Variation in encoding context benefits item recognition.

The current study assesses whether varying the encoding context of a repeated event is a potential strategy to improve recognition memory across retrieval contexts. Context variability, also known as encoding variability, has historically been investigated primarily using recall and cued recall tasks, with the consensus being that encoding variability is not necessarily beneficial for episodic retrieval. However, recent studies (see text) suggest that test type may determine the strategy's effectiveness. Aligned with these recent findings, we found consistent benefits to simple item recognition when a word was studied in more variable contexts compared to less variable contexts across four experiments. This main effect of context variability occurred when crossed with a manipulation of repetition spacing and when crossed with a manipulation of encoding-retrieval context match. Variation in encoding contexts beyond the future retrieval context led to better item recognition than repeated study exposures within the future retrieval context. We argue that the current study and other recent findings indicate a need to re-evaluate the historical consensus on encoding variability as a beneficial strategy for learning.

Cognitive tasks affect the relationship between representational pattern similarity and subsequent item memory in the hippocampus.

Episodic memories are records of personally experienced events, coded neurally via the hippocampus and surrounding medial temporal lobe cortex. Information about the neural signal corresponding to a memory representation can be measured in fMRI data when the pattern across voxels is examined. Prior studies have found that similarity in the voxel patterns across repetition of a to-be-remembered stimulus predicts later memory retrieval, but the results are inconsistent across studies. The current study investigates the possibility that cognitive goals (defined here via the task instructions given to participants) during encoding affect the voxel pattern that will later support memory retrieval, and therefore that neural representations cannot be interpreted based on the stimulus alone. The behavioral results showed that exposure to variable cognitive tasks across repetition of events benefited subsequent memory retrieval. Voxel patterns in the hippocampus indicated a significant interaction between cognitive tasks (variable vs. consistent) and memory (remembered vs. forgotten) such that reduced voxel pattern similarity for repeated events with variable cognitive tasks, but not consistent cognitive tasks, supported later memory success. There was no significant interaction in neural pattern similarity between cognitive tasks and memory success in medial temporal cortices or lateral occipital cortex. Instead, higher similarity in voxel patterns in right medial temporal cortices was associated with later memory retrieval, regardless of cognitive task. In conclusion, we found that the relationship between pattern similarity across repeated encoding and memory success in the hippocampus (but not medial temporal lobe cortex) changes when the cognitive task during encoding does or does not vary across repetitions of the event.

The interaction of relational encoding and unitization: Effects on medial temporal lobe processing during retrieval.

In retrieval of typical episodic memories, recollection leads to retrieval of context details whereas familiarity is only diagnostic for item memory. Unitization is an encoding strategy that allows context details to be processed as item features and therefore increases the involvement of familiarity-based recognition in retrieval of these context details. Relational encoding is a hippocampally-dependent process that stores items and contexts independently. Our previous study Tu and Diana [1] concluded that mixing unitized and non-unitized context details in the same episode reduced the contribution of familiarity to retrieval of any one detail. In the current study, we modified the paradigm by removing visual cues to the context details and the condition-specific blocking during test. Surprisingly, the behavioral data diverged from our 2016 study and indicated that the two manipulated context details in the modified paradigm were processed independently of one another. Neuroimaging data further revealed anterior hippocampal activation was associated with unitization of source information as compared to relational encoding. We also found the predicted increase in bilateral perirhinal cortex activation and decrease in parahippocampal cortex activation during retrieval of unitized color information when compared to relationally-encoded color information. We did not find that same predicted pattern of differences due to unitization of size information.

Neural congruence between intertemporal and interpersonal self-control: Evidence from delay and social discounting.

Behavioral studies using delay and social discounting as indices of self-control and altruism, respectively, have revealed functional similarities between farsighted and social decisions. However, neural evidence for this functional link is lacking. Twenty-five young adults completed a delay and social discounting task during fMRI scanning. A spatiotemporal partial least squares analysis revealed that both forms of discounting were well characterized by a pattern of brain activity in areas comprising frontoparietal control, default, and mesolimbic reward networks. Both forms of discounting appear to draw on common neurocognitive mechanisms, regardless of whether choices involve intertemporal or interpersonal outcomes. We also observed neural profiles differentiating between high and low discounters. High discounters were well characterized by increased medial temporal lobe and limbic activity. In contrast, low discount rates were associated with activity in the medial prefrontal cortex and right temporoparietal junction. This pattern may reflect biological mechanisms underlying behavioral heterogeneity in discount rates.

Event-related potentials during encoding: Comparing unitization to relational processing.

Context details are typically encoded into episodic memory via arbitrary associations to the relevant item, known as relational binding. Subsequent retrieval of those context details is primarily supported by recollection. Research suggests that context retrieval can rely on familiarity if the context details are "unitized" and thereby encoded as features of the item itself in a single new representation. With most investigations into unitization focusing on the contributions of familiarity and recollection during retrieval, little is known about unitization during encoding. In an effort to begin understanding unitization as an encoding process, we used event-related potentials to monitor brain activity while participants were instructed to encode words with color information using relational association or unitization. Results showed that unitization-based encoding elicited significantly more negative potentials in the left parietal region than relational encoding during presentation of the second segment of strategically-specific sentences. This difference continued through presentation of the third sentence segment, becoming less lateralized, and ended before the final two segments were presented. During the mental imagery period, unitization-based encoding elicited significantly more positive potentials than relational encoding in the first 200ms centrally and from 400 through 1000ms in left fronto-temporal and parieto-occipital regions. Our findings indicate that unitization and relational processing diverged at approximately the time that the context item was presented in the relational condition. During mental imagery, unitization diverged from relational processing immediately, suggesting that unitization affected the nature of the item representation, and possibly the brain regions involved, during memory encoding.

The effect of binaural beats on verbal working memory and cortical connectivity.

OBJECTIVE: Synchronization in activated regions of cortical networks affect the brain's frequency response, which has been associated with a wide range of states and abilities, including memory. A non-invasive method for manipulating cortical synchronization is binaural beats. Binaural beats take advantage of the brain's response to two pure tones, delivered independently to each ear, when those tones have a small frequency mismatch. The mismatch between the tones is interpreted as a beat frequency, which may act to synchronize cortical oscillations. Neural synchrony is particularly important for working memory processes, the system controlling online organization and retention of information for successful goal-directed behavior. Therefore, manipulation of synchrony via binaural beats provides a unique window into working memory and associated connectivity of cortical networks. APPROACH: In this study, we examined the effects of different acoustic stimulation conditions during an N-back working memory task, and we measured participant response accuracy and cortical network topology via EEG recordings. Six acoustic stimulation conditions were used: None, Pure Tone, Classical Music, 5 Hz binaural beats, 10 Hz binaural beats, and 15 Hz binaural beats. MAIN RESULTS: We determined that listening to 15 Hz binaural beats during an N-Back working memory task increased the individual participant's accuracy, modulated the cortical frequency response, and changed the cortical network connection strengths during the task. Only the 15 Hz binaural beats produced significant change in relative accuracy compared to the None condition. SIGNIFICANCE: Listening to 15 Hz binaural beats during the N-back task activated salient frequency bands and produced networks characterized by higher information transfer as compared to other auditory stimulation conditions.

The neurocognitive basis of borrowed context information.

Falsely remembered items can be accompanied by episodic context retrieval. This finding is difficult to explain because there is no episode that binds the remembered item to the experimenter-controlled context features. The current study examines the neural correlates of false context retrieval when the context features can be traced to encoding episodes of semantically-similar items. Our neuroimaging results support a "dissociated source" mechanism for context borrowing in false memory. We found that parahippocampal cortex (PHc) activation, thought to indicate context retrieval, was greater during trials that involved context borrowing (an incorrect, but plausible source decision) than during baseline correct context retrieval. In contrast, hippocampal activation, thought to indicate retrieval of an episodic binding, was stronger during correct source retrieval than during context borrowing. Vivid context retrieval during false recollection experiences was also indicated by increased activation in visual perceptual regions for context borrowing as compared to other incorrect source judgments. The pattern of findings suggests that context borrowing can arise when unusually strong activation of a semantically-related item's contextual features drives relatively weak retrieval of the associated episodic binding with failure to confirm the item information within that binding. This dissociated source retrieval mechanism suggests that context-driven episodic retrieval does not necessarily lead to retrieval of specific item details. That is, source information can be retrieved in the absence of item memory.

Neural correlates of temporal context retrieval for abstract scrambled phrases: Reducing narrative and familiarity-based strategies.

Temporal context, memory for the timing of events, can be assessed using non-temporal strategies such as relative familiarity or inference from a semantic narrative. Neuroimaging studies, which have previously encouraged such strategies, find similar patterns of brain regions involved in both temporal and non-temporal context memory. The present study aims to investigate whether previous findings are driven by the use of non-temporal strategies or whether the same pattern of brain regions is identified when relative familiarity and semantic narrative strategies are discouraged. We used abstract phrases (e.g. alone me leave) created by scrambling familiar three-word phrases. The words in the phrases were less concrete than the object image stimuli used in previous studies of temporal context memory (Jenkins and Ranganath, 2010) and were presented quickly while participants read each word aloud. This differed from previous studies in which participants were encouraged to use narrative strategies during encoding (Tubridy and Davachi, 2011) and was designed to discourage use of narrative strategies. The relative familiarity of the words within each phrase was similar and likely not diagnostic of word order during encoding, in order to minimize the use of relative familiarity strategies. Neuroimaging results indicate that temporal context retrieval was associated with the hippocampus, parahippocampal cortex, ventromedial prefrontal cortex, and retrosplenial cortex, which are regions consistent with the retrieval of non-temporal context in episodic memory, suggesting that previous findings were not driven entirely by non-temporal strategies but rather that temporal memory relies on similar brain regions to non-temporal memory.

Parahippocampal Cortex Processes the Nonspatial Context of an Event.

Parahippocampal cortex (PHc) is known to process spatial information, both in perceptual and episodic memory studies. However, recent theories propose an expanded role for PHc in processing context information in general, whether spatial or nonspatial. The current study used a source memory paradigm to investigate encoding and retrieval of nonspatial context information. Human participants were asked to judge lexical aspects of word stimuli and to retrieve those judgments during a later memory test. Anterior PHc showed significantly greater activation for items associated with correct source judgments than items associated with incorrect source judgments during both encoding and retrieval phases. These findings suggest that the role of PHc in episodic memory cannot be limited to spatial information.

The Effect of Binaural Beats on Visuospatial Working Memory and Cortical Connectivity.

Binaural beats utilize a phenomenon that occurs within the cortex when two different frequencies are presented separately to each ear. This procedure produces a third phantom binaural beat, whose frequency is equal to the difference of the two presented tones and which can be manipulated for non-invasive brain stimulation. The effects of binaural beats on working memory, the system in control of temporary retention and online organization of thoughts for successful goal directed behavior, have not been well studied. Furthermore, no studies have evaluated the effects of binaural beats on brain connectivity during working memory tasks. In this study, we determined the effects of different acoustic stimulation conditions on participant response accuracy and cortical network topology, as measured by EEG recordings, during a visuospatial working memory task. Three acoustic stimulation control conditions and three binaural beat stimulation conditions were used: None, Pure Tone, Classical Music, 5Hz binaural beats, 10Hz binaural beats, and 15Hz binaural beats. We found that listening to 15Hz binaural beats during a visuospatial working memory task not only increased the response accuracy, but also modified the strengths of the cortical networks during the task. The three auditory control conditions and the 5Hz and 10Hz binaural beats all decreased accuracy. Based on graphical network analyses, the cortical activity during 15Hz binaural beats produced networks characteristic of high information transfer with consistent connection strengths throughout the visuospatial working memory task.

Frontotemporal function]al connectivity and executive functions contribute to episodic memory performance.

The contributions of hemispheric-specific electrophysiology (electroencephalogram or EEG) and independent executive functions (inhibitory control, working memory, cognitive flexibility) to episodic memory performance were examined using abstract paintings. Right hemisphere frontotemporal functional connectivity during encoding and retrieval, measured via EEG alpha coherence, statistically predicted performance on recency but not recognition judgments for the abstract paintings. Theta coherence, however, did not predict performance. Likewise, cognitive flexibility statistically predicted performance on recency judgments, but not recognition. These findings suggest that recognition and recency operate via separate electrophysiological and executive mechanisms.

Temporal context processing within hippocampal subfields.

The episodic memory system can differentiate similar events based on the temporal information associated with the events. Temporal context, which is at least partially determined by the events that precede or follow the critical event, may be a cue to differentiate events. The purpose of the present study is to investigate whether the hippocampal dentate gyrus (DG)/CA3 and CA1 subfields are sensitive to changes in temporal context and, if so, whether the subregions show a linear or threshold-like response to similar temporal contexts. Participants incidentally encoded a series of object picture triplets and 20 of them were included in final analyses. The third picture in each triplet was operationally defined as the target and the first two pictures served as temporal context for the target picture. Each target picture was presented twice with temporal context manipulated to be either repeated, high similarity, low similarity, or new on the second presentation. We extracted beta parameters for the repeated target as a function of the type of temporal context. We expected to see repetition suppression, a reduction in the beta values, in response to repetition of the target. If temporal context information is included in the representation of the target within a given region, this repetition suppression should be greater for target images that were preceded by their original context than for target images preceded by a new context. Neuroimaging results showed that CA1, but not DG/CA3, modifies the target's representation based on its temporal context. Right CA1 did not distinguish high similarity temporal context from repeated context but did distinguish low similarity temporal context from repeated context. These results indicate that CA1 is sensitive to temporal context and suggest that it does not differentiate between a substantially similar temporal context and an identical temporal context. In contrast, DG/CA3 does not appear to process temporal context as defined in the current experiment.

Two are not better than one: Combining unitization and relational encoding strategies.

In recognition memory, recollection is defined as retrieval of the context associated with an event, whereas familiarity is defined as retrieval based on item strength alone. Recent studies have shown that conventional recollection-based tasks, in which context details are manipulated for source memory assessment at test, can also rely on familiarity when context information is "unitized" with the relevant item information at encoding. Unlike naturalistic episodic memories that include many context details encoded in different ways simultaneously, previous studies have focused on unitization and its effect on the recognition of a single context detail. To further understand how various encoding strategies operate on item and context representations, we independently assigned unitization and relational association to 2 context details (size and color) of each item and tested the contribution of recollection and familiarity to source recognition of each detail. The influence of familiarity on retrieval of each context detail was compared as a function of the encoding strategy used for each detail. Receiver operating characteristic curves suggested that the unitization effect was not additive and that similar levels of familiarity occurred for 1 or multiple details when unitization was the only strategy applied during encoding. On the other hand, a detrimental effect was found when relational encoding and unitization were simultaneously applied to 1 item such that a salient nonunitized context detail interfered with the effortful processing required to unitize an accompanying context detail. However, this detrimental effect was not reciprocal and possibly dependent on the nature of individual context details. (PsycINFO Database Record

Associative memory and its cerebral correlates in Alzheimer׳s disease: evidence for distinct deficits of relational and conjunctive memory.

This study investigated the impact of Alzheimer׳s disease (AD) on conjunctive and relational binding in episodic memory. Mild AD patients and controls had to remember item-color associations by imagining color either as a contextual association (relational memory) or as a feature of the item to be encoded (conjunctive memory). Patients׳ performance in each condition was correlated with cerebral metabolism measured by FDG-PET. The results showed that AD patients had an impaired capacity to remember item-color associations, with deficits in both relational and conjunctive memory. However, performance in the two kinds of associative memory varied independently across patients. Partial Least Square analyses revealed that poor conjunctive memory was related to hypometabolism in an anterior temporal-posterior fusiform brain network, whereas relational memory correlated with metabolism in regions of the default mode network. These findings support the hypothesis of distinct neural systems specialized in different types of associative memory and point to heterogeneous profiles of memory alteration in Alzheimer׳s disease as a function of damage to the respective neural networks.

Parahippocampal cortex activation during context reinstatement predicts item recollection.

Episodic memory is the binding of an event with information about the context in which that event (or item) was experienced. The context of an event may include its spatial and temporal location as well as goal-directed, conscious thoughts evoked during the event. We call this latter type of information cognitive context. The binding of items and context (BIC) theory of medial temporal lobe function proposes that the parahippocampal cortex (PHc) plays a key role in processing cognitive context. Therefore, we predicted that activity in the PHc during reactivation of a previously experienced cognitive context would be correlated with item recollection, even when the associated item and its episodic binding had not yet been retrieved. Using a novel paradigm, we measured brain activation with functional magnetic resonance imaging in response to covert reinstatement of a cognitive context, prior to presenting an item memory probe. Contexts were studied with multiple items to ensure that spontaneous item retrieval would not occur prior to the test probe. At test, contexts were reinstated for 8 s before the test probe was presented. We manipulated whether the reinstated context matched the encoding context of the test probe that followed. For such matching contexts, we found that increased PHc activation prior to the test probe predicted recollection following the test probe. If a context unrelated to the eventual test item probe was reinstated, there was no such association between PHc activation during context reinstatement and eventual memory judgments. These findings suggest that PHc activation is correlated with cognitive context retrieval.

Associative memory in aging: the effect of unitization on source memory.

In normal aging, memory for associations declines more than memory for individual items. Unitization is an encoding process defined by creation of a new single entity to represent a new arbitrary association. The current study tested the hypothesis that age-related differences in associative memory can be reduced by encoding instructions that promote unitization. In two experiments, groups of 20 young and 20 older participants learned new associations between a word and a background color under two conditions. In the item detail condition, they had to imagine that the item is the same color as the background-an instruction promoting unitization of the associations. In the context detail condition, which did not promote unitization, they had to imagine that the item interacted with another colored object. At test, they had to retrieve the color that was associated with each word (source memory). In both experiments, the results showed an age-related decrement in source memory performance in the context detail but not in the item detail condition. Moreover, Experiment 2 examined receiver operating characteristics in older participants and indicated that familiarity contributed more to source memory performance in the item detail than in the context detail condition. These findings suggest that unitization of new associations can overcome the associative memory deficit observed in aging, at least for item-color associations.

Adaptation to cognitive context and item information in the medial temporal lobes.

The medial temporal lobes (MTL) play an essential role in episodic memory, and accumulating evidence indicates that two MTL subregions--the perirhinal (PRc) and parahippocampal (PHc) cortices--might have different functions. According to the binding of item and context theory (Diana, Yonelinas, & Ranganath, 2007; Eichenbaum, Yonelinas, & Ranganath, 2007), PRc is involved in processing item information, the target of memory encoding, whereas PHc is involved in processing context information, peripheral information that identifies the circumstances of the episode. Here, we used functional magnetic resonance imaging (fMRI) adaptation to test the roles of different MTL subregions in the processing of item and context information. Participants were scanned while viewing a series of objects. Each object was presented with a unique semantic encoding question that elicited a salient cognitive context. The object picture, the encoding question, both, or neither were immediately repeated. We found that PRc activity was sensitive to repetition of the object but not the encoding question whereas PHc activity was sensitive to repetition of the encoding question but not the object. These data are consistent with the idea that the PRc and PHc are differentially involved in the representation of item and context information and additionally suggest that the role of the PHc extends to nonspatial, cognitive context information.

ERP correlates of source memory: unitized source information increases familiarity-based retrieval.

Source memory tests typically require subjects to make decisions about the context in which an item was encoded and are thought to depend on recollection of details from the study episode. Although it is generally believed that familiarity does not contribute to source memory, recent behavioral studies have suggested that familiarity may also support source recognition when item and source information are integrated, or "unitized," during study (Diana, Yonelinas, and Ranganath, 2008). However, an alternative explanation of these behavioral findings is that unitization affects the manner in which recollection contributes to performance, rather than increasing familiarity-based source memory. To discriminate between these possibilities, we conducted an event-related potential (ERP) study testing the hypothesis that unitization increases the contribution of familiarity to source recognition. Participants studied associations between words and background colors using tasks that either encouraged or discouraged unitization. ERPs were recorded during a source memory test for background color. The results revealed two distinct neural correlates of source recognition: a frontally distributed positivity that was associated with familiarity-based source memory in the high-unitization condition only and a parietally distributed positivity that was associated with recollection-based source memory in both the high- and low-unitization conditions. The ERP and behavioral findings provide converging evidence for the idea that familiarity can contribute to source recognition, particularly when source information is encoded as an item detail.

Medial temporal lobe activity during source retrieval reflects information type, not memory strength.

The medial temporal lobes (MTLs) are critical for episodic memory but the functions of MTL subregions are controversial. According to memory strength theory, MTL subregions collectively support declarative memory in a graded manner. In contrast, other theories assert that MTL subregions support functionally distinct processes. For instance, one view is that perirhinal cortex (PRc) processes item information, parahippocampal cortex (PHc) processes context information, and the hippocampus binds item and context. Here, we report two experiments that tested competing predictions from these models. In these studies, subjects encoded color-word associations by imagining color either as a contextual association (context detail condition) or as a feature of the item to be encoded (item detail condition). Results showed that encoding color information as an item detail improved source recognition in amnesic patients with recollection deficits. Furthermore, event-related fMRI data from healthy subjects revealed PRc activation associated with successful retrieval of item details, whereas activation in the hippocampus and PHc was associated with recollection-based source retrieval. The qualitatively different patterns of results observed in PRc and hippocampus/PHc are inconsistent with a memory strength account and are consistent with the idea that different MTL regions process different types of episodic information.