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While many theories assume that sleep is critical in stabilizing and strengthening memories, our recent behavioral study (Liu & Ranganath, 2021, Psychonomic Bulletin & Review, 28[6], 2035-2044) suggests that sleep does not simply stabilize memories. Instead, it plays a more complex role, integrating information across two temporally distinct learning episodes. In the current study, we simulated the results of Liu and Ranganath (2021) using our biologically plausible computational model, TEACH, developed based on the complementary learning systems (CLS) framework. Our model suggests that when memories are activated during sleep, the reduced influence of temporal context establishes connections across temporally separated events through mutual training between the hippocampus and neocortex. In addition to providing a compelling mechanistic explanation for the selective effect of sleep, this model offers new examples of the diverse ways in which the cortex and hippocampus can interact during learning.

Recollection of real-world events is often accompanied by a sense of being in the place where the event transpired. Convergent evidence suggests the hippocampus plays a key role in supporting episodic memory by associating information with the time and place it was originally encountered. This representation is reinstated during memory retrieval. However, little is known about the roles of different subfields of the human hippocampus in this process. Research in humans and non-human animal models has suggested that spatial environmental boundaries have a powerful influence on spatial and episodic memory, as well as hippocampal representations of contexts and events. Here, we used high-resolution fMRI to investigate how boundaries influence hippocampal activity patterns during the recollection of objects encountered in different spatial contexts. During the encoding phase, participants viewed objects once in a naturalistic virtual reality task in which they passively explored two rooms in one of two houses. Following the encoding phase, participants were scanned while they recollected items in the absence of any spatial contextual information. Our behavioral results demonstrated that spatial context memory was enhanced for objects encountered near a boundary. Activity patterns in CA1 carried information about the spatial context associated with each of these boundary items. Exploratory analyses revealed that recollection performance was correlated with the fidelity of retrieved spatial context representations in anterior parahippocampal cortex and subiculum. Our results highlight the privileged role of boundaries in CA1 and suggest more generally a close relationship between memory for spatial contexts and representations in the hippocampus and parahippocampal region.

Converging evidence from studies of human and nonhuman animals suggests that the hippocampus contributes to sequence learning by using temporal context to bind sequentially occurring items. The fornix is a white matter pathway containing the major input and output pathways of the hippocampus, including projections from medial septum and to diencephalon, striatum, lateral septum and prefrontal cortex. If the fornix meaningfully contributes to hippocampal function, then individual differences in fornix microstructure might predict sequence memory. Here, we tested this prediction by performing tractography in 51 healthy adults who had undertaken a sequence memory task. Microstructure properties of the fornix were compared with those of tracts connecting medial temporal lobe regions but not predominantly the hippocampus: the Parahippocampal Cingulum bundle (PHC) (conveying retrosplenial projections to parahippocampal cortex) and the Inferior Longitudinal Fasciculus (ILF) (conveying occipital projections to perirhinal cortex). Using principal components analysis, we combined Free-Water Elimination Diffusion Tensor Imaging and Neurite Orientation Dispersion and Density Imaging measures obtained from multi-shell diffusion MRI into two informative indices: the first (PC1) capturing axonal packing/myelin and the second (PC2) capturing microstructural complexity. We found a significant correlation between fornix PC2 and implicit reaction-time indices of sequence memory, indicating that greater fornix microstructural complexity is associated with better sequence memory. No such relationship was found with measures from the PHC and ILF. This study highlights the importance of the fornix in aiding memory for objects within a temporal context, potentially reflecting a role in mediating inter-regional communication within an extended hippocampal system.

Memory is well known to decline over the course of healthy aging. However, memory is not a monolith and draws from different kinds of representations. Historically, much of our understanding of age-related memory decline stems from recognition of isolated studied items. In contrast, real-life events are often remembered as narratives, and this kind of information is generally missed in typical recognition memory studies. Here, we designed a task to tax mnemonic discrimination of event details, directly contrasting perceptual and narrative memory. Older and younger adults watched an episode of a television show and later completed an old/new recognition test featuring targets, novel foils, and similar lures in narrative and perceptual domains. While we observed no age-related differences on basic recognition of repeated targets and novel foils, older adults showed a deficit in correctly rejecting perceptual, but not narrative, lures. These findings provide insight into the vulnerability of different memory domains in aging and may be useful in characterizing individuals at risk for pathological cognitive decline.

We often remember past experiences in the form of a story. We can even form narratives from events which occur at disparate times. Yet, the neuroscience of memory has historically tried to understand how we remember specific periods of time, or “events.”The hippocampus is necessary for remembering specific events. One possibility is that the hippocampus organizes our memories in time.However, we previously found that narratives can bridge the gap between events in our memory, making events easier to remember.Hypothesis: the hippocampus might link two events together in memory when they can form a single, coherent narrative.

Recent work in cognitive and systems neuroscience has suggested that the hippocampus might support planning, imagination, and navigation by forming cognitive maps that capture the abstract structure of physical spaces, tasks, and situations. Navigation involves disambiguating similar contexts, and the planning and execution of a sequence of decisions to reach a goal. Here, we examine hippocampal activity patterns in humans during a goal-directed navigation task to investigate how contextual and goal information are incorporated in the construction and execution of navigational plans. During planning, hippocampal pattern similarity is enhanced across routes that share a context and a goal. During navigation, we observe prospective activation in the hippocampus that reflects the retrieval of pattern information related to a key-decision point. These results suggest that, rather than simply representing overlapping associations or state transitions, hippocampal activity patterns are shaped by context and goals.

Although every life event is unique, there are considerable commonalities across events. However, little is known about whether or how the brain flexibly represents information about different event components at encoding and during remembering. Here, we show that different cortico-hippocampal networks systematically represent specific components of events depicted in videos, both during online experience and during episodic memory retrieval. Regions of an Anterior Temporal Network represented information about people, generalizing across contexts, whereas regions of a Posterior Medial Network represented context information, generalizing across people. Medial prefrontal cortex generalized across videos depicting the same event schema, whereas the hippocampus maintained event-specific representations. Similar effects were seen in real-time and recall, suggesting reuse of event components across overlapping episodic memories. These representational profiles together provide a computationally optimal strategy to scaffold memory for different high-level event components, allowing efficient reuse for event comprehension, recollection, and imagination.

BackgroundTheories suggest that people with schizophrenia (SZ) have problems generating predictions based on past experiences. The dorsolateral prefrontal cortex (DLPFC) and hippocampus participate in memory-based prediction. We used functional magnetic resonance imaging to investigate DLPFC and hippocampal function in healthy control (HC) subjects and people with SZ during memory-based prediction.MethodsPrior to scanning, HC subjects (n = 54) and people with SZ (n = 31) learned 5-object sequences presented in fixed or random orders on each repetition. During scanning, participants made semantic decisions (e.g., "Can this object fit in a shoebox?") on a continuous stream of objects from fixed and random sequences. Sequence prediction was demonstrated by faster semantic decisions for objects in fixed versus random sequences because memory could be used to anticipate and more efficiently process semantic information about upcoming objects in fixed sequences. Representational similarity analyses were used to determine how each sequence type was represented in the posterior hippocampus and DLPFC.ResultsSequence predictions were reduced in individuals with SZ relative to HC subjects. Representational similarity analyses revealed stronger memory-based predictions in the DLPFC of HC subjects than people with SZ, and DLPFC representations correlated with more successful predictions in HC subjects only. For the posterior hippocampus, voxel pattern similarity was increased for fixed versus random sequences in HC subjects only, but no significant between-group differences or correlations with prediction success were observed.ConclusionsIndividuals with SZ are capable of learning temporal sequences; however, they are impaired using memory to predict upcoming events as efficiently as HC subjects. This deficit appears related to disrupted neural representation of sequence information in the DLPFC.

The hippocampus plays a critical role in supporting episodic memory, in large part by binding together experiences and items with surrounding contextual information. At present, however, little is known about the roles of different hippocampal subfields in supporting this item-context binding. To address this question, we constructed a task in which items were affiliated with differing types of context-cognitive associations that vary at the local, item level and membership in temporally organized lists that linked items together at a global level. Participants made item recognition judgments while undergoing high-resolution fMRI. We performed voxel pattern similarity analyses to answer the question of how human hippocampal subfields represent retrieved information about cognitive states and the time at which a past event took place. As participants recollected previously presented items, activity patterns in the CA23DG subregion carried information about prior cognitive states associated with these items. We found no evidence to suggest reinstatement of information about temporal context at the level of list membership, but exploratory analyses revealed representations of temporal context at a coarse level in conjunction with representations of cognitive contexts. Results are consistent with characterizations of CA23DG as a critical site for binding together items and contexts in the service of memory retrieval.

When making a turn at a familiar intersection, we know what items and landmarks will come into view. These perceptual expectations, or predictions, come from our knowledge of the context; however, it is unclear how memory and perceptual systems interact to support the prediction and reactivation of sensory details in cortex. To address this, human participants learned the spatial layout of animals positioned in a cross maze. During fMRI, participants of both sexes navigated between animals to reach a target, and in the process saw a predictable sequence of five animal images. Critically, to isolate activity patterns related to item predictions, rather than bottom-up inputs, one-fourth of trials ended early, with a blank screen presented instead. Using multivariate pattern similarity analysis, we reveal that activity patterns in early visual cortex, posterior medial regions, and the posterior hippocampus showed greater similarity when seeing the same item compared with different items. Further, item effects in posterior hippocampus were specific to the sequence context. Critically, activity patterns associated with seeing an item in visual cortex and posterior medial cortex, were also related to activity patterns when an item was expected, but omitted, suggesting sequence predictions were reinstated in these regions. Finally, multivariate connectivity showed that patterns in the posterior hippocampus at one position in the sequence were related to patterns in early visual cortex and posterior medial cortex at a later position. Together, our results support the idea that hippocampal representations facilitate sensory processing by modulating visual cortical activity in anticipation of expected items.SIGNIFICANCE STATEMENT Our visual world is a series of connected events, where we can predict what we might see next based on our recent past. Understanding the neural circuitry and mechanisms of the perceptual and memory systems that support these expectations is fundamental to revealing how we perceive and act in our world. Using brain imaging, we studied what happens when we expect to see specific visual items, and how such expectations relate to top-down memory signals. We find both visual and memory systems reflect item predictions, and moreover, we show that hippocampal activity supports predictions of future expected items. This demonstrates that the hippocampus acts to predict upcoming items, and reinstates such predictions in cortex.

Many studies suggest that information about past experience, or episodic memory, is divided into discrete units called "events." Yet we can often remember experiences that span multiple events. Events that occur in close succession might simply be linked because of their proximity to one another, but we can also build links between events that occur farther apart in time. Intuitively, some kind of organizing principle should enable temporally distant events to become bridged in memory. We tested the hypothesis that episodic memory exhibits a narrative-level organization, enabling temporally distant events to be better remembered if they form a coherent narrative. Furthermore, we tested whether post-encoding memory consolidation is necessary to integrate temporally distant events. In three experiments, participants learned and subsequently recalled events from fictional stories, in which pairs of temporally distant events involving side characters ("sideplots") either formed one coherent narrative or two unrelated narratives. Across participants, we varied whether recall was assessed immediately after learning, or after a delay: 24 hours, 12 hours between morning and evening ("wake"), or 12 hours between evening and morning ("sleep"). Participants recalled more information about coherent than unrelated narrative events, in most delay conditions, including immediate recall and wake conditions, suggesting that post-encoding consolidation was not necessary to integrate temporally distant events into a larger narrative. Furthermore, post hoc modeling across experiments suggested that narrative coherence facilitated recall over and above any effects of sentence-level semantic similarity. This reliable memory benefit for coherent narrative events supports theoretical accounts which propose that narratives provide a high-level architecture for episodic memory.

A hallmark of human intelligence is the ability to adapt to new situations, by applying learned rules to new content (systematicity) and thereby enabling an open-ended number of inferences and actions (generativity). Here, we propose that the human brain accomplishes these feats through pathways in the parietal cortex that encode the abstract structure of space, events, and tasks, and pathways in the temporal cortex that encode information about specific people, places, and things (content). Recent neural network models show how the separation of structure and content might emerge through a combination of architectural biases and learning, and these networks show dramatic improvements in the ability to capture systematic, generative behavior. We close by considering how the hippocampal formation may form integrative memories that enable rapid learning of new structure and content representations.

The hippocampus plays a critical role in the rapid learning of new episodic memories. Many computational models propose that the hippocampus is an autoassociator that relies on Hebbian learning (i.e., "cells that fire together, wire together"). However, Hebbian learning is computationally suboptimal as it does not learn in a way that is driven toward, and limited by, the objective of achieving effective retrieval. Thus, Hebbian learning results in more interference and a lower overall capacity. Our previous computational models have utilized a powerful, biologically plausible form of error-driven learning in hippocampal CA1 and entorhinal cortex (EC) (functioning as a sparse autoencoder) by contrasting local activity states at different phases in the theta cycle. Based on specific neural data and a recent abstract computational model, we propose a new model called Theremin (Total Hippocampal ERror MINimization) that extends error-driven learning to area CA3-the mnemonic heart of the hippocampal system. In the model, CA3 responds to the EC monosynaptic input prior to the EC disynaptic input through dentate gyrus (DG), giving rise to a temporal difference between these two activation states, which drives error-driven learning in the EC→CA3 and CA3↔CA3 projections. In effect, DG serves as a teacher to CA3, correcting its patterns into more pattern-separated ones, thereby reducing interference. Results showed that Theremin, compared with our original Hebbian-based model, has significantly increased capacity and learning speed. The model makes several novel predictions that can be tested in future studies.

In memory, our continuous experiences are broken up into discrete events. Boundaries between events are known to influence the temporal organization of memory. However, how and through which mechanism event boundaries shape temporal order memory (TOM) remains unknown. Across four experiments, we show that event boundaries exert a dual role: improving TOM for items within an event and impairing TOM for items across events. Decreasing event length in a list enhances TOM, but only for items at earlier local event positions, an effect we term the local primacy effect. A computational model, in which items are associated to a temporal context signal that drifts over time but resets at boundaries captures all behavioural results. Our findings provide a unified algorithmic mechanism for understanding how and why event boundaries affect TOM, reconciling a long-standing paradox of why both contextual similarity and dissimilarity promote TOM.

How are experiences of events used to update knowledge of predictive relations in semantic memory? We examined the roles of anterior-lateral entorhinal cortex (alEC), important for encoding recently experienced temporal relations, and middle temporal gyrus (MTG), involved in familiar event concepts. Participants underwent fMRI during exposure to novel event sequences and a memory probe phase (Session 1) and the same process a week later (Session 2). Across distinct sequences, predictive relations among similar events could either be Consistent, or the roles of the events could swap (Inconsistent). We examined the effect of Consistency on the strength of relational memory content. Areas that integrate across diverse experiences should be aided in the Consistent condition. We found that alEC performed this integrative role in Session 1, and at Session 2, similar effects were also observed in MTG. We suggest that these areas both contribute to building relational knowledge from experience.

An outstanding issue in cognitive science is whether the computational principles that apply to causal reasoning also guide the way that participants encode the relations among events in memory. The outcome density effect is a behavioral pattern in causal reasoning in which participants’ causal ratings linearly decrease as the base rate of the effect also decreases, even while contingency remains 0. It is key evidence for Bayesian models of causal reasoning as it reflects decreasing uncertainty. We queried whether it may also, separately, affect memory for events. We measured both recall and causal ratings in a causal learning task to test whether the outcome density manipulation affects causal judgment, recall, or both. We replicated the outcome density effect on causal judgment, lending support to Bayesian models, but found that memory instead exhibited a U-shaped relationship with base rate, and therefore, causal judgment and memory had divergent signatures.

BackgroundPeople with schizophrenia (SZ) exhibit impaired episodic memory when relating objects to each other in time and space. Empirical studies and computational models suggest that low-frequency neural oscillations may be a mechanism by which the brain keeps track of temporal relationships during encoding and retrieval, with modulation of oscillatory power as sequences are learned. It is unclear whether sequence memory deficits in SZ are associated with altered neural oscillations.MethodsUsing electroencephalography, this study examined neural oscillations in 51 healthy control subjects and 37 people with SZ during a temporal sequence learning task. Multiple 5-object picture sequences were presented across 4 study-test blocks in either fixed or random order. Participants answered semantic questions for each object (e.g., living/nonliving), and sequence memory was operationalized as faster responses for fixed versus random sequences. Differences in oscillatory power between fixed versus random sequences provided a neural index of temporal sequence memory.ResultsAlthough both groups showed reaction time differences in late blocks (blocks 3 and 4), this evidence of sequence memory was reduced in people with SZ relative to healthy control subjects. Decreases in globally distributed prestimulus alpha (8-12 Hz) and beta 1 (13-20 Hz) power for fixed versus random sequences in late blocks were also attenuated in people with SZ relative to healthy control subjects. Moreover, changes in oscillatory power predicted individual reaction time differences and fully mediated the relationship between group and sequence memory.ConclusionsDisrupted modulation of alpha and beta 1 electroencephalography oscillations is a candidate mechanism of temporal sequence memory deficits in people with SZ.

Retrieval practice improves retention of tested information, and it can either impair or facilitate retention of untested information. Here, we investigated how semantic relatedness, episodic context, and sleep-dependent memory consolidation determine the effects of retrieval practice on retention of untested items. Participants studied lists of scene-word associations. Each scene was associated with two different words ("pairmates") that were either semantically related or unrelated and either in the same (temporally close) or different lists (temporally far). In three experiments, retrieval practice of scene-word associations facilitated retention of unpracticed, temporally close pairmates and impaired retention of temporally far, semantically unrelated pairmates. Critically, retrieval practice impaired retention of temporally far, semantically related pairmates if participants were unable to sleep during the retention interval, but it facilitated retention of these items if participants were able to sleep. Our findings suggest that sleep extends the benefits of testing to related information learned in temporally separate episodes.

Life's events are scattered throughout time, yet we often recall different events in the context of an integrated narrative. Prior research suggests that the hippocampus, which supports memory for past events, can support the integration of overlapping associations or separate events in memory. However, the conditions that lead to hippocampus-dependent memory integration are unclear. We used functional brain imaging to test whether the opportunity to form a larger narrative (narrative coherence) drives hippocampal memory integration. During encoding of fictional stories, patterns of hippocampal activity, including activity at boundaries between events, were more similar between distant events that formed one coherent narrative, compared with overlapping events taken from unrelated narratives. One day later, the hippocampus preferentially supported detailed recall of coherent narrative events, through reinstatement of hippocampal activity patterns from encoding. These findings demonstrate a key function of the hippocampus: the integration of events into a narrative structure for memory.

The hippocampus, well known for its role in episodic memory, might also be an important brain region for extracting structure from our experiences in order to guide future decisions. Recent evidence in rodents suggests that the hippocampus supports decision making by representing task structure in cooperation with the orbitofrontal cortex (OFC). Here, we examine how the human hippocampus and OFC represent task structure during an associative learning task that required learning of both context-determined and context-invariant probabilistic associations. We find that after learning, hippocampal and lateral OFC representations differentiated between context-determined and context-invariant task structures. The degree of this differentiation within the hippocampus and lateral OFC is highly correlated. These results advance our understanding of the hippocampus and suggest that the hippocampus and OFC support goal-directed behavior by representing information that guides the selection of appropriate decision strategies.

Curiosity states benefit memory for target information, but also incidental information presented during curiosity states. However, it is not known whether incidental curiosity-enhanced memory depends on when incidental information during curiosity states is encountered. Here, participants incidentally encoded unrelated face images at different time points while they anticipated answers to trivia questions. Across two experiments, we found memory enhancements for unrelated faces presented during high-curiosity compared with low-curiosity states, but only when presented shortly after a trivia question. This suggests processes associated with the elicitation of curiosity-but not sustained anticipation or the satisfaction of curiosity-enhance memory for incidental information.

Episodic memory depends on interactions between the hippocampus and interconnected neocortical regions. Here, using data-driven analyses of resting-state functional magnetic resonance imaging (fMRI) data, we identified the networks that interact with the hippocampus-the default mode network (DMN) and a "medial temporal network" (MTN) that included regions in the medial temporal lobe (MTL) and precuneus. We observed that the MTN plays a critical role in connecting the visual network to the DMN and hippocampus. The DMN could be further divided into 3 subnetworks: a "posterior medial" (PM) subnetwork comprised of posterior cingulate and lateral parietal cortices; an "anterior temporal" (AT) subnetwork comprised of regions in the temporopolar and dorsomedial prefrontal cortex; and a "medial prefrontal" (MP) subnetwork comprised of regions primarily in the medial prefrontal cortex (mPFC). These networks vary in their functional connectivity (FC) along the hippocampal long axis and represent different kinds of information during memory-guided decision-making. Finally, a Neurosynth meta-analysis of fMRI studies suggests new hypotheses regarding the functions of the MTN and DMN subnetworks, providing a framework to guide future research on the neural architecture of episodic memory.

Transcranial direct current stimulation (tDCS) to the left prefrontal cortex has been shown to produce broad behavioral effects including enhanced learning and vigilance. Still, the neural mechanisms underlying such effects are not fully understood. Furthermore, the neural underpinnings of repeated stimulation remain understudied. In this work, we evaluated the effects of the repetition and intensity of tDCS on cerebral perfusion [cerebral blood flow (CBF)]. A cohort of 47 subjects was randomly assigned to one of the three groups. tDCS of 1- or 2-mA was applied to the left prefrontal cortex on three consecutive days, and resting CBF was quantified before and after stimulation using the arterial spin labeling MRI and then compared with a group that received sham stimulation. A widespread decreased CBF was found in a group receiving sham stimulation across the three post-stimulation measures when compared with baseline. In contrast, only slight decreases were observed in the group receiving 2-mA stimulation in the second and third post-stimulation measurements, but more prominent increased CBF was observed across several brain regions including the locus coeruleus (LC). The LC is an integral region in the production of norepinephrine and the noradrenergic system, and an increased norepinephrine/noradrenergic activity could explain the various behavioral findings from the anodal prefrontal tDCS. A decreased CBF was observed in the 1-mA group across the first two post-stimulation measurements, similar to the sham group. This decreased CBF was apparent in only a few small clusters in the third post-stimulation scan but was accompanied by an increased CBF, indicating that the neural effects of stimulation may persist for at least 24 h and that the repeated stimulation may produce cumulative effects.

The goal of this study was to use transcranial direct current stimulation (tDCS) to examine the role of the prefrontal cortex (PFC) in neural oscillatory activity associated with proactive cognitive control in schizophrenia. To do so, we tested the impact of PFC-targeted tDCS on behavioral and electrophysiological markers of proactive cognitive control engagement in individuals with schizophrenia. Using a within-participants, double-blinded, sham-controlled crossover design, we recorded EEG while participants with schizophrenia completed a proactive cognitive control task (the Dot Pattern Expectancy (DPX) Task), after receiving 20 min of active prefrontal stimulation at 2 mA or sham stimulation. We hypothesized that active stimulation would enhance proactive cognitive control, leading to changes in behavioral performance on the DPX task and in activity in the gamma frequency band during key periods of the task designed to tax proactive cognitive control. The results showed significant changes in the pattern of error rates and increases in EEG gamma power as a function of tDCS condition (active or sham), that were indicative of enhanced proactive cognitive control. These findings, considered alongside our previous work in healthy adults, provides novel support for the role gamma oscillations in proactive cognitive control and they suggest that frontal tDCS may be a promising approach to enhance proactive cognitive control in schizophrenia.

Cognitive maps enable efficient inferences from limited experience that can guide novel decisions. We tested whether the hippocampus (HC), entorhinal cortex (EC), and ventromedial prefrontal cortex (vmPFC)/medial orbitofrontal cortex (mOFC) organize abstract and discrete relational information into a cognitive map to guide novel inferences. Subjects learned the status of people in two unseen 2D social hierarchies, with each dimension learned on a separate day. Although one dimension was behaviorally relevant, multivariate activity patterns in HC, EC, and vmPFC/mOFC were linearly related to the Euclidean distance between people in the mentally reconstructed 2D space. Hubs created unique comparisons between the hierarchies, enabling inferences between novel pairs. We found that both behavior and neural activity in EC and vmPFC/mOFC reflected the Euclidean distance to the retrieved hub, which was reinstated in HC. These findings reveal how abstract and discrete relational structures are represented, are combined, and enable novel inferences in the human brain.

BackgroundProviding early psychosis (EP) individuals with family psychoeducation (FPE) can reduce symptoms and improve clinical outcomes. However, relational memory problems may limit prospective utilization of FPE information. This study examines whether memory for FPE can be improved by testing participants during the initial FPE workshop presentation.MethodData were obtained from 20 people with EP and 20 demographically matched healthy comparison subjects (HC). During session one, FPE was presented in small group workshops, with half of the information re-studied twice (re-study condition) and the remaining information tested twice using cued recall tasks (retrieval practice condition). One week later (session two), delayed cued recall was tested for all FPE information. "Testing effects" (i.e., better memory following retrieval practice versus re-study) were examined across all items (standard analysis) and also limited to items successfully retrieved during session one (conditionalized analysis).ResultsHC had better initial recall and learned more over the two retrieval practice trials than EP. However, HC also lost more information than EP over the one-week delay. Both groups produced a significant testing effect. This effect was smaller in EP versus HC across all test items, but did not differ for the conditionalized analysis. Negative symptoms were inversely correlated with delayed cued recall in EP.ConclusionsEP participants benefit from retrieval practice, with participants with less severe negative symptoms showing the greatest benefit. These results encourage use of memory tests during group psychoeducation to improve subsequent long-term recall of clinically relevant information.

Recent research has highlighted a role for the hippocampus and a Posterior Medial cortical network in signaling event boundaries. However, little is known about whether or how these neural processes change over the course of healthy aging. Here, 546 cognitively normal participants 18-88 years old viewed a short movie while brain activity was measured using fMRI. The hippocampus and regions of the Posterior Medial network show increased activity at event boundaries, but these boundary-evoked responses decrease with age. Boundary-evoked activity in the posterior hippocampus predicts performance on a separate test of memory for stories, suggesting that hippocampal activity during event segmentation may be a broad indicator of individual differences in episodic memory ability. In contrast, boundary-evoked responses in the medial prefrontal cortex and middle temporal gyrus increase across the age range. These findings suggest that aging may alter neural processes for segmenting and remembering continuous real-world experiences.

When acute stress is experienced immediately after memory encoding (i.e., post-encoding stress) it can significantly impact subsequent memory for that event. For example, recent work has suggested that post-encoding stress occurring in a different context from encoding impairs memory. However, the neural processes underlying these effects are poorly understood. We aimed to expand this understanding by conducting an analysis of resting functional connectivity in the period following post-encoding stress that occurred in a different context than encoding, using seed regions in the medial temporal lobes known for their roles in memory. In the current study of 44 males randomized to stress (n = 23) or control (n = 21) groups, we found that stress increased cortisol, impaired recollection of neutral materials, and altered functional connectivity with medial temporal lobe regions. Although stress did not significantly alter hippocampus-amygdala functional connectivity, relative to participants in the control group, participants in the post-encoding stress group showed lower functional connectivity between the hippocampus and a region with a peak in the superior temporal gyrus. Across participants in both groups, functional connectivity between these regions was related to greater increases in cortisol, and it was also inversely related to recollection of neutral materials. In contrast, the stress group showed greater parahippocampal cortex functional connectivity with a region in the left middle temporal gyrus than the control group. Moreover, greater functional connectivity between the parahippocampal cortex and the observed cluster in the middle temporal gyrus was associated with greater cortisol changes from pre- to post-manipulation, but was not related to differences in memory. The results show that post-encoding stress can alter the resting-state functional connectivity between the medial temporal lobe and neocortex, which may help explain how stress impacts memory.

Episodic memory reflects the ability to recollect the temporal and spatial context of past experiences. Episodic memories depend on the hippocampus but have been proposed to undergo rapid forgetting unless consolidated during offline periods such as sleep to neocortical areas for long-term storage. Here, we propose an alternative to this standard systems consolidation theory (SSCT) - a contextual binding account - in which the hippocampus binds item-related and context-related information. We compare these accounts in light of behavioural, lesion, neuroimaging and sleep studies of episodic memory and contend that forgetting is largely due to contextual interference, episodic memory remains dependent on the hippocampus across time, contextual drift produces post-encoding activity and sleep benefits memory by reducing contextual interference.

This study examined the effects of stimulation targeting dorsolateral prefrontal cortex (DLPFC) on behavioral and neural oscillatory markers of proactive cognitive control in healthy adults. We hypothesized that active stimulation targeting the DLPFC would enhance proactive control compared to sham, leading to changes in the pattern of error rates and gamma-band power on the Dot Pattern Expectancy (DPX) task. We recorded EEG while participants completed the DPX, after receiving either 20 minutes of active DLPFC stimulation at 2 mA or sham stimulation in a counterbalanced within-participants design. The results showed significant tDCS-induced changes in the pattern of error rates on the DPX task indicative of enhanced proactive control, as well as predicted increases in gamma power associated with the engagement of proactive control. These results provide support for the role of DLPFC-mediated gamma activity in proactive cognitive control, and further, indicate that proactive control can be enhanced with non-invasive neurostimulation.

The efficacy of cognitive training is controversial, and research progress in the field requires an understanding of factors that promote transfer of training gains and their relationship to changes in brain activity. One such factor may be adaptive task difficulty, as adaptivity is predicted to facilitate more efficient processing by creating a prolonged mismatch between the supply of, and the demand upon, neural resources. To test this hypothesis, we measured behavioral and neural plasticity in fMRI sessions before and after 10 sessions of working memory updating (WMU) training, in which the difficulty of practiced tasks either adaptively increased in response to performance or was fixed. Adaptive training resulted in transfer to an untrained episodic memory task and activation decreases in striatum and hippocampus on a trained WMU task, and the amount of training task improvement was associated with near transfer to other WMU tasks and with hippocampal activation changes on both near and far transfer tasks. These findings suggest that cognitive training programs should incorporate adaptive task difficulty to broaden transfer of training gains and maximize efficiency of task-related brain activity.

Emotional experiences are typically remembered with a greater sense of recollection than neutral experiences, but memory benefits for emotional items do not typically extend to their source contexts. Item and source memory have been attributed to different subregions of the medial temporal lobes (MTL), but it is unclear how emotional item recollection fits into existing models of MTL function and, in particular, what is the role of the hippocampus. To address these issues, we used high-resolution functional magnetic resonance imaging (fMRI) to examine MTL contributions to successful emotional item and context encoding. The results showed that emotional items were recollected more often than neutral items. Whereas amygdala and perirhinal cortex (PRC) activity supported the recollection advantage for emotional items, hippocampal and parahippocampal cortex activity predicted subsequent source memory for both types of items, reflecting a double dissociation between anterior and posterior MTL regions. In addition, amygdala activity during encoding modulated the relationships of PRC activity and hippocampal activity to subsequent item recollection and source memory, respectively. Specifically, whereas PRC activity best predicted subsequent item recollection when amygdala activity was relatively low, hippocampal activity best predicted source memory when amygdala activity was relatively high. We interpret these findings in terms of complementary compared to synergistic amygdala-MTL interactions. The results suggest that emotion-related enhancements in item recollection are supported by an amygdala-PRC pathway, which is separable from the hippocampal pathway that binds items to their source context.

Episodic memory is known to rely on the hippocampus, but how the hippocampus organizes different episodes to permit their subsequent retrieval remains controversial. One major area of debate hinges on a discrepancy between two hypothesized roles of the hippocampus: differentiating between similar events to reduce interference and assigning similar representations to events that share overlapping items and contextual information. Here, we used multivariate analyses of activity patterns measured with fMRI to characterize how the hippocampus distinguishes between memories based on similarity at the level of items and/or context. Hippocampal activity patterns discriminated between events that shared either item or context information but generalized across events that shared similar item-context associations. The current findings provide evidence that, whereas the hippocampus can reduce mnemonic interference by separating events that generalize along a single attribute dimension, overlapping hippocampal codes may support memory for events with overlapping item-context relations. This lends new insights into the way the hippocampus may balance multiple mnemonic operations in adaptively guiding behavior.

Remembering is a complex process that involves recalling specific details, such as who you were with when you celebrated your last birthday, as well as contextual information, such as the place where you celebrated. It is well established that the act of remembering enhances long-term retention of the retrieved information, but the neural and cognitive mechanisms that drive memory enhancement are not yet understood. One possibility is that the process of remembering results in reactivation of the broader episodic context. Consistent with this idea, in two experiments, we found that multiple retrieval attempts enhanced long-term retention of both the retrieved object and the nontarget object that shared scene context, compared with a restudy control. Using representational similarity analysis of fMRI data in experiment 2, we found that retrieval resulted in greater neural reactivation of both the target objects and contextually linked objects compared with restudy. Furthermore, this reactivation occurred in a network of medial and lateral parietal lobe regions that have been linked to episodic recollection. The results demonstrate that retrieving a memory can enhance retention of information that is linked in the broader event context and the hippocampus and a posterior medial network of parietal cortical areas (also known as the Default Network) play complementary roles in supporting the reactivation of episodically linked information during retrieval.

Historically, research on the cognitive processes that support human memory proceeded, to a large extent, independently of research on the neural basis of memory. Accumulating evidence from neuroimaging, however, has enabled the field to develop a broader and more integrative perspective. Here, we briefly outline how advances in cognitive neuroscience can potentially shed light on concepts and controversies in human memory research. We argue that research on the functional properties of cortico-hippocampal networks informs us about how memories might be organized in the brain, which, in turn, helps to reconcile seemingly disparate perspectives in cognitive psychology. Finally, we discuss several open questions and directions for future research.

People can employ adaptive strategies to increase the likelihood that previously encoded information will be successfully retrieved. One such strategy is to constrain retrieval toward relevant information by reimplementing the neurocognitive processes that were engaged during encoding. Using EEG, we examined the temporal dynamics with which constraining retrieval toward semantic versus nonsemantic information affects the processing of new "foil" information encountered during a memory test. Time-frequency analysis of EEG data acquired during an initial study phase revealed that semantic compared with nonsemantic processing was associated with alpha decreases in a left frontal electrode cluster from around 600 msec after stimulus onset. Successful encoding of semantic versus nonsemantic foils during a subsequent memory test was related to decreases in alpha oscillatory activity in the same left frontal electrode cluster, which emerged relatively late in the trial at around 1000-1600 msec after stimulus onset. Across participants, left frontal alpha power elicited by semantic processing during the study phase correlated significantly with left frontal alpha power associated with semantic foil encoding during the memory test. Furthermore, larger left frontal alpha power decreases elicited by semantic foil encoding during the memory test predicted better subsequent semantic foil recognition in an additional surprise foil memory test, although this effect did not reach significance. These findings indicate that constraining retrieval toward semantic information involves reimplementing semantic encoding operations that are mediated by alpha oscillations and that such reimplementation occurs at a late stage of memory retrieval, perhaps reflecting additional monitoring processes.

The hippocampus plays a critical role in spatial and episodic memory. Mechanistic models predict that hippocampal subfields have computational specializations that differentially support memory. However, there is little empirical evidence suggesting differences between the subfields, particularly in humans. To clarify how hippocampal subfields support human spatial and episodic memory, we developed a virtual reality paradigm where participants passively navigated through houses (spatial contexts) across a series of videos (episodic contexts). We then used multivariate analyses of high-resolution fMRI data to identify neural representations of contextual information during recollection. Multi-voxel pattern similarity analyses revealed that CA1 represented objects that shared an episodic context as more similar than those from different episodic contexts. CA23DG showed the opposite pattern, differentiating between objects encountered in the same episodic context. The complementary characteristics of these subfields explain how we can parse our experiences into cohesive episodes while retaining the specific details that support vivid recollection.

The development and application of concepts is a critical component of cognition. Although concepts can be formed on the basis of simple perceptual or semantic features, conceptual representations can also capitalize on similarities across feature relationships. By representing these types of higher-order relationships, concepts can simplify the learning problem and facilitate decisions. Despite this, little is known about the neural mechanisms that support the construction and deployment of these kinds of higher-order concepts during learning. To address this question, we combined a carefully designed associative learning task with computational model-based functional magnetic resonance imaging (fMRI). Participants were scanned as they learned and made decisions about sixteen pairs of cues and associated outcomes. Associations were structured such that individual cues shared feature relationships, operationalized as shared patterns of cue pair-outcome associations. In order to capture the large number of possible conceptual representational structures that participants might employ and to evaluate how conceptual representations are used during learning, we leveraged a well-specified Bayesian computational model of category learning [1]. Behavioral and model-based results revealed that participants who displayed a tendency to link experiences in memory benefitted from faster learning rates, suggesting that the use of the conceptual structure in the task facilitated decisions about cue pair-outcome associations. Model-based fMRI analyses revealed that trial-by-trial integration of cue information into higher-order conceptual representations was supported by an anterior temporal (AT) network of regions previously implicated in representing complex conjunctions of features and meaning-based information.

Life's episodes unfold against a context that changes with time. Recent neuroimaging studies have revealed significant findings about how specific areas of the human brain may support the representation of temporal information in memory. A consistent theme in these studies is that the hippocampus appears to play a central role in representing temporal context, as operationalized in neuroimaging studies of arbitrary lists of items, sequences of items, or meaningful, lifelike events. Additionally, activity in a posterior medial cortical network may reflect the representation of generalized temporal information for meaningful events. The hippocampus, posterior medial network, and other regions-particularly in prefrontal cortex-appear to play complementary roles in memory for temporal context.