α-Band activity tracks a two-dimensional spotlight of attention during spatial working memory maintenance.

Covert spatial attention is thought to facilitate the maintenance of locations in working memory, and EEG α-band activity (8-12 Hz) is proposed to track the focus of covert attention. Recent work has shown that multivariate patterns of α-band activity track the polar angle of remembered locations relative to fixation. However, a defining feature of covert spatial attention is that it facilitates processing in a specific region of the visual field, and prior work has not determined whether patterns of α-band activity track the two-dimensional (2-D) coordinates of remembered stimuli within a visual hemifield or are instead maximally sensitive to the polar angle of remembered locations around fixation. Here, we used a lateralized spatial estimation task, in which observers remembered the location of one or two target dots presented to one side of fixation, to test this question. By applying a linear discriminant classifier to the topography of α-band activity, we found that we were able to decode the location of remembered stimuli. Critically, model comparison revealed that the pattern of classifier choices observed across remembered positions was best explained by a model assuming that α-band activity tracks the 2-D coordinates of remembered locations rather than a model assuming that α-band activity tracks the polar angle of remembered locations relative to fixation. These results support the hypothesis that this α-band activity is involved in the spotlight of attention, and arises from mid- to lower-level visual areas involved in maintaining spatial locations in working memory.NEW & NOTEWORTHY A substantial body of work has shown that patterns of EEG α-band activity track the angular coordinates of attended and remembered stimuli around fixation, but whether these patterns track the two-dimensional coordinates of stimuli presented within a visual hemifield remains an open question. Here, we demonstrate that α-band activity tracks the two-dimensional coordinates of remembered stimuli within a hemifield, showing that α-band activity reflects a spotlight of attention focused on locations maintained in working memory.

Decoding selective attention to context memory: An aging study.

Emerging evidence has suggested that the tendency for older adults to bind too much contextual information during encoding (i.e., hyper-binding) may contribute to poorer memory for relevant contextual information during retrieval. While these findings are consistent with theories of age-related declines in selective attention and inhibitory control, the degree to which older adults are able to selectively attend to relevant contextual information during encoding is unknown. To better understand the neural dynamics associated with selective attention during encoding, the current study applied multivariate pattern analyses (MVPA) to oscillatory EEG in order to track moment-to-moment shifts of attention between relevant and irrelevant contextual information during encoding. Young and older adults studied pictures of objects in the presence of two contextual features: a color and a scene, and their attention was directed to the object's relationship with one of those contexts (i.e., target context). Results showed that patterns of oscillatory power successfully predicted whether selective attention was directed to a scene or color, across age groups. Individual differences in overall classification performance were associated with individual differences in target context memory accuracy during retrieval. However, changes in classification performance within a trial, suggestive of fluctuations in selective attention, predicted individual differences in hyper-binding. To the best of our knowledge, this is the first study to use MPVA techniques to decode attention during episodic encoding and the impact of attentional shifts toward distracting information on age-related context memory impairments and hyper-binding. These results are consistent with the as-of-yet unsubstantiated theory that age-related declines in context memory may be attributable to poorer selective attention and/or greater inhibitory deficits in older adults.

Beta-band activity represents the recent past during episodic encoding.

While much research has focused on understanding how individual stimuli are encoded in episodic memory, less is known about how a series of events is bound into a coherent episode. Cognitive models of episodic memory propose that information about presented stimuli is integrated into a composite representation reflecting one's past experience, allowing events separated in time to become associated. Recent evidence suggests that neural oscillatory activity may be critically involved in this process. To examine how oscillatory activity contributes to binding of information across events, we measured scalp EEG as participants studied categorized lists of people, places, and objects. We assessed their memory for the lists using free recall, allowing us to characterize the temporal and semantic organization of the studied items in memory. Using pattern classification, we identified EEG activity during encoding at a range of frequencies and scalp locations that was sensitive to the category of presented stimuli. In the beta band (16-25Hz) at right posterior electrodes, we observed activity that was also sensitive to the category of recently presented stimuli. This neural activity showed two characteristics consistent with a representation of the recent past: It became stronger when multiple items from the same category were presented in succession, and it contained a fading trace of the previous category after a category shift. When items were separated by an inter-item distraction task, this integrative beta-band activity was disrupted. Distraction also led to decreased semantic organization of the studied materials without affecting their temporal organization; this suggests that distraction disrupts the integration of semantic information over time, preventing encoding of items in terms of the semantic context of earlier items. Our results provide evidence that beta-band activity is involved in maintaining information about recent events, allowing construction of a coherent representation of a temporally extended episode in memory.

Neural activity in the medial temporal lobe reveals the fidelity of mental time travel.

Neural circuitry in the medial temporal lobe (MTL) is critically involved in mental time travel, which involves the vivid retrieval of the details of past experience. Neuroscientific theories propose that the MTL supports memory of the past by retrieving previously encoded episodic information, as well as by reactivating a temporal code specifying the position of a particular event within an episode. However, the neural computations supporting these abilities are underspecified. To test hypotheses regarding the computational mechanisms supported by different MTL subregions during mental time travel, we developed a computational model that linked a blood oxygenation level-dependent signal to cognitive operations, allowing us to predict human performance in a memory search task. Activity in the posterior MTL, including parahippocampal cortex, reflected how strongly one reactivates the temporal context of a retrieved memory, allowing the model to predict whether the next memory will correspond to a nearby moment in the study episode. A signal in the anterior MTL, including perirhinal cortex, indicated the successful retrieval of list items, without providing information regarding temporal organization. A hippocampal signal reflected both processes, consistent with theories that this region binds item and context information together to form episodic memories. These findings provide evidence for modern theories that describe complementary roles of the hippocampus and surrounding parahippocampal and perirhinal cortices during the retrieval of episodic memories, shaping how humans revisit the past.

Decoding Episodic Retrieval Processes: Frontoparietal and Medial Temporal Lobe Contributions to Free Recall.

Neuroimaging studies of recognition memory have identified distinct patterns of cortical activity associated with two sets of cognitive processes: Recollective processes supporting retrieval of information specifying a probe item's original source are associated with the posterior hippocampus, ventral posterior parietal cortex, and medial pFC. Familiarity processes supporting the correct identification of previously studied probes (in the absence of a recollective response) are associated with activity in anterior medial temporal lobe (MTL) structures including the perirhinal cortex and anterior hippocampus, in addition to lateral prefrontal and dorsal posterior parietal cortex. Here, we address an open question in the cognitive neuroscientific literature: To what extent are these same neurocognitive processes engaged during an internally directed memory search task like free recall? We recorded fMRI activity while participants performed a series of free recall and source recognition trials, and we used a combination of univariate and multivariate analysis techniques to compare neural activation profiles across the two tasks. Univariate analyses showed that posterior MTL regions were commonly associated with recollective processes during source recognition and with free recall responses. Prefrontal and posterior parietal regions were commonly associated with familiarity processes and free recall responses, whereas anterior MTL regions were only associated with familiarity processes during recognition. In contrast with the univariate results, free recall activity patterns characterized using multivariate pattern analysis did not reliably match the neural patterns associated with recollective processes. However, these free recall patterns did reliably match patterns associated with familiarity processes, supporting theories of memory in which common cognitive mechanisms support both item recognition and free recall.

Functional interactions between large-scale networks during memory search.

Neuroimaging studies have identified two major large-scale brain networks, the default mode network (DMN) and the dorsal attention network (DAN), which are engaged for internally and externally directed cognitive tasks respectively, and which show anticorrelated activity during cognitively demanding tests and at rest. We identified these brain networks using independent component analysis (ICA) of functional magnetic resonance imaging data, and examined their interactions during the free-recall task, a self-initiated memory search task in which retrieval is performed in the absence of external cues. Despite the internally directed nature of the task, the DAN showed transient engagement in the seconds leading up to successful retrieval. ICA revealed a fractionation of the DMN into 3 components. A posteromedial network increased engagement during memory search, while the two others showed suppressed activity during memory search. Cooperative interactions between this posteromedial network, a right-lateralized frontoparietal control network, and a medial prefrontal network were maintained during memory search. The DAN demonstrated heterogeneous task-dependent shifts in functional coupling with various subnetworks within the DMN. This functional reorganization suggests a broader role of the DAN in the absence of externally directed cognition, and highlights the contribution of the posteromedial network to episodic retrieval.

Temporal context and the organisational impairment of memory search in schizophrenia.

INTRODUCTION: An influential theory of schizophrenic deficits in executive function suggests that patients have difficulty maintaining and utilising an internal contextual representation, whose function is to ensure that stimuli are processed in a task-appropriate manner. In basic research on episodic memory, retrieved-context theories propose that an internal contextual representation is critically involved in memory search, facilitating the retrieval of task-appropriate memories. This contextual machinery is thought to give rise to temporal organisation during free recall: the tendency for successive recall responses to correspond to items from nearby positions on the study list. If patients with schizophrenia have a generalised contextual deficit, then this leads to the prediction that these patients will exhibit reduced temporal organisation in free recall. METHODS: Using a combination of classic and recently developed organisational measures, we characterised recall organisation in 75 patients with schizophrenia and 72 nondisordered control participants performing a multi-trial free-recall task. RESULTS: Patients with schizophrenia showed diminished temporal organisation, as well as diminished subjective organisation of their recall sequences relative to control participants. The two groups showed similar amounts of semantic organisation during recall. CONCLUSIONS: The observation of reduced temporal organisation in the patient group is consistent with the proposal that the memory deficit in schizophrenia can be characterised as a deficit in contextual processing.

Brain rhythms in mental time travel.

The memory theorist Endel Tulving referred to the ability to search through one's memories, and revisit events and episodes from one's past, as mental time travel. This process involves the reactivation of past mental states reflecting the perceptual and conceptual characteristics of the original experience. Widely distributed neural circuitry is engaged in the service of memory search, and the dynamics of these circuits are reflected in rhythmic oscillatory signals at widespread frequencies, recorded both in the local field around neurons and more globally at the scalp. Retrieved-context theory provides a theoretical bridge between the behavioral phenomena exhibited by participants in memory search tasks, and the neural signals reflecting the dynamics of the underlying circuitry. Computational models based on this theory make broad predictions regarding the representational structure of neural activity recorded during these tasks. In recent work, researchers have used multivariate analytic techniques on topographic patterns of oscillatory neural activity to confirm critical predictions of retrieved-context theory. We review the cognitive theory motivating this recent work, and the analytic techniques being developed to create integrated neural-behavioral models of human memory search.

Category-specific neural oscillations predict recall organization during memory search.

Retrieved-context models of human memory propose that as material is studied, retrieval cues are constructed that allow one to target particular aspects of past experience. We examined the neural predictions of these models by using electrocorticographic/depth recordings and scalp electroencephalography (EEG) to characterize category-specific oscillatory activity, while participants studied and recalled items from distinct, neurally discriminable categories. During study, these category-specific patterns predict whether a studied item will be recalled. In the scalp EEG experiment, category-specific activity during study also predicts whether a given item will be recalled adjacent to other same-category items, consistent with the proposal that a category-specific retrieval cue is used to guide memory search. Retrieved-context models suggest that integrative neural circuitry is involved in the construction and maintenance of the retrieval cue. Consistent with this hypothesis, we observe category-specific patterns that rise in strength as multiple same-category items are studied sequentially, and find that individual differences in this category-specific neural integration during study predict the degree to which a participant will use category information to organize memory search. Finally, we track the deployment of this retrieval cue during memory search: Category-specific patterns are stronger when participants organize their responses according to the category of the studied material.

Oscillatory patterns in temporal lobe reveal context reinstatement during memory search.

Psychological theories of memory posit that when people recall a past event, they not only recover the features of the event itself, but also recover information associated with other events that occurred nearby in time. The events surrounding a target event, and the thoughts they evoke, may be considered to represent a context for the target event, helping to distinguish that event from similar events experienced at different times. The ability to reinstate this contextual information during memory search has been considered a hallmark of episodic, or event-based, memory. We sought to determine whether context reinstatement may be observed in electrical signals recorded from the human brain during episodic recall. Analyzing electrocorticographic recordings taken as 69 neurosurgical patients studied and recalled lists of words, we uncovered a neural signature of context reinstatement. Upon recalling a studied item, we found that the recorded patterns of brain activity were not only similar to the patterns observed when the item was studied, but were also similar to the patterns observed during study of neighboring list items, with similarity decreasing reliably with positional distance. The degree to which individual patients displayed this neural signature of context reinstatement was correlated with their tendency to recall neighboring list items successively. These effects were particularly strong in temporal lobe recordings. Our findings show that recalling a past event evokes a neural signature of the temporal context in which the event occurred, thus pointing to a neural basis for episodic memory.

Items Outperform Adjectives in a Computational Model of Binary Semantic Classification.

Semantic memory encompasses one's knowledge about the world. Distributional semantic models, which construct vector spaces with embedded words, are a proposed framework for understanding the representational structure of human semantic knowledge. Unlike some classic semantic models, distributional semantic models lack a mechanism for specifying the properties of concepts, which raises questions regarding their utility for a general theory of semantic knowledge. Here, we develop a computational model of a binary semantic classification task, in which participants judged target words for the referent's size or animacy. We created a family of models, evaluating multiple distributional semantic models, and mechanisms for performing the classification. The most successful model constructed two composite representations for each extreme of the decision axis (e.g., one averaging together representations of characteristically big things and another of characteristically small things). Next, the target item was compared to each composite representation, allowing the model to classify more than 1,500 words with human-range performance and to predict response times. We propose that when making a decision on a binary semantic classification task, humans use task prompts to retrieve instances representative of the extremes on that semantic dimension and compare the probe to those instances. This proposal is consistent with the principles of the instance theory of semantic memory.

The modulation and elimination of temporal organization in free recall.

Experiences occur in a continual succession, and the temporal structure of those experiences is often preserved in memory. The temporal contiguity effect of free recall reveals the temporal structure of memory: when a particular item is remembered, the next response is likely to come from a nearby list position. This effect is remarkably robust, appearing across a wide variety of methodological variations of the task. The temporal contiguity effect is also central to retrieved-context models, which propose temporal organization arises from the interaction of a temporal context representation with the contents of memory. Across six experiments, we demonstrate methodological manipulations that dramatically modulate and even eliminate temporal organization in free recall. We find that temporal organization is strongly modulated and in some cases potentially eliminated by strong semantic structure, the presence of retrieval practice, and a long list length. Other factors such as orienting task, paired-associate item structure, and retention interval duration have more subtle effects on temporal organization. In an accompanying set of simulations, we show that the modulation and elimination of the temporal organization follows lawful patterns predicted by the context maintenance and retrieval (CMR) retrieved-context model. We also find cases where CMR does not specifically predict the modulation of temporal organization, and in these cases our analysis suggests how the theory might be developed to account for these effects. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Children's memory "in the wild": examining the temporal organization of free recall from a week-long camp at a local zoo.

Free-recall paradigms have greatly influenced our understanding of memory. The majority of this research involves laboratory-based events (e.g., word lists) that are studied and tested within minutes. This literature shows that adults recall events in a temporally organized way, with successive responses often coming from neighboring list positions (i.e., temporal clustering) and with enhanced memorability of items from the end of a list (i.e., recency). Temporal clustering effects are so robust that temporal organization is described as a fundamental memory property. Yet relatively little is known about the development of this temporal structure across childhood, and even less about children's memory search for real-world events occurring over an extended period. In the present work, children (N = 144; 3 age groups: 4-5-year-olds, 6-7-year-olds, 8-10-year-olds) took part in a 5-day summer camp at a local zoo. The camp involved various dynamic events, including daily animal exhibit visits. On day 5, children were asked to recall all the animals they visited. We found that overall recall performance, in terms of number of animals recalled, improved steadily across childhood. Temporal organization and recency effects showed different developmental patterns. Temporal clustering was evident in the response sequences for all age groups and became progressively stronger across childhood. In contrast, the recency advantage, when characterized as a proportion of total responses, was stable across age groups. Thus, recall dynamics in early childhood parallel that seen in adulthood, with continued development of temporal organization across middle to late childhood.

A context maintenance and retrieval model of organizational processes in free recall.

The authors present the context maintenance and retrieval (CMR) model of memory search, a generalized version of the temporal context model of M. W. Howard and M. J. Kahana (2002a), which proposes that memory search is driven by an internally maintained context representation composed of stimulus-related and source-related features. In the CMR model, organizational effects (the tendency for related items to cluster during the recall sequence) arise as a consequence of associations between active context elements and features of the studied material. Semantic clustering is due to longstanding context-to-item associations, whereas temporal clustering and source clustering are both due to associations formed during the study episode. A behavioral investigation of the three forms of organization provides data to constrain the CMR model, revealing interactions between the organizational factors. Finally, the authors discuss the implications of CMR for their understanding of a broad class of episodic memory phenomena and suggest ways in which this theory may guide exploration of the neural correlates of memory search.

Memory search and the neural representation of context.

A challenge for theories of episodic memory is to determine how we focus memory search on a set of recently learned items. Cognitive theories suggest that the recall of an item representation is driven by an internally maintained context representation that integrates incoming information with a long time-scale. Neural investigations have shown that recalling an item revives the pattern of brain activity present during its study. To link these neural and cognitive approaches, we propose a framework in which context is maintained and updated in prefrontal cortex, and is associated with item information through hippocampal projections. The proposed framework is broadly consistent with neurobiological studies of temporal integration and with studies of memory deficits in individuals with prefrontal damage.

Examining the episodic context account: does retrieval practice enhance memory for context?

Retrieval practice, such as filling in blanks or taking quizzes, is firmly established as an effective study strategy. However, the underlying mechanism of how retrieval practice benefits memory is still unclear. One current theory, the episodic context account, proposes that retrieval enhances memory by reinstating a prior learning context. This retrieved context is then strengthened and updated to include context at the time of recall, which later serves as an effective retrieval cue. However, few studies have directly tested this hypothesis. We did so by examining participants' memory for the initial study context. Across three experiments, participants encoded cue-target pairs presented in different colors and either restudied or practiced retrieving the targets. If retrieval practice benefits memory by reinstating the prior episodic context, participants who successfully retrieved the items during practice should have enhanced memory for context details (i.e. font color) compared to participants who restudied the pairs. Contrary to this prediction, memory for font colors did not differ between the restudy condition and the retrieval practice condition. Even when font color was actively attended to and integrated with the to-be-remembered items, retrieval practice did not increase memory for this aspect of context. Our results suggest that the context reinstated during retrieval practice is limited in nature. Aspects of the context that are not essential to retrieval of the item are not strengthened by retrieval practice.

Searching for Semantic Knowledge: A Vector Space Semantic Analysis of the Feature Generation Task.

A recent neuropsychological study found that amnesic patients with hippocampal damage (HP) and severe declarative memory impairment produce markedly fewer responses than healthy comparison (CO) participants in a semantic feature generation task (Klooster and Duff, 2015), consistent with the idea that hippocampal damage is associated with semantic cognitive deficits. Participants were presented with a target word and asked to produce as many features of that word as possible (e.g., for target word "book," "read words on a page"). Here, we use the response sequences collected by Klooster and Duff (2015) to develop a vector space model of semantic search. We use this model to characterize the dynamics of semantic feature generation and consider the role of the hippocampus in this search process. Both HP and CO groups tended to initiate the search process with features close in semantic space to the target word, with a gradual decline in similarity to the target word over the first several responses. Adjacent features in the response sequence showed stronger similarity to each other than to non-adjacent features, suggesting that the search process follows a local trajectory in semantic space. Overall, HP patients generated features that were closer in semantic space to the representation of the target word, as compared to the features generated by the CO group, which ranged more widely in semantic space. These results are consistent with a model in which a compound retrieval cue (containing a representation of the target word and a representation of the previous response) is used to probe semantic memory. The model suggests that the HP group's search process is restricted from ranging as far in semantic space from the target word, relative to the CO group. These results place strong constraints on the structure of models of semantic memory search, and on the role of hippocampus in probing semantic memory.

Beyond mind-reading: multi-voxel pattern analysis of fMRI data.

A key challenge for cognitive neuroscience is determining how mental representations map onto patterns of neural activity. Recently, researchers have started to address this question by applying sophisticated pattern-classification algorithms to distributed (multi-voxel) patterns of functional MRI data, with the goal of decoding the information that is represented in the subject's brain at a particular point in time. This multi-voxel pattern analysis (MVPA) approach has led to several impressive feats of mind reading. More importantly, MVPA methods constitute a useful new tool for advancing our understanding of neural information processing. We review how researchers are using MVPA methods to characterize neural coding and information processing in domains ranging from visual perception to memory search.

Lingering representations of stimuli influence recall organization.

Several prominent theories posit that information about recent experiences lingers in the brain and organizes memories for current experiences, by forming a temporal context that is linked to those memories at encoding. According to these theories, if the thoughts preceding an experience X resemble the thoughts preceding an experience Y, then X and Y should show an elevated probability of being recalled together. We tested this prediction by using multi-voxel pattern analysis (MVPA) of fMRI data to measure neural evidence for lingering processing of preceding stimuli. As predicted, memories encoded with similar lingering thoughts about the category of preceding stimuli were more likely to be recalled together. Our results demonstrate that the "fading embers" of previous stimuli help to organize recall, confirming a key prediction of computational models of episodic memory.

A predictive framework for evaluating models of semantic organization in free recall.

Research in free recall has demonstrated that semantic associations reliably influence the organization of search through episodic memory. However, the specific structure of these associations and the mechanisms by which they influence memory search remain unclear. We introduce a likelihood-based model-comparison technique, which embeds a model of semantic structure within the context maintenance and retrieval (CMR) model of human memory search. Within this framework, model variants are evaluated in terms of their ability to predict the specific sequence in which items are recalled. We compare three models of semantic structure, latent semantic analysis (LSA), global vectors (GloVe), and word association spaces (WAS), and find that models using WAS have the greatest predictive power. Furthermore, we find evidence that semantic and temporal organization is driven by distinct item and context cues, rather than a single context cue. This finding provides important constraint for theories of memory search.