Although the perirhinal cortex (PRC), parahippocampal cortex (PHC) and retrosplenial cortex (RSC) have an essential role in memory, the precise functions of these areas are poorly understood. Here, we review the anatomical and functional characteristics of these areas based on studies in humans, monkeys and rats. Our Review suggests that the PRC and PHC-RSC are core components of two separate large-scale cortical networks that are dissociable by neuroanatomy, susceptibility to disease and function. These networks not only support different types of memory but also appear to support different aspects of cognition.

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.

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Emotional events are remembered better than neutral events, and this emotion advantage becomes particularly pronounced over time. The time dependent effects of emotion impact recollection rather than familiarity-based recognition, and they influence recollection of item-specific details rather than contextual details. Moreover, the amygdala, but not the hippocampus, is critical in producing these effects. Time-dependent effects of emotion have been attributed to an emotional consolidation process whereby the amygdala gradually facilitates the storage of emotional memories by other medial temporal lobe regions. However, here we propose that these effects can be better understood by an emotional binding account whereby the amygdala mediates the recollection of item-emotion bindings that are forgotten more slowly than item-context bindings supported by the hippocampus.

Reward motivation is known to modulate memory encoding, and this effect depends on interactions between the substantia nigra/ ventral tegmental area complex (SN/VTA) and the hippocampus. It is unknown, however, whether these interactions influence offline neural activity in the human brain that is thought to promote memory consolidation. Here, we used functional magnetic resonance imaging (fMRI) to test the effect of reward motivation on post-learning neural dynamics and subsequent memory for objects that were learned in high- or low-reward motivation contexts. We found that post-learning increases in resting-state functional connectivity between the SN/VTA and hippocampus predicted preferential retention of objects that were learned in high-reward contexts. In addition, multivariate pattern classification revealed that hippocampal representations of high-reward contexts were preferentially reactivated during post-learning rest, and the number of hippocampal reactivations was predictive of preferential retention of items learned in high-reward contexts. These findings indicate that reward motivation alters offline post-learning dynamics between the SN/VTA and hippocampus, providing novel evidence for a potential mechanism by which reward could influence memory consolidation.

Over the past decade, fMRI techniques have been increasingly used to interrogate the neural correlates of successful emotional memory encoding. These investigations have typically aimed to either characterize the contributions of the amygdala and medial temporal lobe (MTL) memory system, replicating results in animals, or delineate the neural correlates of specific behavioral phenomena. It has remained difficult, however, to synthesize these findings into a systems neuroscience account of how networks across the whole brain support the enhancing effects of emotion on memory encoding. To this end, the present study employed a meta-analytic approach using activation likelihood estimates to assess the anatomical specificity and reliability of event-related fMRI activations related to successful memory encoding for emotional versus neutral information. The meta-analysis revealed consistent clusters within bilateral amygdala, anterior hippocampus, anterior and posterior parahippocampal gyrus, the ventral visual stream, left lateral prefrontal cortex and right ventral parietal cortex. The results within the amygdala and MTL support a wealth of findings from the animal literature linking these regions to arousal-mediated memory effects. The consistency of findings in cortical targets, including the visual, prefrontal, and parietal cortices, underscores the importance of generating hypotheses regarding their participation in emotional memory formation. In particular, we propose that the amygdala interacts with these structures to promote enhancements in perceptual processing, semantic elaboration, and attention, which serve to benefit subsequent memory for emotional material. These findings may motivate future research on emotional modulation of widespread neural systems and the implications of this modulation for cognition.

The authors manipulated emotion regulation strategies at encoding and administered explicit and implicit memory tests. In Experiment 1, participants used reappraisal to enhance and decrease the personal relevance of unpleasant and neutral pictures. In Experiment 2, decrease cues were replaced with suppress cues that directed participants to inhibit emotion-expressive behavior. Across experiments, using reappraisal to enhance the personal relevance of pictures improved free recall. By contrast, attempting to suppress emotional displays tended to impair recall, especially compared to the enhance condition. Using reappraisal to decrease the personal relevance of pictures had different effects depending on picture type. Paired with unpleasant pictures, the decrease cue tended to improve recall. Paired with neutral stimuli, the decrease cue tended to impair recall. Emotion regulation did not affect perceptual priming. Results highlight dissociable effects of emotion regulation on explicit and implicit memory, as well as dissociations between regulation strategies with respect to explicit memory.Keywords: affect, cognitive control, emotion, emotion regulation, memory On a daily basis, lives are shaped by emotions, including joy at successes, sadness due to losses, and fear in the face of threats. However, emotions are often not experienced passively. Instead, individuals engage in emotion regulation (Gross, Richards, & John, 2006), attempting to modulate the behavioral, experiential, or physiological components of emotions (Gross, 1998b). Successful emotion regulation is associated with positive outcomes, including development of social competence (Eisenberg, Fabes, Guthrie, & Reiser, 2000) and improved subjective well-being (Larsen & Prizmic, 2004), whereas emotion dysregulation figures prominently in psychopathology (Gross & Munoz, 1995) and may be a precursor to problematic behaviors, including violence (Davidson, Putnam, & Larson, 2000).Various emotion regulation strategies have different behavioral effects, as demonstrated by investigations of reappraisal and expressive suppression. Reappraisal refers to cognitive attempts to change the meaning of stimuli, while expressive suppression refers to inhibition of emotionally expressive behavior (Gross, 1998b). Several studies have examined the effects of these strategies on stimulus encoding. Reappraising gruesome films as less distressing reduces self-reported negative emotion and physiological arousal (Lazarus & Alfert, 1964). Furthermore, using reappraisal to enhance and decrease responses to unpleasant pictures leads to increased and reduced startle responses, respectively (Jackson, Malmstadt, Larson, & Davidson, 2000), as well as increased and reduced activity in the amgydala (Ochsner et al., 2004). Less is known about the neural correlates of expressive suppression, but this strategy leads to increased sympathetic nervous system activity and does not effectively modulate unpleasant emotional experience (Gross, 1998a;Gross & Levenson, 1993.Thus, reappraisal and e...

a b s t r a c tMajor 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 ¼ 22) versus controls (N ¼ 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.