[show abstract][hide abstract] ABSTRACT: Although it is well established that the perirhinal cortex (PRC) makes an important contribution to recognition memory, the specific nature of this contribution remains uncertain. The finding that PRC activity is reduced for old compared with new items is typically attributed to the recovery of a long-term memory (LTM) signal. However, because old items are processed more easily or fluently than new items, reduced PRC activity could reflect increased fluency rather than LTM retrieval per se. We tested this hypothesis in humans using fMRI and a well-validated method to manipulate fluency: the masked priming paradigm. Some words during an old-new recognition test were preceded by conceptually related words (primes) that were subliminally presented (masked). The behavioral results replicated previous findings using this paradigm, whereby the fluency manipulation increased "oldness" responses to both old and new items. The fMRI analyses yielded two main sets of results. First, in the case of new items, which are independent from LTM retrieval, masked priming reduced PRC activity and predicted behavioral misattribution of fluency to oldness. Second, in the case of old items, the same PRC region showing fluency-related reductions for new items also contributed to "old" responding to old items. Individual differences in PRC attenuation also predicted oldness ratings to old items, and fluency modulated PRC connectivity with other brain regions associated with processing oldness signals, including visual cortex and right lateral prefrontal cortex. These results support a broader view in which the PRC serves a function more general than memory.
Journal of Neuroscience 09/2013; 33(36):14466-74. · 6.91 Impact Factor
[show abstract][hide abstract] ABSTRACT: 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 contribition of the PFC regions and the architecture of memory.
[show abstract][hide abstract] ABSTRACT: Restudying material is a common method for learning new information, but not necessarily an effective one. Research on the testing effect shows that practice involving retrieval from memory can facilitate later memory in contrast to passive restudy. Despite extensive behavioral work, the brain processes that make retrieval an effective learning strategy remain unclear. In the present experiment, we explored how initially retrieving items affected memory a day later as compared to a condition involving traditional restudy. In contrast to restudy, initial testing that contributed to future memory success was associated with engagement of several regions including the anterior hippocampus, lateral temporal cortices, and medial prefrontal cortex (PFC). Additionally, testing enhanced hippocampal connectivity with ventrolateral PFC and midline regions. These findings indicate that the testing effect may be contingent on processes that are typically thought to support memory success at encoding (e.g. relational binding, selection and elaboration of semantically-related information) in addition to those more often associated with retrieval (e.g. memory search).
[show abstract][hide abstract] ABSTRACT: A fundamental question in the emotional memory literature is why emotion enhances memory in some conditions but disrupts memory in other conditions. For example, separate studies have shown that emotional stimuli tend to be better remembered in long-term episodic memory (EM), whereas emotional distracters tend to impair working memory (WM) maintenance. The first goal of this study was to directly compare the neural correlates of EM enhancement (EME) and WM impairing (WMI) effects, and the second goal was to explore individual differences in these mechanisms. During event-related functional magnetic resonance imaging (fMRI), participants maintained faces in WM while being distracted by emotional or neutral pictures presented during the delay period. EM for the distracting pictures was tested after scanning and was used to identify successful encoding activity for the picture distracters. The first goal yielded two findings: (1) emotional pictures that disrupted face WM but enhanced subsequent EM were associated with increased amygdala (AMY) and hippocampal activity (ventral system) coupled with reduced dorsolateral PFC (dlPFC) activity (dorsal system); (2) trials in which emotion enhanced EM without disrupting WM were associated with increased ventrolateral PFC activity. The ventral-dorsal switch can explain EME and WMI, while the ventrolateral PFC effect suggests a coping mechanism. The second goal yielded two additional findings: (3) participants who were more susceptible to WMI showed greater amygdala increases and PFC reductions; (4) AMY activity increased and dlPFC activity decreased with measures of attentional impulsivity. Taken together, these results clarify the mechanisms linking the enhancing and impairing effects of emotion on memory, and provide insights into the role of individual differences in the impact of emotional distraction.
[show abstract][hide abstract] ABSTRACT: In the 1980s and 1990s, there was a major theoretical debate in the memory domain regarding the multiple memory systems and processing modes frameworks. The components of processing framework argued for a middle ground: Instead of neatly divided memory systems or processing modes, this framework proposed the existence of numerous processing components that are recruited in different combinations by memory tasks and yield complex patterns of associations and dissociations. Because behavioral evidence was not sufficient to decide among these three frameworks, the debate was largely abandoned. However, functional neuroimaging evidence accumulated during the last two decades resolves the stalemate, because this evidence is more consistent with the components framework than with the other two frameworks. For example, functional neuroimaging evidence shows that brain regions attributed to one memory system can contribute to tasks associated with other memory systems and that brain regions attributed to the same processing mode (perceptual or conceptual) can be dissociated from each other. Functional neuroimaging evidence suggests that memory processes are supported by transient interactions between a few regions called process-specific alliances. These conceptual developments are an example of how functional neuroimaging can contribute to theoretical debates in cognitive psychology.
Perspectives on Psychological Science 01/2013; 8(1):49-55. · 4.89 Impact Factor
[show abstract][hide abstract] ABSTRACT: A fundamental principle in memory research is that memory is a function of the similarity between encoding and retrieval operations. Consistent with this principle, many neurobiological models of declarative memory assume that memory traces are stored in cortical regions, and the hippocampus facilitates the reactivation of these traces during retrieval. The present investigation tested the novel prediction that encoding-retrieval similarity can be observed and related to memory at the level of individual items. Multivariate representational similarity analysis was applied to functional magnetic resonance imaging data collected during encoding and retrieval of emotional and neutral scenes. Memory success tracked fluctuations in encoding-retrieval similarity across frontal and posterior cortices. Importantly, memory effects in posterior regions reflected increased similarity between item-specific representations during successful recognition. Mediation analyses revealed that the hippocampus mediated the link between cortical similarity and memory success, providing crucial evidence for hippocampal-cortical interactions during retrieval. Finally, because emotional arousal is known to modulate both perceptual and memory processes, similarity effects were compared for emotional and neutral scenes. Emotional arousal was associated with enhanced similarity between encoding and retrieval patterns. These findings speak to the promise of pattern similarity measures for evaluating memory representations and hippocampal-cortical interactions.
[show abstract][hide abstract] ABSTRACT: Although ventral parietal cortex (VPC) activations can be found in a variety of cognitive domains, these activations have been typically attributed to cognitive operations specific to each domain. In this article, we propose a hypothesis that can account for VPC activations across all the cognitive domains reviewed. We first review VPC activations in the domains of perceptual and motor reorienting, episodic memory retrieval, language and number processing, theory of mind, and episodic memory encoding. Then, we consider the localization of VPC activations across domains and conclude that they are largely overlapping with some differences around the edges. Finally, we assess how well four different hypotheses of VPC function can explain findings in various domains and conclude that a bottom-up attention hypothesis provides the most complete and parsimonious account.
Trends in Cognitive Sciences 05/2012; 16(6):338-52. · 16.01 Impact Factor
[show abstract][hide abstract] ABSTRACT: Abstract The article "More than a feeling: Pervasive influences of memory without awareness of retrieval" reviews evidence from ERP studies of recognition memory that the FN400 effect typically ascribed to familiarity may index implicit memory that occurs during recognition testing. We find their argument compelling, and contend that this potential "implicit contamination" is not unique to ERP studies. We suggest an analogous problem affecting fMRI studies, focusing particularly on the perirhinal cortex. Resolving this issue is critical for understanding the relationship between memory and the medial temporal lobes.
[show abstract][hide abstract] ABSTRACT: 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.
Neurobiology of aging 12/2011; 32(12):2318.e17-30. · 5.94 Impact Factor
[show abstract][hide abstract] ABSTRACT: 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.
[show abstract][hide abstract] ABSTRACT: 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.
Journal of Cognitive Neuroscience 07/2011; 23(12):3959-71. · 4.49 Impact Factor
[show abstract][hide abstract] ABSTRACT: 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.
[show abstract][hide abstract] ABSTRACT: 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.
Journal of Cognitive Neuroscience 06/2011; 23(6):1275-84. · 4.49 Impact Factor
[show abstract][hide abstract] ABSTRACT: 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).
Journal of Cognitive Neuroscience 05/2011; 23(11):3209-17. · 4.49 Impact Factor
[show abstract][hide abstract] ABSTRACT: Emotional events tend to be better remembered than non-emotional events. One goal of cognitive and affective neuroscientists is to understand the neural mechanisms underlying this enhancing effect of emotion on memory. A method that has proven particularly influential in the investigation of the memory-enhancing effect of emotion is the so-called subsequent memory paradigm (SMP). This method was originally used to investigate the neural correlates of non-emotional memories, and more recently we and others also applied it successfully to studies of emotional memory (reviewed in). Here, we describe a protocol that allows investigation of the neural correlates of the memory-enhancing effect of emotion using the SMP in conjunction with event-related functional magnetic resonance imaging (fMRI). An important feature of the SMP is that it allows separation of brain activity specifically associated with memory from more general activity associated with perception. Moreover, in the context of investigating the impact of emotional stimuli, SMP allows identification of brain regions whose activity is susceptible to emotional modulation of both general/perceptual and memory-specific processing. This protocol can be used in healthy subjects, as well as in clinical patients where there are alterations in the neural correlates of emotion perception and biases in remembering emotional events, such as those suffering from depression and post-traumatic stress disorder (PTSD).
[show abstract][hide abstract] ABSTRACT: This video and the associated PDF (Shafer et al., 2011) illustrate an experimental protocol that allows investigation of the neural correlates of the memory-enhancing effect of emotion, using fMRI.
[show abstract][hide abstract] ABSTRACT: 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.
Annals of the New York Academy of Sciences 03/2011; 1224(1):174 - 190. · 4.38 Impact Factor