings provide strong evidence that HCR versus LCR, and by implication, recollection versus familiarity processes, are represented in common as well as specific brain regions.

Encoding and Retrieving Faces and Places: Distinguishing Process- and Stimulus-specific Differences in Brain Activity

Neuropsychologia. Sep, 2009  |  Pubmed ID: 19524092

Among the most fundamental issues in cognitive neuroscience is how the brain may be organized into process-specific and stimulus-specific regions. In the episodic memory domain, most functional neuroimaging studies have focused on the former dimension, typically investigating the neural correlates of various memory processes. Thus, there is little information about what role stimulus-specific brain regions play in successful memory processes. To address this issue, the present event-related fMRI study used a factorial design to focus on the role of stimulus-specific brain regions, such as the fusiform face area (FFA) and parahippocampal place area (PPA) in successful encoding and retrieval processes. Searching within regions sensitive to faces or places, we identified areas similarly involved in encoding and retrieval, as well as areas differentially involved in encoding or retrieval. Finally, we isolated regions associated with successful memory, regardless of stimulus and process type. There were three main findings. Within face sensitive regions, anterior medial PFC and right FFA displayed equivalent encoding and retrieval success processes whereas left FFA was associated with successful encoding rather than retrieval. Within place sensitive regions, left PPA displayed equivalent encoding and retrieval success processes whereas right PPA was associated with successful encoding rather than retrieval. Finally, medial temporal and prefrontal regions were associated with general memory success, regardless of stimulus or process type. Taken together, our results clarify the contribution of different brain regions to stimulus- and process-specific episodic memory mechanisms.

Posterior Midline and Ventral Parietal Activity is Associated with Retrieval Success and Encoding Failure

Frontiers in Human Neuroscience. 2009  |  Pubmed ID: 19680466

The ventral part of lateral posterior parietal cortex (VPC) and the posterior midline region (PMR), including the posterior cingulate cortex and precuneus, tend to show deactivation during demanding cognitive tasks, and have been associated with the default mode of the brain. Interestingly, PMR and VPC activity has been associated with successful episodic retrieval but also with unsuccessful episodic encoding. However, the differential contributions of PMR and VPC to retrieval vs. encoding has never been demonstrated within-subjects and within the same experiment. Here, we directly tested the prediction that PMR and VPC activity should be associated with retrieval success but with encoding failure. Consistent with this prediction, we found across five different fMRI experiments that, during retrieval, activity in these regions is greater for hits than misses, whereas during encoding, it is greater for subsequent misses than hits. We also found that these regions overlap with the ones that show deactivations during conscious rest. Our findings further aid in clarifying the role of the default mode regions in learning and memory.

Functional Neuroimaging Studies of Aging and Emotion: Fronto-amygdalar Differences During Emotional Perception and Episodic Memory

Journal of the International Neuropsychological Society : JINS. Nov, 2009  |  Pubmed ID: 19703320

Emotional processes are enhanced in aging, such that aging is characterized by superior emotional regulation. This article provides a brief review of the neural bases supporting this effect with a focus on functional neuroimaging studies of perception and episodic memory. The most consistent finding across these studies is that older adults show an alteration in the recruitment of the amygdala, but greater recruitment of the frontal cortex. These Fronto-amygdalar Age-related Differences in Emotion (FADE) may reflect emotional regulation strategies mediated by frontal brain regions that dampen emotion-related activations in the amygdala.

Cerebral White Matter Integrity Mediates Adult Age Differences in Cognitive Performance

Journal of Cognitive Neuroscience. Feb, 2009  |  Pubmed ID: 18564054

Previous research has established that age-related decline occurs in measures of cerebral white matter integrity, but the role of this decline in age-related cognitive changes is not clear. To conclude that white matter integrity has a mediating (causal) contribution, it is necessary to demonstrate that statistical control of the white matter-cognition relation reduces the magnitude of age-cognition relation. In this research, we tested the mediating role of white matter integrity, in the context of a task-switching paradigm involving word categorization. Participants were 20 healthy, community-dwelling older adults (60-85 years), and 20 younger adults (18-27 years). From diffusion tensor imaging tractography, we obtained fractional anisotropy (FA) as an index of white matter integrity in the genu and splenium of the corpus callosum and the superior longitudinal fasciculus (SLF). Mean FA values exhibited age-related decline consistent with a decrease in white matter integrity. From a model of reaction time distributions, we obtained independent estimates of the decisional and nondecisional (perceptual-motor) components of task performance. Age-related decline was evident in both components. Critically, age differences in task performance were mediated by FA in two regions: the central portion of the genu, and splenium-parietal fibers in the right hemisphere. This relation held only for the decisional component and was not evident in the nondecisional component. This result is the first demonstration that the integrity of specific white matter tracts is a mediator of age-related changes in cognitive performance.

Brain-derived Neurotrophic Factor Val66met Polymorphism and Hippocampal Activation During Episodic Encoding and Retrieval Tasks

Hippocampus. May, 2010  |  Pubmed ID: 20865733

Brain-derived neurotrophic factor (BDNF) is a neurotrophin which has been shown to regulate cell survival and proliferation, as well as synaptic growth and hippocampal long-term potentiation. A naturally occurring single nucleotide polymorphism in the human BDNF gene (val66met) has been associated with altered intercellular trafficking and regulated secretion of BDNF in met compared to val carriers. Additionally, previous studies have found a relationship between the BDNF val66met genotype and functional activity in the hippocampus during episodic and working memory tasks in healthy young adults. Specifically, studies have found that met carriers exhibit both poorer performance and reduced neural activity within the medial temporal lobe (MTL) when performing episodic memory tasks. However, these studies have not been well replicated and have not considered the role of behavioral differences in the interpretation of neural differences. The current study sought to control for cognitive performance in investigating the role of the BDNF val66met genotype on neural activity associated with episodic memory. Across item and relational memory tests, met carriers exhibited increased MTL activation during both encoding and retrieval stages, compared to noncarriers. The results suggest that met carriers are able to recruit MTL activity to support successful memory processes, and reductions in cognitive performance observed in prior studies are not a ubiquitous effect associated with variants of the BDNF val66met genotype. © 2010 Wiley-Liss, Inc.

Age-related Effects on the Neural Correlates of Autobiographical Memory Retrieval

Neurobiology of Aging. Dec, 2010  |  Pubmed ID: 21190759

Older adults recall less episodically rich autobiographical memories (AM), however, the neural basis of this effect is not clear. Using functional MRI, we examined the effects of age during search and elaboration phases of AM retrieval. Our results suggest that the age-related attenuation in the episodic richness of AMs is associated with difficulty in the strategic retrieval processes underlying recovery of information during elaboration. First, age effects on AM activity were more pronounced during elaboration than search, with older adults showing less sustained recruitment of the hippocampus and ventrolateral prefrontal cortex (VLPFC) for less episodically rich AMs. Second, there was an age-related reduction in the modulation of top-down coupling of the VLPFC on the hippocampus for episodically rich AMs. In sum, the present study shows that changes in the sustained response and coupling of the hippocampus and prefrontal cortex (PFC) underlie age-related reductions in episodic richness of the personal past.

Temporal Lobe Functional Activity and Connectivity in Young Adult APOE Varepsilon4 Carriers

Alzheimer's & Dementia : the Journal of the Alzheimer's Association. Jul, 2010  |  Pubmed ID: 19744893

We sought to determine if the APOE epsilon4 allele influences both the functional activation and connectivity of the medial temporal lobes (MTLs) during successful memory encoding in young adults.

Neural Mechanisms of Context Effects on Face Recognition: Automatic Binding and Context Shift Decrements

Journal of Cognitive Neuroscience. Nov, 2010  |  Pubmed ID: 19925208

Although people do not normally try to remember associations between faces and physical contexts, these associations are established automatically, as indicated by the difficulty of recognizing familiar faces in different contexts ("butcher-on-the-bus" phenomenon). The present fMRI study investigated the automatic binding of faces and scenes. In the face-face (F-F) condition, faces were presented alone during both encoding and retrieval, whereas in the face/scene-face (FS-F) condition, they were presented overlaid on scenes during encoding but alone during retrieval (context change). Although participants were instructed to focus only on the faces during both encoding and retrieval, recognition performance was worse in the FS-F than in the F-F condition ("context shift decrement" [CSD]), confirming automatic face-scene binding during encoding. This binding was mediated by the hippocampus as indicated by greater subsequent memory effects (remembered > forgotten) in this region for the FS-F than the F-F condition. Scene memory was mediated by right parahippocampal cortex, which was reactivated during successful retrieval when the faces were associated with a scene during encoding (FS-F condition). Analyses using the CSD as a regressor yielded a clear hemispheric asymmetry in medial temporal lobe activity during encoding: Left hippocampal and parahippocampal activity was associated with a smaller CSD, indicating more flexible memory representations immune to context changes, whereas right hippocampal/rhinal activity was associated with a larger CSD, indicating less flexible representations sensitive to context change. Taken together, the results clarify the neural mechanisms of context effects on face recognition.

Mental Hoop Diaries: Emotional Memories of a College Basketball Game in Rival Fans

The Journal of Neuroscience : the Official Journal of the Society for Neuroscience. Feb, 2010  |  Pubmed ID: 20147540

The rivalry between the men's basketball teams of Duke University and the University of North Carolina-Chapel Hill (UNC) is one of the most storied traditions in college sports. A subculture of students at each university form social bonds with fellow fans, develop expertise in college basketball rules, team statistics, and individual players, and self-identify as a member of a fan group. The present study capitalized on the high personal investment of these fans and the strong affective tenor of a Duke-UNC basketball game to examine the neural correlates of emotional memory retrieval for a complex sporting event. Male fans watched a competitive, archived game in a social setting. During a subsequent functional magnetic resonance imaging session, participants viewed video clips depicting individual plays of the game that ended with the ball being released toward the basket. For each play, participants recalled whether or not the shot went into the basket. Hemodynamic signal changes time locked to correct memory decisions were analyzed as a function of emotional intensity and valence, according to the fan's perspective. Results showed intensity-modulated retrieval activity in midline cortical structures, sensorimotor cortex, the striatum, and the medial temporal lobe, including the amygdala. Positively valent memories specifically recruited processing in dorsal frontoparietal regions, and additional activity in the insula and medial temporal lobe for positively valent shots recalled with high confidence. This novel paradigm reveals how brain regions implicated in emotion, memory retrieval, visuomotor imagery, and social cognition contribute to the recollection of specific plays in the mind of a sports fan.

Adult Age Differences in Functional Connectivity During Executive Control

NeuroImage. Aug, 2010  |  Pubmed ID: 20434565

Task switching requires executive control processes that undergo age-related decline. Previous neuroimaging studies have identified age-related differences in brain activation associated with global switching effects (dual-task blocks versus single-task blocks), but age-related differences in activation during local switching effects (switch trials versus repeat trials, within blocks) have not been investigated. This experiment used functional magnetic resonance imaging (fMRI), and diffusion tensor imaging (DTI), to examine adult age differences in task switching across adjacent trials (i.e., local task switching). During fMRI scanning, participants performed a cued, word categorization task. From interspersed cue-only trials, switch-related processing associated with the cue was estimated separately from the target. Activation associated with task switching, within a distributed frontoparietal network, differed for cue- and target-related processing. The magnitude of event-related activation for task switching was similar for younger adults (n=20; 18-27years) and older adults (n=20; 60-85years), although activation sustained throughout the on-tasks periods exhibited some age-related decline. Critically, the functional connectivity of switch-related regions, during cue processing, was higher for younger adults than for older adults, whereas functional connectivity during target processing was comparable across the age groups. Further, individual differences in cue-related functional connectivity shared a substantial portion of the age-related variability in the efficiency of target categorization response (drift rate). This age-related difference in functional connectivity, however, was independent of white matter integrity within task-relevant regions. These findings highlight the functional connectivity of frontoparietal activation as a potential source of age-related decline in executive control.

COMT Val108/158 Met Genotype Affects Neural but Not Cognitive Processing in Healthy Individuals

Cerebral Cortex (New York, N.Y. : 1991). Mar, 2010  |  Pubmed ID: 19641018

The relationship between cognition and a functional polymorphism in the catechol-O-methlytransferase (COMT) gene, val108/158met, is one of debate in the literature. Furthermore, based on the dopaminergic differences associated with the COMT val108/158met genotype, neural differences during cognition may be present, regardless of genotypic differences in cognitive performance. To investigate these issues the current study aimed to 1) examine the effects of COMT genotype using a large sample of healthy individuals (n = 496-1218) and multiple cognitive measures, and using a subset of the sample (n = 22), 2) examine whether COMT genotype effects medial temporal lobe (MTL) and frontal activity during successful relational memory processing, and 3) investigate group differences in functional connectivity associated with successful relational memory processing. Results revealed no significant group difference in cognitive performance between COMT genotypes in any of the 19 cognitive measures. However, in the subset sample, COMT val homozygotes exhibited significantly decreased MTL and increased prefrontal activity during both successful relational encoding and retrieval, and reduced connectivity between these regions compared with met homozygotes. Taken together, the results suggest that although the COMT val108/158met genotype has no effect on cognitive behavioral measures in healthy individuals, it is associated with differences in neural process underlying cognitive output.

Overlapping Brain Activity Between Episodic Memory Encoding and Retrieval: Roles of the Task-positive and Task-negative Networks

NeuroImage. Jan, 2010  |  Pubmed ID: 19647800

The notion that the brain is organized into two complementary networks, one that is task-positive and supports externally-oriented processing, and the other that is task-negative and supports internally-oriented processing, has recently attracted increasing attention. The goal of the present study was to investigate involvement of the task-positive and task-negative networks in overlapping activity between episodic memory encoding and retrieval. To this end, we performed a functional MRI study that included both encoding and retrieval tasks. We hypothesized that during the study phase, encoding success activity (remembered > forgotten) involves mainly the task-positive network, whereas encoding failure activity (forgotten > remembered) involves mainly the task-negative network. We also hypothesized that during the test phase, retrieval success activity (old > new) involves mainly the task-negative network, whereas novelty detection activity (new > old) involves mainly the task-positive network. Based on these hypotheses, we made 3 predictions regarding study-test overlap. First, there would be relatively high level of overlap between encoding success and novelty detection activity involving the task-positive network. Second, there would be relatively high level of overlap between encoding failure and retrieval success activity involving the task-negative network. Third, there would be relatively low level of overlap between encoding success and retrieval success activity as well as between encoding failure and novelty detection activity. The results fully confirmed our 3 predictions. Taken together, the present findings clarify roles of the task-positive and task-negative networks in encoding and retrieval and the function of overlapping brain activity between encoding and retrieval.

Effects of Aging on Functional Connectivity of the Amygdala During Negative Evaluation: a Network Analysis of FMRI Data

Neurobiology of Aging. Feb, 2010  |  Pubmed ID: 18455837

Previous evidence has suggested both preserved emotional function in aging and age-related differences in emotional processing, but the neural networks underlying such processing alterations in the context of preserved affective function are not clear. Using event-related fMRI, we scanned young and older adults while they made valence ratings for emotional pictures. Behavioral results showed a similar pattern of emotional evaluation, but older adults experienced negatively valenced pictures as being less negative. Consistent with behavioral findings, we identified common activity in the right amygdala, but age-related differences in the functional connectivity of this region with the rest of the brain. Compared to young adults, older adults had greater functional connectivity between the right amygdala and ventral anterior cingulate cortex, possibly reflecting increased emotional regulation. Conversely, older adults showed decreased functional connectivity with posterior brain regions, likely reflecting decreased perceptual processing. Thus, age-related differences in evaluating negatively valenced stimuli might reflect decreased perceptual processing of these stimuli, as well as the engagement of control processes that inhibit the response to negative emotion.

Age-related Dedifferentiation of Learning Systems: an FMRI Study of Implicit and Explicit Learning

Neurobiology of Aging. Dec, 2011  |  Pubmed ID: 20471139

Abundant research finds that in young adults explicit learning (EL) is more dependent on the medial temporal lobes (MTL) whereas implicit learning (IL) is more dependent on the striatum. Using fMRI, we investigated age differences in each task and whether this differentiation is preserved in older adults. Results indicated that, while young recruited the MTL for EL and striatum for IL, both activations were significantly reduced in older adults. Additionally, results indicated that older adults recruited the MTL for IL, and this activation was significantly greater in older compared with young adults. A significant Task × Age interaction was found in both regions-with young preferentially recruiting the MTL for EL and striatum for IL, and older adults showing no preferential recruit for either task. Finally, young adults demonstrated significant negative correlations between activity in the striatum and MTL during both the EL and IL tasks. These correlations were attenuated in older adults. Taken together results support dedifferentiation in aging across memory systems.

Watching My Mind Unfold Versus Yours: an FMRI Study Using a Novel Camera Technology to Examine Neural Differences in Self-projection of Self Versus Other Perspectives

Journal of Cognitive Neuroscience. Jun, 2011  |  Pubmed ID: 20521858

Self-projection, the capacity to re-experience the personal past and to mentally infer another person's perspective, has been linked to medial prefrontal cortex (mPFC). In particular, ventral mPFC is associated with inferences about one's own self, whereas dorsal mPFC is associated with inferences about another individual. In the present fMRI study, we examined self-projection using a novel camera technology, which employs a sensor and timer to automatically take hundreds of photographs when worn, in order to create dynamic visuospatial cues taken from a first-person perspective. This allowed us to ask participants to self-project into the personal past or into the life of another person. We predicted that self-projection to the personal past would elicit greater activity in ventral mPFC, whereas self-projection of another perspective would rely on dorsal mPFC. There were three main findings supporting this prediction. First, we found that self-projection to the personal past recruited greater ventral mPFC, whereas observing another person's perspective recruited dorsal mPFC. Second, activity in ventral versus dorsal mPFC was sensitive to parametric modulation on each trial by the ability to relive the personal past or to understand another's perspective, respectively. Third, task-related functional connectivity analysis revealed that ventral mPFC contributed to the medial temporal lobe network linked to memory processes, whereas dorsal mPFC contributed to the fronto-parietal network linked to controlled processes. In sum, these results suggest that ventral-dorsal subregions of the anterior midline are functionally dissociable and may differentially contribute to self-projection of self versus other.

Remembering Beauty: Roles of Orbitofrontal and Hippocampal Regions in Successful Memory Encoding of Attractive Faces

NeuroImage. Jan, 2011  |  Pubmed ID: 20659568

Behavioral data have shown that attractive faces are better remembered but the neural mechanisms of this effect are largely unknown. To investigate this issue, female participants were scanned with event-related functional MRI (fMRI) while rating the attractiveness of male faces. Memory for the faces was tested after fMRI scanning and was used to identify successful encoding activity (subsequent memory paradigm). As expected, attractive faces were remembered better than other faces. The study yielded three main fMRI findings. First, activity in the right orbitofrontal cortex increased linearly as a function of attractiveness ratings. Second, activity in the left hippocampus increased as a function of subsequent memory (subsequent misses

Shared Brain Activity for Aesthetic and Moral Judgments: Implications for the Beauty-is-Good Stereotype

Social Cognitive and Affective Neuroscience. Jan, 2011  |  Pubmed ID: 20231177

The Beauty-is-Good stereotype refers to the assumption that attractive people possess sociably desirable personalities and higher moral standards. The existence of this bias suggests that the neural mechanisms for judging facial attractiveness and moral goodness overlap. To investigate this idea, we scanned participants with functional magnetic resonance imaging while they made attractiveness judgments about faces and goodness judgments about hypothetical actions. Activity in the medial orbitofrontal cortex increased as a function of both attractiveness and goodness ratings, whereas activity in the insular cortex decreased with both attractiveness and goodness ratings. Within each of these regions, the activations elicited by attractiveness and goodness judgments were strongly correlated with each other, supporting the idea of similar contributions of each region to both judgments. Moreover, activations in orbitofrontal and insular cortices were negatively correlated with each other, suggesting an opposing relationship between these regions during attractiveness and goodness judgments. These findings have implications for understanding the neural mechanisms of the Beauty-is-Good stereotype.

Level of Processing Modulates the Neural Correlates of Emotional Memory Formation

Journal of Cognitive Neuroscience. Apr, 2011  |  Pubmed ID: 20350176

Emotion is known to influence multiple aspects of memory formation, including the initial encoding of the memory trace and its consolidation over time. However, the neural mechanisms whereby emotion impacts memory encoding remain largely unexplored. The present study used a levels-of-processing manipulation to characterize the impact of emotion on encoding with and without the influence of elaborative processes. Participants viewed emotionally negative, neutral, and positive scenes under two conditions: a shallow condition focused on the perceptual features of the scenes and a deep condition that queried their semantic meaning. Recognition memory was tested 2 days later. Results showed that emotional memory enhancements were greatest in the shallow condition. fMRI analyses revealed that the right amygdala predicted subsequent emotional memory in the shallow more than deep condition, whereas the right ventrolateral PFC demonstrated the reverse pattern. Furthermore, the association of these regions with the hippocampus was modulated by valence: the amygdala-hippocampal link was strongest for negative stimuli, whereas the prefrontal-hippocampal link was strongest for positive stimuli. Taken together, these results suggest two distinct activation patterns underlying emotional memory formation: an amygdala component that promotes memory during shallow encoding, especially for negative information, and a prefrontal component that provides extra benefits during deep encoding, especially for positive information.

Cognitive and Neural Contributors to Emotion Regulation in Aging

Social Cognitive and Affective Neuroscience. Apr, 2011  |  Pubmed ID: 20385663

Older adults, compared to younger adults, focus on emotional well-being. While the lifespan trajectory of emotional processing and its regulation has been characterized behaviorally, few studies have investigated the underlying neural mechanisms. Here, older adults (range: 59-73 years) and younger adults (range: 19-33 years) participated in a cognitive reappraisal task during functional magnetic resonance imaging (fMRI) scanning. On each trial, participants viewed positive, negative or neutral pictures and either naturally experienced the image ('Experience' condition) or attempted to detach themselves from the image ('Reappraise' condition). Across both age groups, cognitive reappraisal activated prefrontal regions similar to those reported in prior studies of emotion regulation, while emotional experience activated the bilateral amygdala. Psychophysiological interaction analyses revealed that the left inferior frontal gyrus (IFG) and amygdala demonstrated greater inverse connectivity during the 'Reappraise' condition relative to the 'Experience' condition. The only regions exhibiting significant age differences were the left IFG and the left superior temporal gyrus, for which greater regulation-related activation was observed in younger adults. Controlling for age, increased performance on measures of cognition predicted greater regulation-related decreases in amygdala activation. Thus, while older and younger adults use similar brain structures for emotion regulation and experience, the functional efficacy of those structures depends on underlying cognitive ability.

The Porous Boundaries Between Explicit and Implicit Memory: Behavioral and Neural Evidence

Annals of the New York Academy of Sciences. Apr, 2011  |  Pubmed ID: 21486300

Explicit memory refers to the conscious retrieval of past information or experiences, whereas implicit memory refers to an unintentional or nonconscious form of retrieval. Much of the literature in cognitive psychology and cognitive neuroscience has focused on differences between explicit and implicit memory, and the traditional view is that they rely on distinct brain systems. However, the potential interplay between implicit and explicit memory is not always clear. This review draws from behavioral and functional neuroimaging evidence to evaluate three areas in which implicit and explicit memory may be interrelated. First, we discuss views of familiarity-based recognition in terms of its relationship with implicit memory. Second, we review the challenges of distinguishing between implicit memory and involuntary aware memory, at both behavioral and neural levels. Finally, we examine evidence indicating that implicit and explicit retrieval of relational information may rely on a common neural mechanism. Taken together, these areas indicate that, under certain circumstances, there may be an important and influential relationship between conscious and nonconscious expressions of memory.

The Hippocampus is Coupled with the Default Network During Memory Retrieval but Not During Memory Encoding

PloS One. 2011  |  Pubmed ID: 21494597

The brain's default mode network (DMN) is activated during internally-oriented tasks and shows strong coherence in spontaneous rest activity. Despite a surge of recent interest, the functional role of the DMN remains poorly understood. Interestingly, the DMN activates during retrieval of past events but deactivates during encoding of novel events into memory. One hypothesis is that these opposing effects reflect a difference between attentional orienting towards internal events, such as retrieved memories, vs. external events, such as to-be-encoded stimuli. Another hypothesis is that hippocampal regions are coupled with the DMN during retrieval but decoupled from the DMN during encoding. The present fMRI study investigated these two hypotheses by combining a resting-state coherence analysis with a task that measured the encoding and retrieval of both internally-generated and externally-presented events. Results revealed that the main DMN regions were activated during retrieval but deactivated during encoding. Counter to the internal orienting hypothesis, this pattern was not modulated by whether memory events were internal or external. Consistent with the hippocampal coupling hypothesis, the hippocampus behaved like other DMN regions during retrieval but not during encoding. Taken together, our findings clarify the relationship between the DMN and the neural correlates of memory retrieval and encoding.

Overlapping Parietal Activity in Memory and Perception: Evidence for the Attention to Memory Model

Journal of Cognitive Neuroscience. Nov, 2011  |  Pubmed ID: 21568633

The specific role of different parietal regions to episodic retrieval is a topic of intense debate. According to the Attention to Memory (AtoM) model, dorsal parietal cortex (DPC) mediates top-down attention processes guided by retrieval goals, whereas ventral parietal cortex (VPC) mediates bottom-up attention processes captured by the retrieval output or the retrieval cue. This model also hypothesizes that the attentional functions of DPC and VPC are similar for memory and perception. To investigate this last hypothesis, we scanned participants with event-related fMRI whereas they performed memory and perception tasks, each comprising an orienting phase (top-down attention) and a detection phase (bottom-up attention). The study yielded two main findings. First, consistent with the AtoM model, orienting-related activity for memory and perception overlapped in DPC, whereas detection-related activity for memory and perception overlapped in VPC. The DPC overlap was greater in the left intraparietal sulcus, and the VPC overlap in the left TPJ. Around overlapping areas, there were differences in the spatial distribution of memory and perception activations, which were consistent with trends reported in the literature. Second, both DPC and VPC showed stronger connectivity with medial-temporal lobe during the memory task and with visual cortex during the perception task. These findings suggest that, during memory tasks, some parietal regions mediate similar attentional control processes to those involved in perception tasks (orienting in DPC vs. detection in VPC), although on different types of information (mnemonic vs. sensory).

Emotion Processing in the Aging Brain is Modulated by Semantic Elaboration

Neuropsychologia. Mar, 2011  |  Pubmed ID: 20869375

The neural correlates of emotion processing have been shown to vary with age: older adults (OAs) exhibit increased frontal activations and, under some circumstances, decreased amygdala activations relative to young adults (YAs) during emotion processing. Some of these differences are additionally modulated by valence, with age-related biases toward positive versus negative stimuli, and are thought to depend on OAs' capacity for controlled elaboration. However, the role of semantic elaboration in mediating valence effects in the aging brain has not yet been explicitly tested. In the present study, YAs and OAs were scanned while they viewed negative, neutral, and positive pictures during either a deep, elaborative task or a shallow, perceptual task. fMRI results reveal that emotion-related activity in the amygdala is preserved in aging and insensitive to elaboration demands. This study provides novel evidence that differences in valence processing are modulated by elaboration: relative to YAs, OAs show enhanced activity in the medial prefrontal cortex (PFC) and ventrolateral PFC in response to positive versus negative stimuli, but only during elaborative processing. These positive valence effects are predicted by individual differences in executive function in OAs for the deep but not shallow task. Finally, psychophysiological interaction analyses reveal age effects on valence-dependent functional connectivity between medial PFC and ventral striatum, as well as age and task effects on medial PFC-retrosplenial cortex interactions. Altogether, these findings provide support for the hypothesis that valence shifts in the aging brain are mediated by controlled processes such as semantic elaboration, self-referential processing, and emotion regulation.

Neural Correlates of Emotional Processing in Depression: Changes with Cognitive Behavioral Therapy and Predictors of Treatment Response

Journal of Psychiatric Research. May, 2011  |  Pubmed ID: 20934190

Major depressive disorder (MDD) is characterized by the presence of disturbances in emotional processing. However, the neural correlates of these alterations, and how they may be affected by therapeutic interventions, remain unclear. The present study addressed these issues in a preliminary investigation using functional magnetic resonance imaging (fMRI) to examine neural responses to positive, negative, and neutral pictures in unmedicated MDD patients (N = 22) versus controls (N = 14). After this initial scan, MDD patients were treated with cognitive behavioral therapy (CBT) and scanned again after treatment. Within regions that showed pre-treatment differences between patients and controls, we tested the association between pre-treatment activity and subsequent treatment response as well as activity changes from pre- to post-treatment. This study yielded three main findings. First, prior to treatment and relative to controls, patients exhibited overall reduced activity in the ventromedial prefrontal cortex (PFC), diminished discrimination between emotional and neutral items in the amygdala, caudate, and hippocampus, and enhanced responses to negative versus positive stimuli in the left anterior temporal lobe (ATL) and right dorsolateral PFC. Second, CBT-related symptom improvement in MDD patients was predicted by increased activity at baseline in ventromedial PFC as well as the valence effects in the ATL and dorsolateral PFC. Third, from pre- to post-treatment, MDD patients exhibited overall increases in ventromedial PFC activation, enhanced arousal responses in the amygdala, caudate, and hippocampus, and a reversal of valence effects in the ATL. The study was limited by the relatively small sample that was able to complete both scan sessions, as well as an inability to determine the influence of comorbid disorders within the current sample. Nevertheless, components of the neural networks corresponding to emotion processing disturbances in MDD appear to resolve following treatment and are predictive of treatment response, possibly reflecting improvements in emotion regulation processes in response to CBT.

Gender Differences in Autobiographical Memory for Everyday Events: Retrieval Elicited by SenseCam Images Versus Verbal Cues

Memory (Hove, England). Oct, 2011  |  Pubmed ID: 20981611

Gender differences are frequently observed in autobiographical memory (AM). However, few studies have investigated the neural basis of potential gender differences in AM. In the present functional MRI (fMRI) study we investigated gender differences in AMs elicited using dynamic visual images vs verbal cues. We used a novel technology called a SenseCam, a wearable device that automatically takes thousands of photographs. SenseCam differs considerably from other prospective methods of generating retrieval cues because it does not disrupt the ongoing experience. This allowed us to control for potential gender differences in emotional processing and elaborative rehearsal, while manipulating how the AMs were elicited. We predicted that males would retrieve more richly experienced AMs elicited by the SenseCam images vs the verbal cues, whereas females would show equal sensitivity to both cues. The behavioural results indicated that there were no gender differences in subjective ratings of reliving, importance, vividness, emotion, and uniqueness, suggesting that gender differences in brain activity were not due to differences in these measures of phenomenological experience. Consistent with our predictions, the fMRI results revealed that males showed a greater difference in functional activity associated with the rich experience of SenseCam vs verbal cues, than did females.

Neural Correlates of Confidence During Item Recognition and Source Memory Retrieval: Evidence for Both Dual-process and Strength Memory Theories

Journal of Cognitive Neuroscience. Dec, 2011  |  Pubmed ID: 21736454

Although the medial-temporal lobes (MTL), PFC, and parietal cortex are considered primary nodes in the episodic memory network, there is much debate regarding the contributions of MTL, PFC, and parietal subregions to recollection versus familiarity (dual-process theory) and the feasibility of accounts on the basis of a single memory strength process (strength theory). To investigate these issues, the current fMRI study measured activity during retrieval of memories that differed quantitatively in terms of strength (high vs. low-confidence trials) and qualitatively in terms of recollection versus familiarity (source vs. item memory tasks). Support for each theory varied depending on which node of the episodic memory network was considered. Results from MTL best fit a dual-process account, as a dissociation was found between a right hippocampal region showing high-confidence activity during the source memory task and bilateral rhinal regions showing high-confidence activity during the item memory task. Within PFC, several left-lateralized regions showed greater activity for source than item memory, consistent with recollective orienting, whereas a right-lateralized ventrolateral area showed low-confidence activity in both tasks, consistent with monitoring processes. Parietal findings were generally consistent with strength theory, with dorsal areas showing low-confidence activity and ventral areas showing high-confidence activity in both tasks. This dissociation fits with an attentional account of parietal functions during episodic retrieval. The results suggest that both dual-process and strength theories are partly correct, highlighting the need for an integrated model that links to more general cognitive theories to account for observed neural activity during episodic memory retrieval.

A "snapshot" of Declarative Memory: Differing Developmental Trajectories in Episodic and Autobiographical Memory

Memory (Hove, England). Nov, 2011  |  Pubmed ID: 21942825

Episodic and autobiographical memory are clearly related, yet in both the adult and developmental literatures it is difficult to compare them because of differences in how the constructs are assessed, including differences in content, levels of control, and time since experience. To address these issues, we directly compared children's and adults' autobiographical and episodic memory using the same controlled paradigm. Participants engaged in a photo-taking activity in a museum (autobiographical encoding) and viewed others' photographs of the same museum exhibits (episodic encoding). At test, participants classified photos as ones they took, viewed, or novel. In the autobiographical condition older children and adults performed similarly; younger children's performance was lower than adults'. In contrast, in the episodic condition both groups of children performed more poorly than adults. The findings suggest the developmental primacy of autobiographical relative to episodic memory, and that traditional episodic tasks may underestimate older children's declarative memory abilities.

Where Is ELSA? The Early to Late Shift in Aging

Cerebral Cortex (New York, N.Y. : 1991). Nov, 2011  |  Pubmed ID: 22114083

Studies of cognitive and neural aging have recently provided evidence of a shift from an early- to late-onset cognitive control strategy, linked with temporally extended activity in the prefrontal cortex (PFC). It has been uncertain, however, whether this age-related shift is unique to PFC and executive control tasks or whether the functional location might vary depending on the particular cognitive processes that are altered. The present study tested whether an early-to-late shift in aging (ELSA) might emerge in the medial temporal lobes (MTL) during a protracted context memory task comprising both anticipatory cue (retrieval preparation) and retrieval probe (retrieval completion) phases. First, we found reduced MTL activity in older adults during the early retrieval preparation phase coupled with increased MTL activity during the late retrieval completion phase. Second, we found that functional connectivity between MTL and PFC regions was higher during retrieval preparation in young adults but higher during retrieval completion in older adults, suggesting an important interactive relationship between the ELSA pattern in MTL and PFC. Taken together, these results critically suggest that aging results in temporally lagged activity even in regions not typically associated with cognitive control, such as the MTL.

The Architecture of Cross-hemispheric Communication in the Aging Brain: Linking Behavior to Functional and Structural Connectivity

Cerebral Cortex (New York, N.Y. : 1991). Jan, 2012  |  Pubmed ID: 21653286

Contralateral recruitment remains a controversial phenomenon in both the clinical and normative populations. To investigate the neural correlates of this phenomenon, we explored the tendency for older adults to recruit prefrontal cortex (PFC) regions contralateral to those most active in younger adults. Participants were scanned with diffusion tensor imaging and functional magnetic rresonance imaging during a lateralized word matching task (unilateral vs. bilateral). Cross-hemispheric communication was measured behaviorally as greater accuracy for bilateral than unilateral trials (bilateral processing advantage [BPA]) and at the neural level by functional and structural connectivity between contralateral PFC. Compared with the young, older adults exhibited 1) greater BPAs in the behavioral task, 2) greater compensatory activity in contralateral PFC during the bilateral condition, 3) greater functional connectivity between contralateral PFC during bilateral trials, and 4) a positive correlation between fractional anisotropy in the corpus callosum and both the BPA and the functional connectivity between contralateral PFC, indicating that older adults' ability to distribute processing across hemispheres is constrained by white matter integrity. These results clarify how older adults' ability to recruit extra regions in response to the demands of aging is mediated by existing structural architecture, and how this architecture engenders corresponding functional changes that allow subjects to meet those task demands.