Visual recognition memory of scenes is driven by categorical, not sensory, visual representations.

When we perceive a scene, our brain processes various types of visual information simultaneously, ranging from sensory features, such as line orientations and colors, to categorical features, such as objects and their arrangements. Whereas the role of sensory and categorical visual representations in predicting subsequent memory has been studied using isolated objects, their impact on memory for complex scenes remains largely unknown. To address this gap, we conducted an fMRI study in which female and male participants encoded pictures of familiar scenes (e.g., an airport picture) and later recalled them, while rating the vividness of their visual recall. Outside the scanner, participants had to distinguish each seen scene (e.g., an airport picture) from three similar lures (e.g., 3 airport pictures). We modeled the sensory and categorical visual features of multiple scenes using both early and late layers of a deep convolutional neural network. Then, we applied representational similarity analysis to determine which brain regions represented stimuli in accordance with the sensory and categorical models. We found that categorical, but not sensory, representations predicted subsequent memory. In line with the previous result, only for the categorical model, the average recognition performance of each scene exhibited a positive correlation with the average visual dissimilarity between the item in question and its respective lures. These results strongly suggest that even in memory tests that ostensibly rely solely on visual cues (such as force-choice visual recognition with similar distractors), memory decisions for scenes may be primarily influenced by categorical rather than sensory representations.Significance Statement Our memory for real-world scenes often comprises a tableau of complex visual features, but recent findings challenge the view that our memories of such stimuli rely on purely visual information. Instead, it appears that our memory for scenes is heavily influenced by higher-level categorical information. Analyzing cortical representations in regions responsive to both categorical and sensory features, we discovered that only the former can reliably predict memory outcomes. Moreover, the distinctiveness of scenes in terms of their categoric features among similar examples is positively associated with our ability to accurately recognize previously encountered scenes. In essence, this study sheds light on how our brains rely on categorical information to recognize natural scenes.

The structure of prior knowledge enhances memory in experts by reducing interference.

The influence of prior knowledge on memory is ubiquitous, making the specific mechanisms of this relationship difficult to disentangle. Here, we show that expert knowledge produces a fundamental shift in the way that interitem similarity (i.e., the perceived resemblance between items in a set) biases episodic recognition. Within a group of expert birdwatchers and matched controls, we characterized the psychological similarity space for a set of well-known local species and a set of less familiar, nonlocal species. In experts, interitem similarity was influenced most strongly by taxonomic features, whereas in controls, similarity judgments reflected bird color. In controls, perceived episodic oldness during a recognition memory task increased along with measures of global similarity between items, consistent with classic models of episodic recognition. Surprisingly, for experts, high global similarity did not drive oldness signals. Instead, for local birds memory tracked the availability of species-level name knowledge, whereas for nonlocal birds, it was mediated by the organization of generalized conceptual space. These findings demonstrate that episodic memory in experts can benefit from detailed subcategory knowledge, or, lacking that, from the overall relational structure of concepts. Expertise reshapes psychological similarity space, helping to resolve mnemonic separation challenges arising from high interitem overlap. Thus, even in the absence of knowledge about item-specific details or labels, the presence of generalized knowledge appears to support episodic recognition in domains of expertise by altering the typical relationship between psychological similarity and memory.

The Role of the Ventromedial Prefrontal Cortex and Basal Forebrain in Relational Memory and Inference.

The ventromedial prefrontal cortex (vmPFC) is involved in diverse cognitive operations, from inhibitory control to processing of semantic schemas. When accompanied by damage to the basal forebrain, vmPFC lesions can also impair relational memory, the ability to form and recall relations among items. Impairments in establishing direct relations among items (e.g., A is related to B, B is related to C) can also hinder the transitive processing of indirect relationships (e.g., inferring that A and C are related through direct relations that each contain B). Past work has found that transitive inference improves when the direct relations are organized within an existing knowledge structure, or schema. This type of semantic support is most effective for individuals whose relational memory deficits are mild (e.g., healthy age-related decline) rather than pronounced (e.g., hippocampal amnesia, amnestic mild cognitive impairment). Given that vmPFC damage can produce both relational memory and schema processing deficits, such damage may pose a particular challenge in establishing the type of relational structure required for transitive inference, even when supported by preexisting knowledge. To examine this idea, we tested individuals with lesions to the mPFC on multiple conditions that varied in pre-experimental semantic support and explored the extent to which they could identify both previously studied (direct) and novel transitive (indirect) relations. Most of the mPFC cases showed marked transitive inference deficits and even showed impaired knowledge of preexisting, direct, semantic relations, consistent with disruptions to schema-related processes. However, one case with more dorsal mPFC damage showed preserved ability to identify direct relations and make novel inferences, particularly when pre-experimental knowledge could be used to support performance. These results suggest that damage to the mPFC and basal forebrain can impede establishment of ad hoc relational schemas upon which transitive inference is based, but that appealing to prior knowledge may still be useful for those neurological cases that have some degree of preserved relational memory.

Age-related dedifferentiation and hyperdifferentiation of perceptual and mnemonic representations.

Preliminary evidence indicates that occipito-temporal activation patterns for different visual stimuli are less distinct in older (OAs) than younger (YAs) adults, suggesting a dedifferentiation of visual representations with aging. Yet, it is unclear if this deficit (1) affects only sensory or also categorical aspects of representations during visual perception (perceptual representations), and (2) affects only perceptual or also mnemonic representations. To investigate these issues, we fMRI-scanned YAs and OAs viewing and then remembering visual scenes. First, using representational similarity analyses, we distinguished sensory vs. categorical features of perceptual representations. We found that, compared to YAs, sensory features in early visual cortex were less differentiated in OAs (i.e., age-related dedifferentiation), replicating previous research, whereas categorical features in anterior temporal lobe (ATL) were more differentiated in OAs. This is, to our knowledge, the first report of an age-related hyperdifferentiation. Second, we assessed the quality of mnemonic representations by measuring encoding-retrieval similarity (ERS) in activation patterns. We found that aging impaired mnemonic representations in early visual cortex and hippocampus but enhanced mnemonic representations in ATL. Thus, both perceptual and mnemonic representations in ATL were enhanced by aging. In sum, our findings suggest that aging impairs visual and mnemonic representations in posterior brain regions but enhances them in anterior regions.

Age-Related Compensatory Reconfiguration of PFC Connections during Episodic Memory Retrieval.

During demanding cognitive tasks, older adults (OAs) frequently show greater prefrontal cortex (PFC) activity than younger adults (YAs). This age-related increase in PFC activity is often associated with enhanced cognitive performance, suggesting functional compensation. However, the brain is a complex network of interconnected regions, and it is unclear how network connectivity of PFC regions differs for OAs versus YAs. To investigate this, we examined the age-related difference on the functional brain networks mediating episodic memory retrieval. YAs and OAs participants encoded and recalled visual scenes, and age-related differences in network topology during memory retrieval were investigated as a function of memory performance. We measured both changes in functional integration and reconfiguration in connectivity patterns. The study yielded three main findings. First, PFC regions were more functionally integrated with the rest of the brain network in OAs. Critically, this age-related increase in PFC integration was associated with better retrieval performance. Second, PFC regions showed stronger performance-related reconfiguration of connectivity patterns in OAs. Finally, the PFC reconfiguration increases in OAs tracked reconfiguration reductions in the medial temporal lobe (MTL)-a core episodic memory region, suggesting that PFC connectivity in OAs may be compensating for MTL deficits.

Visual and Semantic Representations Predict Subsequent Memory in Perceptual and Conceptual Memory Tests.

It is generally assumed that the encoding of a single event generates multiple memory representations, which contribute differently to subsequent episodic memory. We used functional magnetic resonance imaging (fMRI) and representational similarity analysis to examine how visual and semantic representations predicted subsequent memory for single item encoding (e.g., seeing an orange). Three levels of visual representations corresponding to early, middle, and late visual processing stages were based on a deep neural network. Three levels of semantic representations were based on normative observed ("is round"), taxonomic ("is a fruit"), and encyclopedic features ("is sweet"). We identified brain regions where each representation type predicted later perceptual memory, conceptual memory, or both (general memory). Participants encoded objects during fMRI, and then completed both a word-based conceptual and picture-based perceptual memory test. Visual representations predicted subsequent perceptual memory in visual cortices, but also facilitated conceptual and general memory in more anterior regions. Semantic representations, in turn, predicted perceptual memory in visual cortex, conceptual memory in the perirhinal and inferior prefrontal cortex, and general memory in the angular gyrus. These results suggest that the contribution of visual and semantic representations to subsequent memory effects depends on a complex interaction between representation, test type, and storage location.

Application of long-interval paired-pulse transcranial magnetic stimulation to motion-sensitive visual cortex does not lead to changes in motion discrimination.

The perception of visual motion is dependent on a set of occipitotemporal regions that are readily accessible to neuromodulation. The current study tested if paired-pulse Transcranial Magnetic Stimulation (ppTMS) could modulate motion perception by stimulating the occipital cortex as participants viewed near-threshold motion dot stimuli. In this sham-controlled study, fifteen subjects completed two sessions. On the first visit, resting motor threshold (RMT) was assessed, and participants performed an adaptive direction discrimination task to determine individual motion sensitivity. During the second visit, subjects performed the task with three difficulty levels as TMS pulses were delivered 150 and 50 ms prior to motion stimulus onset at 120% RMT, under the logic that the cumulative inhibitory effect of these pulses would alter motion sensitivity. ppTMS was delivered at one of two locations: 3 cm dorsal and 5 cm lateral to inion (scalp-based coordinate), or at the site of peak activation for "motion" according to the NeuroSynth fMRI database (meta-analytic coordinate). Sham stimulation was delivered on one-third of trials by tilting the coil 90°. Analyses showed no significant active-versus-sham effects of ppTMS when stimulation was delivered to the meta-analytic (p = 0.15) or scalp-based coordinates (p = 0.17), which were separated by 29 mm on average. Active-versus-sham stimulation differences did not interact with either stimulation location (p = 0.12) or difficulty (p = 0.33). These findings fail to support the hypothesis that long-interval ppTMS recruits inhibitory processes in motion-sensitive cortex but must be considered within the limited parameters used in this design.

Cortical Overlap and Cortical-Hippocampal Interactions Predict Subsequent True and False Memory.

The declarative memory system allows us to accurately recognize a countless number of items and events, particularly those strengthened by repeated exposure. However, increased familiarity due to repetition can also lead to false recognition of related but new items, particularly when mechanisms supporting fine-grain mnemonic discrimination fail. The hippocampus is thought to be particularly important in separating overlapping cortical inputs during encoding so that similar experiences can be differentiated. In the current study of male and female human subjects, we examine how neural pattern similarity between repeated exemplars of a given concept (e.g., apple) influences true and false memory for target or lure images. Consistent with past work, we found that subsequent true recognition was related to pattern similarity between concept exemplars and the entire encoding set (global encoding similarity), particularly in ventral visual stream. In addition, memory for an individual target exemplar (a specific apple) could be predicted solely by the degree of pattern overlap between the other exemplars (different apple pictures) of that concept (concept-specific encoding similarity). Critically, subsequent false memory for lures was mitigated when high concept-specific similarity in cortical areas was accompanied by differentiated hippocampal representations of the corresponding exemplars. Furthermore, both true and false memory entailed the reinstatement of concept-related information at varying levels of specificity. These results link both true and false memory to a measure of concept strength expressed in the overlap of cortical representations, and importantly, illustrate how the hippocampus serves to separate concurrent cortical overlap in the service of detailed memory.SIGNIFICANCE STATEMENT In some instances, the same processes that help promote memory for a general idea or concept can also hinder more detailed memory judgments, which may involve differentiating between closely related items. The current study shows that increased overlap in cortical representations for conceptually-related pictures is associated with increased recognition of repeated concept pictures. Whether similar lure items were falsely remembered as old further depended on the hippocampus, where the presence of more distinct representations protected against later false memory. This work suggests that the differentiability of brain patterns during perception is related to the differentiability of items in memory, but that fine-grain discrimination depends on the interaction between cortex and hippocampus.

Neural basis of goal-driven changes in knowledge activation.

Depending on a person's goals, different aspects of stored knowledge are accessed. Decades of behavioral work document the flexible use of knowledge, but little neuroimaging work speaks to these questions. We used representational similarity analysis to investigate whether the relationship between brain activity and semantic structure of statements varied in two tasks hypothesized to differ in the degree to which knowledge is accessed: judging truth (semantic task) and judging oldness (episodic task). During truth judgments, but not old/new recognition judgments, a left-lateralized network previously associated with semantic memory exhibited correlations with semantic structure. At a neural level, people activate knowledge representations in different ways when focused on different goals. The present results demonstrate the potential of multivariate approaches in characterizing knowledge storage and retrieval, as well as the ways that it shapes our understanding and long-term memory.

Excitatory TMS modulates memory representations.

Brain stimulation technologies have seen increasing application in basic science investigations, specifically toward the goal of improving memory function. However, proposals concerning the neural mechanisms underlying cognitive enhancement often rely on simplified notions of excitation. As a result, most applications examining the effects of transcranial magnetic stimulation (TMS) on functional neuroimaging measures have been limited to univariate analyses of brain activity. We present here analyses using representational similarity analysis (RSA) and encoding-retrieval similarity (ERS) analysis to quantify the effect of TMS on memory representations. To test whether an increase in local excitability in PFC can have measurable influences on upstream representations in earlier temporal memory regions, we compared 1 and 5Hz stimulation to the left dorsolateral PFC (DLPFC). We found that 5Hz rTMS, relative to 1Hz, had multiple effects on neural representations: 1) greater representational similarity during both encoding and retrieval in ventral stream regions, 2) greater ERS in the hippocampus, and, critically, 3) increasing ERS in MTL was correlated with increasing univariate activity in DLPFC, and greater functional connectivity for hits than misses between these regions. These results provide the first evidence of rTMS modulating semantic representations and strengthen the idea that rTMS may affect the reinstatement of previously experienced events in upstream regions.

Contributions of the ventral parietal cortex to declarative memory.

Our understanding of the role that ventral parietal cortex (VPC) plays in declarative memory processes has changed dramatically over the last two decades. The goal of this chapter is to provide a concise overview data concerning VPC involvement in episodic memory (EM), and to connect this data to several key theories of VPC function. We review evidence from five methodological domains in cognitive neuroscience: neuropsychological lesion evidence, univariate activation studies, multivoxel pattern analyses, functional connectivity studies, and brain stimulation experiments. We discuss how the body of empirical work bears on putative mnemonic functions of VPC related to attention and stimulus representation, and detail the strengths and weaknesses of related theories. Lastly, we identify several broad conceptual questions raised by recent investigations, and outline directions for future research.

Knowledge supports memory retrieval through familiarity, not recollection.

Semantic memory, or general knowledge of the world, guides learning and supports the formation and retrieval of new episodic memories. Behavioral evidence suggests that this knowledge effect is supported by recollection-a more controlled form of memory retrieval generally accompanied by contextual details-to a greater degree than familiarity-a more automatic form of memory retrieval generally absent of contextual details. In the current study, we used functional magnetic resonance imaging (fMRI) to investigate the role that regions associated with recollection and familiarity play in retrieving recent instances of known (e.g., The Summer Olympic Games are held four years apart) and unknown (e.g., A flaky deposit found in port bottles is beeswing) statements. Our results revealed a surprising pattern: Episodic retrieval of known statements recruited regions associated with familiarity, but not recollection. Instead, retrieval of unknown statements recruited regions associated with recollection. These data, in combination with quicker reaction times for the retrieval of known than unknown statements, suggest that known statements can be successfully retrieved on the basis of familiarity, whereas unknown statements were retrieved on the basis of recollection. Our results provide insight into how knowledge influences episodic retrieval and demonstrate the role of neuroimaging in providing insights into cognitive processes in the absence of explicit behavioral responses.

Neural mechanisms underlying subsequent memory for personal beliefs:An fMRI study.

Many fMRI studies have examined the neural mechanisms supporting emotional memory for stimuli that generate emotion rather automatically (e.g., a picture of a dangerous animal or of appetizing food). However, far fewer studies have examined how memory is influenced by emotion related to social and political issues (e.g., a proposal for large changes in taxation policy), which clearly vary across individuals. In order to investigate the neural substrates of affective and mnemonic processes associated with personal opinions, we employed an fMRI task wherein participants rated the intensity of agreement/disagreement to sociopolitical belief statements paired with neural face pictures. Following the rating phase, participants performed an associative recognition test in which they distinguished identical versus recombined face-statement pairs. The study yielded three main findings: behaviorally, the intensity of agreement ratings was linked to greater subjective emotional arousal as well as enhanced high-confidence subsequent memory. Neurally, statements that elicited strong (vs. weak) agreement or disagreement were associated with greater activation of the amygdala. Finally, a subsequent memory analysis showed that the behavioral memory advantage for statements generating stronger ratings was dependent on the medial prefrontal cortex (mPFC). Together, these results both underscore consistencies in neural systems supporting emotional arousal and suggest a modulation of arousal-related encoding mechanisms when emotion is contingent on referencing personal beliefs.

Search and recovery of autobiographical and laboratory memories: Shared and distinct neural components.

Functional neuroimaging evidence suggests that there are differences in the neural correlates of episodic memory for laboratory stimuli (laboratory memory) and for events from one's own life (autobiographical memory). However, this evidence is scarce and often confounded with differences in memory testing procedures. Here, we directly compared the neural mechanisms underlying the search and recovery of autobiographical and laboratory memories while minimizing testing differences. Before scanning, participants completed a laboratory memory encoding task in which they studied four-word "chains" spread across three word pairs. During scanning, participants completed a laboratory memory retrieval task, in which they recalled the word chains, and an autobiographical memory retrieval task, in which they recalled specific personal events associated with word cues. Importantly, response times were similar in the two tasks, allowing for a direct comparison of the activation time courses. We found that during memory search (searching for the memory target), similar brain regions were activated during both the autobiographical and laboratory tasks, whereas during memory recovery (accessing the memory traces; i.e., ecphory), clear differences emerged: regions of the default mode network (DMN) were activated greater during autobiographical than laboratory memory, whereas the bilateral superior parietal lobules were activated greater during laboratory than autobiographical memory. Also, multivariate functional connectivity analyses revealed that regardless of memory stage, the DMN and ventral attention network exhibited a more integrated topology in the functional network underlying autobiographical (vs. laboratory) memory retrieval, whereas the fronto-parietal task control network exhibited a more integrated topology in the functional network underlying laboratory (vs. autobiographical) memory retrieval. These findings further characterize the shared and distinct neural components underlying autobiographical and laboratory memories, and suggest that differences in autobiographical vs. laboratory memory brain activation previously reported in the literature reflect memory recovery rather than search differences.

Hippocampal Contributions to the Large-Scale Episodic Memory Network Predict Vivid Visual Memories.

A common approach in memory research is to isolate the function(s) of individual brain regions, such as the hippocampus, without addressing how those regions interact with the larger network. To investigate the properties of the hippocampus embedded within large-scale networks, we used functional magnetic resonance imaging and graph theory to characterize complex hippocampal interactions during the active retrieval of vivid versus dim visual memories. The study yielded 4 main findings. First, the right hippocampus displayed greater communication efficiency with the network (shorter path length) and became a more convergent structure for information integration (higher centrality measures) for vivid than dim memories. Second, vivid minus dim differences in our graph theory measures of interest were greater in magnitude for the right hippocampus than for any other region in the 90-region network. Moreover, the right hippocampus significantly reorganized its set of direct connections from dim to vivid memory retrieval. Finally, beyond the hippocampus, communication throughout the whole-brain network was more efficient (shorter global path length) for vivid than dim memories. In sum, our findings illustrate how multivariate network analyses can be used to investigate the roles of specific regions within the large-scale network, while also accounting for global network changes.

On Known Unknowns: Fluency and the Neural Mechanisms of Illusory Truth.

The "illusory truth" effect refers to the phenomenon whereby repetition of a statement increases its likelihood of being judged true. This phenomenon has important implications for how we come to believe oft-repeated information that may be misleading or unknown. Behavioral evidence indicates that fluency, the subjective ease experienced while processing information, underlies this effect. This suggests that illusory truth should be mediated by brain regions previously linked to fluency, such as the perirhinal cortex (PRC). To investigate this possibility, we scanned participants with fMRI while they rated the truth of unknown statements, half of which were presented earlier (i.e., repeated). The only brain region that showed an interaction between repetition and ratings of perceived truth was PRC, where activity increased with truth ratings for repeated, but not for new, statements. This finding supports the hypothesis that illusory truth is mediated by a fluency mechanism and further strengthens the link between PRC and fluency.

The influence of self-awareness on emotional memory formation: an fMRI study.

Evidence from functional neuroimaging studies of emotional perception shows that when attention is focused on external features of emotional stimuli (external perceptual orienting--EPO), the amygdala is primarily engaged, but when attention is turned inwards towards one's own emotional state (interoceptive self-orienting--ISO), regions of the salience network, such as the anterior insula (AI) and the dorsal anterior cingulate cortex (dACC), also play a major role. Yet, it is unknown if ISO boosts the contributions of AI and dACC not only to emotional 'perception' but also to emotional 'memory'. To investigate this issue, participants were scanned with functional magnetic resonance imaging (fMRI) while viewing emotional and neutral pictures under ISO or EPO, and memory was tested several days later. The study yielded three main findings: (i) emotion boosted perception-related activity in the amygdala during both ISO and EPO and in the right AI exclusively during ISO; (ii) emotion augmented activity predicting subsequent memory in AI and dACC during ISO but not during EPO and (iii) high confidence memory was associated with increased amygdala-dACC connectivity, selectively for ISO encoding. These findings show, for the first time, that ISO promotes emotional memory formation via regions associated with interoceptive awareness of emotional experience, such as AI and dACC.

Reinstatement of individual past events revealed by the similarity of distributed activation patterns during encoding and retrieval.

Neurobiological memory models assume memory traces are stored in neocortex, with pointers in the hippocampus, and are then reactivated during retrieval, yielding the experience of remembering. Whereas most prior neuroimaging studies on reactivation have focused on the reactivation of sets or categories of items, the current study sought to identify cortical patterns pertaining to memory for individual scenes. During encoding, participants viewed pictures of scenes paired with matching labels (e.g., "barn," "tunnel"), and, during retrieval, they recalled the scenes in response to the labels and rated the quality of their visual memories. Using representational similarity analyses, we interrogated the similarity between activation patterns during encoding and retrieval both at the item level (individual scenes) and the set level (all scenes). The study yielded four main findings. First, in occipitotemporal cortex, memory success increased with encoding-retrieval similarity (ERS) at the item level but not at the set level, indicating the reactivation of individual scenes. Second, in ventrolateral pFC, memory increased with ERS for both item and set levels, indicating the recapitulation of memory processes that benefit encoding and retrieval of all scenes. Third, in retrosplenial/posterior cingulate cortex, ERS was sensitive to individual scene information irrespective of memory success, suggesting automatic activation of scene contexts. Finally, consistent with neurobiological models, hippocampal activity during encoding predicted the subsequent reactivation of individual items. These findings show the promise of studying memory with greater specificity by isolating individual mnemonic representations and determining their relationship to factors like the detail with which past events are remembered.

The neural basis of involuntary episodic memories.

Voluntary episodic memories require an intentional memory search, whereas involuntary episodic memories come to mind spontaneously without conscious effort. Cognitive neuroscience has largely focused on voluntary memory, leaving the neural mechanisms of involuntary memory largely unknown. We hypothesized that, because the main difference between voluntary and involuntary memory is the controlled retrieval processes required by the former, there would be greater frontal activity for voluntary than involuntary memories. Conversely, we predicted that other components of the episodic retrieval network would be similarly engaged in the two types of memory. During encoding, all participants heard sounds, half paired with pictures of complex scenes and half presented alone. During retrieval, paired and unpaired sounds were presented, panned to the left or to the right. Participants in the involuntary group were instructed to indicate the spatial location of the sound, whereas participants in the voluntary group were asked to additionally recall the pictures that had been paired with the sounds. All participants reported the incidence of their memories in a postscan session. Consistent with our predictions, voluntary memories elicited greater activity in dorsal frontal regions than involuntary memories, whereas other components of the retrieval network, including medial-temporal, ventral occipitotemporal, and ventral parietal regions were similarly engaged by both types of memories. These results clarify the distinct role of dorsal frontal and ventral occipitotemporal regions in predicting strategic retrieval and recalled information, respectively, and suggest that, although there are neural differences in retrieval, involuntary memories share neural components with established voluntary memory systems.

Neural correlates of retrieval-based memory enhancement: an fMRI study of the testing effect.

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).