FMRI signals associated with memory strength in the medial temporal lobes: a meta-analysis

To identify patterns of memory-related neural activity in the medial temporal lobes (MTL), a quantitative meta-analysis of 17 functional magnetic resonance imaging (fMRI) studies was performed. The analysis shows that increased activity in the hippocampus and the parahippocampal cortex predicts subsequent memory strength. During retrieval, activity in the hippocampus increases in association with strong memory. In the perirhinal cortex, increased activity predicts subsequent recognition, whether based on weak or strong memory, whereas during retrieval activity decreases below the level for misses in association with both weak and strong memory. The results are consistent with the claim that the hippocampus selectively subserves recollection, whereas adjacent structures subserve familiarity [Eichenbaum, H., Yonelinas, A., & Ranganath, C. (2007). The medial temporal lobe and recognition memory. The Annual Review of Neuroscience, 30, 123-152]. However, this conclusion depends on a specific dual-process theory of recognition memory that has been used to interpret the results. An alternative dual-process model holds that the behavioral methods used to differentiate recollection from familiarity instead separate strong memories from weak memories. When the fMRI data are interpreted in terms of the alternative theory, the fMRI results do not point to selective roles for the hippocampus or the adjacent MTL structures. The fMRI data alone cannot distinguish between these two models, so other methods are needed to resolve the issue.     

Semantic memory recognition is supported by intrinsic recollection-like processes: “The butcher on the bus” revisited

Dual-process models suggest that recognition memory is independently supported by recollection and familiarity. Current theories attribute recollection solely to hippocampally mediated episodic memory (EM), and familiarity to both episodic and semantic memory (SM) supported by medial temporal lobe cortex (MTLC) and prefrontal cortex. We tested whether, contrary to this view, recollection-like processes also intrinsically support SM recognition and whether MTL structures are involved in their execution. A semantic Process Dissociation Procedure (PDP) with famous and non-famous names was used in three experiments. Experiment 1 revealed that recollection-like processes in semantic memory were not associated with episodic memory for the public events, were predicted by performance on standard SM tasks and were independent of EM tasks, suggesting they are intrinsic to SM. Experiment 2 demonstrated the independence of the two process estimates by showing only familiarity was affected by shifting response criterion while only recollection estimates were significantly altered under divided-attention. Finally experiment 3 tested amnesic patients with varying degrees of hippocampal and MTLC damage. Despite normal overall fame recognition performance, recollection estimates were specifically affected by MTL damage. When damage was primarily hippocampal, only retrograde recollection estimates were reduced, while more extensive MTLC damage led to both retrograde and anterograde recollection deficits. We conclude that recognition of semantic information is supported by at least two independent processes akin to the ones that support EM recognition. Recollection-like processes are intrinsic to SM and likely do not reflect EM contribution to SM performance. Together with previous studies of recollection in remote memory, these data suggest that recollection is not a unitary phenomenon. In EM it involves autonoetic re-experiencing, and is supported by interaction of fronto-temporal networks; in EM and SM it supports retrieval of contextual/associative information regardless of consciousness type, and is dependent on intact MTL function. Familiarity processes and neural substrates may also differ between lifetime familiarity and within-session familiarity.     

fMRI evidence for separable and lateralized prefrontal memory monitoring processes

Source memory research suggests that attempting to remember specific contextual aspects surrounding prior stimulus encounters results in greater left prefrontal cortex (PFC) activity than simple item-based old/new recognition judgments. Here, we tested a complementary hypothesis that predicts increases in the right PFC with tasks requiring close monitoring of item familiarity. More specifically, we compared a judgment of frequency (JOF) task to an item memory task, in which the former required estimating the number of previous picture encounters and the latter required discriminating old from new exemplars of previously seen items. In comparison to standard old/new recognition, both source memory and the JOF task examined here require more precise mnemonic judgments. However, in contrast to source memory, cognitive models suggest the JOF task relies heavily upon item familiarity, not specific contextual recollections. Event-related fMRI demonstrated greater recruitment of right, not left, dorsolateral and frontopolar PFC regions during the JOF compared to item memory task. These data suggest a role for right PFC in the close monitoring of the familiarity of objects, which becomes critical when contextual recollection is ineffective in satisfying a memory demand.     

Lateralization of prefrontal activity during episodic memory retrieval: evidence for the production-monitoring hypothesis

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.     

Associative reinstatement: A novel approach to assessing associative memory in patients with unilateral temporal lobe excisions

We investigated whether unilateral medial temporal lobe (MTL) damage disrupts associative reinstatement, which represents the gain in item memory when the studied associative information is reinstated at retrieval. We were interested to see whether associative reinstatement relies on the same relational binding operations that support other types of associative memory (associative identification and recollection) thought to be subserved by the MTL. In addition, we examined whether such damage affects the different types of associative memory to a greater extent than item memory and item familiarity, and whether a different pattern is seen in patients with language dominant relative to non-dominant temporal lobe resection when the studied material consists of verbal information. To do so, we used a word pair recognition paradigm composed of two tasks: (1) a pair recognition task that provides measures of associative reinstatement and item memory, and (2) an associative identification recognition task that provides a measure of associative identification memory. Estimates of item familiarity and recollection were derived from performance on both tasks using a variant of the process-dissociation procedure. Our results showed that associative reinstatement, like other types of associative memory measures, was impaired in patients with unilateral resection, irrespective of the side of damage. Item familiarity, however, was impaired solely following language dominant resection. The lack of a laterality effect in our relational measures was likely due to using an encoding task that promoted formation of both verbal and visual associations, whereas item-based familiarity could rely exclusively on verbal operations. We propose that associative reinstatement provides a sensitive measure of relational memory that is less dependent on strategic processing and therefore more appropriate for evaluating MTL function in patients.     

A single‐trace dual‐process model of episodic memory: A novel computational account of familiarity and recollection

Dual‐process theories of episodic memory state that retrieval is contingent on two independent processes: familiarity (providing a sense of oldness) and recollection (recovering events and their context). A variety of studies have reported distinct neural signatures for familiarity and recollection, supporting dual‐process theory. One outstanding question is whether these signatures reflect the activation of distinct memory traces or the operation of different retrieval mechanisms on a single memory trace. We present a computational model that uses a single neuronal network to store memory traces, but two distinct and independent retrieval processes access the memory. The model is capable of performing familiarity and recollection‐based discrimination between old and new patterns, demonstrating that dual‐process models need not to rely on multiple independent memory traces, but can use a single trace. Importantly, our putative familiarity and recollection processes exhibit distinct characteristics analogous to those found in empirical data; they diverge in capacity and sensitivity to sparse and correlated patterns, exhibit distinct ROC curves, and account for performance on both item and associative recognition tests. The demonstration that a single‐trace, dual‐process model can account for a range of empirical findings highlights the importance of distinguishing between neuronal processes and the neuronal representations on which they operate. © 2009 Wiley‐Liss, Inc.     

Pre-experimental familiarization increases hippocampal activity for both targets and lures in recognition memory: an fMRI study

In the present study, items pre-exposed in a familiarization series were included in a list discrimination task to manipulate memory strength. At test, participants were required to discriminate strong targets and strong lures from weak targets and new lures. This resulted in a concordant pattern of increased "old" responses to strong targets and lures. Model estimates attributed this pattern to either equivalent increases in memory strength across the two types of items (unequal variance signal detection model) or equivalent increases in both familiarity and recollection (dual process signal detection [DPSD] model). Hippocampal activity associated with strong targets and lures showed equivalent increases compared with missed items. This remained the case when analyses were restricted to high-confidence responses considered by the DPSD model to reflect predominantly recollection. A similar pattern of activity was observed in parahippocampal cortex for high-confidence responses. The present results are incompatible with "noncriterial" or "false" recollection being reflected solely in inflated DPSD familiarity estimates and support a positive correlation between hippocampal activity and memory strength irrespective of the accuracy of list discrimination, consistent with the unequal variance signal detection model account.     

Medial temporal lobe function and recognition memory: a novel approach to separating the contribution of recollection and familiarity

Human neuroimaging studies of recognition memory have often been interpreted to mean that the hippocampus supports recollection but not familiarity. This interpretation is complicated by the fact that recollection-based decisions are typically associated with stronger memories than familiarity-based decisions. Some studies of source memory controlled for this difference in memory strength and found that hippocampal activity during learning predicted subsequent item memory strength while recollection-based memory (performance on source memory questions) was held at chance. This result suggests that the hippocampus is important for familiarity. However, a difficulty with this approach is that when source memory is assessed by asking specific, task-relevant source memory questions, participants who fail to answer the prescribed questions might nevertheless have available other (task-irrelevant) source information. Accordingly, successful item memory could still be associated with recollection. The present study used a novel method to assess item memory and source memory. Instead of responding to specific source questions, participants rated their source memory strength based on any information about the learning episode that was available to them. When subsequent source memory strength was held constant at the lowest possible level, we identified regions bilaterally in hippocampus, as well as in perirhinal cortex, where activity during learning increased as subsequent item memory increased in strength. In addition, activity in cortical regions (including prefrontal cortex) was related to source memory success independently of item memory strength. These findings suggest that activity in the hippocampus is related to the encoding of familiarity-based item memory, independent of subsequent recollection-based success.     

Characterizing episodic memory retrieval: electrophysiological evidence for diminished familiarity following unitization

Episodic memory relies on both recollection and familiarity; why these processes are differentially engaged during retrieval remains unclear. Traditionally, recollection has been considered necessary for tasks requiring associative retrieval, whereas familiarity supports recognition of items. Recently, however, familiarity has been shown to contribute to associative recognition if stimuli are "unitized" at encoding (a single representation is created from multiple elements)-the "benefit" of unitization. Here, we ask if there is also a "cost" of unitization; are the elements of unitized representations less accessible via familiarity? We manipulated unitization during encoding and used ERPs to index familiarity and recollection at retrieval. The data revealed a selective reduction in the neural correlate of familiarity for individual words originally encoded in unitized compared with nonunitized word pairs. This finding reveals a measurable cost of unitization, suggesting that the nature of to-be-remembered stimuli is critical in determining whether familiarity contributes to episodic memory.     

Sustained and transient neural modulations in prefrontal cortex related to declarative long-term memory, working memory, and attention

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.     

The medial temporal lobes distinguish between within‐item and item‐context relations during autobiographical memory retrieval

During autobiographical memory retrieval, the medial temporal lobes (MTL) relate together multiple event elements, including object (within‐item relations) and context (item‐context relations) information, to create a cohesive memory. There is consistent support for a functional specialization within the MTL according to these relational processes, much of which comes from recognition memory experiments. In this study, we compared brain activation patterns associated with retrieving within‐item relations (i.e., associating conceptual and sensory‐perceptual object features) and item‐context relations (i.e., spatial relations among objects) with respect to naturalistic autobiographical retrieval. We developed a novel paradigm that cued participants to retrieve information about past autobiographical events, non‐episodic within‐item relations, and non‐episodic item‐context relations with the perceptuomotor aspects of retrieval equated across these conditions. We used multivariate analysis techniques to extract common and distinct patterns of activity among these conditions within the MTL and across the whole brain, both in terms of spatial and temporal patterns of activity. The anterior MTL (perirhinal cortex and anterior hippocampus) was preferentially recruited for generating within‐item relations later in retrieval whereas the posterior MTL (posterior parahippocampal cortex and posterior hippocampus) was preferentially recruited for generating item‐context relations across the retrieval phase. These findings provide novel evidence for functional specialization within the MTL with respect to naturalistic memory retrieval. © 2015 Wiley Periodicals, Inc.     

Induced electroencephalogram oscillations during source memory: familiarity is reflected in the gamma band, recollection in the theta band

Modulations of oscillatory electroencephalogram (EEG) activity in the induced gamma and theta frequency ranges (induced gamma and theta band responses; iGBRs: >30 Hz; iTBRs: approximately 6 Hz) have been associated with retrieval of information from long-term memory. However, the specific functional role of these two forms of oscillatory activity remains unclear. The present study examines theta- and gamma-oscillations within a dual-process framework, which defines "familiarity" and "recollection" as the two component processes of recognition memory. During encoding, participants were instructed to make "bigger/smaller than a shoebox" or "living/nonliving" decisions for different object pictures. During retrieval "old/new" recognition was followed (for items judged old) by a source discrimination task regarding the decision made for each item at encoding. iGBRs (35-80 Hz; 210-330 msec) were higher for correctly identified "old" relative to "new" objects. Importantly, they did not distinguish between successful and unsuccessful source judgments. In contrast, iTBRs (4-7.5 Hz; 600-1200 msec) were sensitive to source discrimination. We propose that iGBRs mirror early associative processes linked to familiarity-related retrieval processes, whereas iTBRs reflect later onsetting, episodic, recollection-related mechanisms.     

Memory orientation and success: separable neurocognitive components underlying episodic recognition

Episodic recognition can be based on recollection of contextual details, on a sense of recent encounter, or some combination of the two. According to several cognitive models, selectively attending to these distinct aspects of memory may require different retrieval orientations and result in different neural responses depending upon whether or not retrieval is successful. Using event-related fMRI, we examined retrieval orientation by having subjects discriminate between two test words in one of two manners. During source recollection, they selected the member of the pair previously associated with a particular encoding task. In contrast, recency judgment required selection of the most recently encountered item of the pair, regardless of how it had been encoded. Furthermore, successful and unsuccessful trials within each retrieval task were contrasted to determine whether retrieval success effects occurred in overlapping or dissimilar neural populations compared to those associated with each retrieval orientation. The results revealed distinct lateral prefrontal and parietal activations that distinguished attempted source recollection from judgments of relative recency; these orientation effects were largely independent of retrieval success. In contrast, medial temporal lobe structures (hippocampus and parahippocampal gyrus) were differentially more active during successful recollection of encoding context, showing similar reduced responses during failed source recollection and judgments of recency. These results indicate that different memory orientations recruit distinct prefrontal and parietal networks and that the recovery of episodic context is associated with the hippocampus and surrounding medial temporal cortices.     

Contribution of familiarity and recollection to associative recognition memory: insights from event-related potentials

Within the dual-process perspective of recognition memory, it has been claimed that familiarity is sufficient to support recognition of single items, but recollection is necessary for associative recognition of item pairs. However, there are some reports suggesting that familiarity might support associative recognition judgments when the items form an easy to access bound representation. In contrast, recollection seems to be required for the recognition of bindings that might be flexibly rearranged in novel situations. We investigated whether both forms of binding are mediated by different mechanisms as reflected by a qualitatively different spatiotemporal eventrelated potential (ERP) pattern. In a recognition memory experiment, subjects gave old/new judgments to words learned by focusing either on interitem associations or on size relation of word triplets. Results revealed higher hit rates in the relational condition as compared to the associative condition. In addition, the proportion of triplets from which all three items were remembered was significantly larger in the relational condition suggesting that memory retrieval in this condition relies primarily on bound representations of word triplets. The ERP revealed a late parietal old/new effect for both conditions, with relational processing resulting in a greater effect. In contrast, an early frontal old/new effect was solely present in the associative condition. Taken together, these data provide evidence that familiarity might support associative recognition if the associated components are coherently encoded into a bound representation. Recollection might foster the recognition of relational bindings among items. This indicates that the contribution of familiarity and recollection to associative recognition depends on the kind of binding operations performed on the items rather than on the single versus multiple item distinction.     

Impairment of verbal recollection following ischemic damage to the right anterior hippocampus

Damage to the left medial temporal lobe (MTL) leads to an impairment of verbal recognition memory, affecting both the process of conscious recollection and familiarity-based recognition. Neuroimaging evidence, on the other hand, suggests a bilateral MTL contribution to verbal recollection. We investigated verbal recognition memory in three patients with focal ischemic lesions to the right MTL. The dual-process signal detection model and the process-dissociation procedure were applied to assess the contributions of recollection and familiarity to recognition memory. Compared to a group of 27 healthy age-matched controls, patients were impaired at recollection while familiarity was intact, and this effect was found for both estimation procedures. Detailed single-case analyses confirmed this pattern in two of the three right MTL patients. The findings suggest that, when task demands are high, as during recollective recognition, the right anterior hippocampus may also contribute to verbal recollection, thereby confirming neuroimaging evidence of a joint involvement of the left and the right MTL in verbal recollection.     

Recognition memory for single items and for associations in amnesic patients

Recognition memory performance reflects two distinct processes or types of memory referred to as recollection and familiarity. According to theoretical claims about the two types of memory, single item and associative recognition tasks can be used as an experimental method to distinguish recollection and familiarity processes. Associative recognition decisions can be used as an index of recollection while memory for single items is mostly based on familiarity judgement. We employed this procedure to examine a possible dissociation in the memory performance of amnesic patients between spared single item and impaired associative recognition. Twelve amnesic patients, six with damage confined to the hippocampus proper, and six with damage elsewhere in the brain, were recruited for the present study. The findings showed that hippocampal amnesics exhibit relative sparing of single item learning but are consistently deficient in the learning of all kinds of between-item associations. These results are consistent with the view that hippocampal formation contributes differently to declarative tasks that require recollective or familiarity processes.     

Does repetition engender the same retrieval processes in young and older adults?

Aging differentially affects retrieval processes underlying recognition memory: familiarity is maintained, whereas recollection declines. We determined whether word repetition across two study-test phases enhanced older adults' use of recollection. During Test 1, frontal episodic memory effects, suggestive of familiarity-based processes, were age invariant, whereas only the young showed a parietal episodic memory effect, suggestive of recollection. Repetition did not modulate the frontal episodic memory effect in either group, but increased the parietal episodic memory effect in the young. Importantly, older adults showed a parietal episodic memory effect at Test 2, suggesting that repetition did enable recollection. Only older adults, however, showed a left frontal negativity, implying that they may have used additional processes to recover episodic memories.     

Selective and shared contributions of the hippocampus and perirhinal cortex to episodic item and associative encoding

Although the general role of the medial-temporal lobe (MTL) in episodic memory is well established, controversy surrounds the precise division of labor between distinct MTL subregions. The perirhinal cortex (PrC) has been hypothesized to support nonassociative item encoding that contributes to later familiarity, whereas the hippocampus supports associative encoding that selectively contributes to later recollection. However, because previous paradigms have predominantly used recollection of the item context as a measure of associative encoding, it remains unclear whether recollection of different kinds of episodic detail depends on the same or different MTL encoding operations. In our current functional magnetic resonance imaging study, we devised a subsequent memory paradigm that assessed successful item encoding in addition to the encoding of two distinct episodic details: an item-color and an item-context detail. Hippocampal encoding activation was selectively enhanced during trials leading to successful recovery of either an item-color or item-context association. Moreover, the magnitude of hippocampal activation correlated with the number, and not the kind, of associated details successfully bound, providing strong evidence for a role of the hippocampus in domain-general associative encoding. By contrast, PrC encoding activation correlated with both nonassociative item encoding as well as associative item-color binding, but not with item-context binding. This pattern suggests that the PrC contributions to memory encoding may be domain-specific and limited to the binding of items with presented item-related features. Critically, together with a separately conducted behavioral study, these data raise the possibility that PrC encoding operations -- in conjunction with hippocampal mechanisms -- contribute to later recollection of presented item details.     

The role of recollection and familiarity in the functional differentiation of the medial temporal lobes

The components of the medial temporal lobes (MTL) receive different kinds of input. The perirhinal cortex receives primarily object/item information, the parahippocampal cortex receives contextual information, and the hippocampus receives high-level inputs that include object/item, context, and other information. Critically, the perirhinal and parahippocampal cortices have similar cytoarchitectonics, which differ considerably from that of the hippocampus and suggest that these cortices process their inputs differently from the way that the hippocampus processes its inputs. Much evidence indicates that the hippocampus is designed to rapidly bind together pattern-separated representations that support recall/recollection well. In contrast, the newer MTL cortices rapidly create poorly pattern-separated memories that support familiarity well, but recall/recollection very poorly. For over a decade, there has been disagreement about whether recall/recollection is primarily mediated by the hippocampus and familiarity by the evolutionarily newer MTL cortices or whether the MTL mediates these kinds of memory in an integrated, homogeneous fashion. Common misconceptions about familiarity, recollection, item, and associative memory are discussed as are methodological problems with MTL lesion and functional imaging research. The possible confound of familiarity with weaker memory and recollection with stronger memory is discussed and the implications of the Montaldi et al. (2006) functional Magnetic Resonance Imaging (fMRI) study, which matched memory strength between strong familiarity and recollection, finding that only recollection activated the hippocampus, are discussed. A suggestion is made about how the long-running conflict of findings in the human hippocampal lesion literature may be resolved.     

Event-related fMRI studies of episodic encoding and retrieval: Meta-analyses using activation likelihood estimation

The recent surge in event-related fMRI studies of episodic memory has generated a wealth of information about the neural correlates of encoding and retrieval processes. However, interpretation of individual studies is hampered by methodological differences, and by the fact that sample sizes are typically small. We submitted results from studies of episodic memory in healthy young adults, published between 1998 and 2007, to a voxel-wise quantitative meta-analysis using activation likelihood estimation [Laird, A. R., McMillan, K. M., Lancaster, J. L., Kochunov, P., Turkeltaub, P. E., & Pardo, J. V., et al. (2005). A comparison of label-based review and ALE meta-analysis in the stroop task. Human Brain Mapping, 25, 6–21]. We conducted separate meta-analyses for four contrasts of interest: episodic encoding success as measured in the subsequent-memory paradigm (subsequent Hit vs. Miss), episodic retrieval success (Hit vs. Correct Rejection), objective recollection (e.g., Source Hit vs. Item Hit), and subjective recollection (e.g., Remember vs. Know). Concordance maps revealed significant cross-study overlap for each contrast. In each case, the left hemisphere showed greater concordance than the right hemisphere. Both encoding and retrieval success were associated with activation in medial-temporal, prefrontal, and parietal regions. Left ventrolateral prefrontal cortex (PFC) and medial-temporal regions were more strongly involved in encoding, whereas left superior parietal and dorsolateral and anterior PFC regions were more strongly involved in retrieval. Objective recollection was associated with activation in multiple PFC regions, as well as multiple posterior parietal and medial-temporal areas, but not hippocampus. Subjective recollection, in contrast, showed left hippocampal involvement. In summary, these results identify broadly consistent activation patterns associated with episodic encoding and retrieval, and subjective and objective recollection, but also subtle differences among these processes.