Neurocognitive subprocesses of working memory performance.

Working memory (WM) has been defined as the active maintenance and flexible updating of goal-relevant information in a form that has limited capacity and resists interference. Complex measures of WM recruit multiple subprocesses, making it difficult to isolate specific contributions of putatively independent subsystems. The present study was designed to determine whether neurophysiological indicators of proposed subprocesses of WM predict WM performance. We recruited 200 individuals defined by care-seeking status and measured neural responses using electroencephalography (EEG), while participants performed four WM tasks. We extracted spectral and time-domain EEG features from each task to quantify each of the hypothesized WM subprocesses: maintenance (storage of content), goal maintenance, and updating. We then used EEG measures of each subprocess as predictors of task performance to evaluate their contribution to WM. Significant predictors of WM capacity included contralateral delay activity and frontal theta, features typically associated with maintenance (storage of content) processes. In contrast, significant predictors of reaction time and its variability included contingent negative variation and the P3b, features typically associated with goal maintenance and updating. Broadly, these results suggest two principal dimensions that contribute to WM performance, tonic processes during maintenance contributing to capacity, and phasic processes during stimulus processing that contribute to response speed and variability. The analyses additionally highlight that reliability of features across tasks was greater (and comparable to that of WM performance) for features associated with stimulus processing (P3b and alpha), than with maintenance (gamma, theta and cross-frequency coupling).

Multivariate resting-state functional connectivity predicts response to cognitive behavioral therapy in obsessive-compulsive disorder.

Cognitive behavioral therapy (CBT) is an effective treatment for many with obsessive-compulsive disorder (OCD). However, response varies considerably among individuals. Attaining a means to predict an individual's potential response would permit clinicians to more prudently allocate resources for this often stressful and time-consuming treatment. We collected resting-state functional magnetic resonance imaging from adults with OCD before and after 4 weeks of intensive daily CBT. We leveraged machine learning with cross-validation to assess the power of functional connectivity (FC) patterns to predict individual posttreatment OCD symptom severity. Pretreatment FC patterns within the default mode network and visual network significantly predicted posttreatment OCD severity, explaining up to 67% of the variance. These networks were stronger predictors than pretreatment clinical scores. Results have clinical implications for developing personalized medicine approaches to identifying individual OCD patients who will maximally benefit from intensive CBT.

Anodal Transcranial Direct Current Stimulation to the Left Rostrolateral Prefrontal Cortex Selectively Improves Source Memory Retrieval.

Functional neuroimaging studies have consistently implicated the left rostrolateral prefrontal cortex (RLPFC) as playing a crucial role in the cognitive operations supporting episodic memory and analogical reasoning. However, the degree to which the left RLPFC causally contributes to these processes remains underspecified. We aimed to assess whether targeted anodal stimulation-thought to boost cortical excitability-of the left RLPFC with transcranial direct current stimulation (tDCS) would lead to augmentation of episodic memory retrieval and analogical reasoning task performance in comparison to cathodal stimulation or sham stimulation. Seventy-two healthy adult participants were evenly divided into three experimental groups. All participants performed a memory encoding task on Day 1, and then on Day 2, they performed continuously alternating tasks of episodic memory retrieval, analogical reasoning, and visuospatial perception across two consecutive 30-min experimental sessions. All groups received sham stimulation for the first experimental session, but the groups differed in the stimulation delivered to the left RLPFC during the second session (either sham, 1.5 mA anodal tDCS, or 1.5 mA cathodal tDCS). The experimental group that received anodal tDCS to the left RLPFC during the second session demonstrated significantly improved episodic memory source retrieval performance, relative to both their first session performance and relative to performance changes observed in the other two experimental groups. Performance on the analogical reasoning and visuospatial perception tasks did not exhibit reliable changes as a result of tDCS. As such, our results demonstrate that anodal tDCS to the left RLPFC leads to a selective and robust improvement in episodic source memory retrieval.

Episodic Memory Retrieval Benefits from a Less Modular Brain Network Organization.

Most complex cognitive tasks require the coordinated interplay of multiple brain networks, but the act of retrieving an episodic memory may place especially heavy demands for communication between the frontoparietal control network (FPCN) and the default mode network (DMN), two networks that do not strongly interact with one another in many task contexts. We applied graph theoretical analysis to task-related fMRI functional connectivity data from 20 human participants and found that global brain modularity-a measure of network segregation-is markedly reduced during episodic memory retrieval relative to closely matched analogical reasoning and visuospatial perception tasks. Individual differences in modularity were correlated with memory task performance, such that lower modularity levels were associated with a lower false alarm rate. Moreover, the FPCN and DMN showed significantly elevated coupling with each other during the memory task, which correlated with the global reduction in brain modularity. Both networks also strengthened their functional connectivity with the hippocampus during the memory task. Together, these results provide a novel demonstration that reduced modularity is conducive to effective episodic retrieval, which requires close collaboration between goal-directed control processes supported by the FPCN and internally oriented self-referential processing supported by the DMN.SIGNIFICANCE STATEMENT Modularity, an index of the degree to which nodes of a complex system are organized into discrete communities, has emerged as an important construct in the characterization of brain connectivity dynamics. We provide novel evidence that the modularity of the human brain is reduced when individuals engage in episodic memory retrieval, relative to other cognitive tasks, and that this state of lower modularity is associated with improved memory performance. We propose a neural systems mechanism for this finding where the nodes of the frontoparietal control network and default mode network strengthen their interaction with one another during episodic retrieval. Such across-network communication likely facilitates effective access to internally generated representations of past event knowledge.

Multi-voxel pattern classification differentiates personally experienced event memories from secondhand event knowledge.

Studies of autobiographical memory retrieval often use photographs to probe participants' memories for past events. Recent neuroimaging work has shown that viewing photographs depicting events from one's own life evokes a characteristic pattern of brain activity across a network of frontal, parietal, and medial temporal lobe regions that can be readily distinguished from brain activity associated with viewing photographs from someone else's life (Rissman, Chow, Reggente, and Wagner, 2016). However, it is unclear whether the neural signatures associated with remembering a personally experienced event are distinct from those associated with recognizing previously encountered photographs of an event. The present experiment used a novel functional magnetic resonance imaging (fMRI) paradigm to investigate putative differences in brain activity patterns associated with these distinct expressions of memory retrieval. Eighteen participants wore necklace-mounted digital cameras to capture events from their everyday lives over the course of three weeks. One week later, participants underwent fMRI scanning, where on each trial they viewed a sequence of photographs depicting either an event from their own life or from another participant's life and judged their memory for this event. Importantly, half of the trials featured photographic sequences that had been shown to participants during a laboratory session administered the previous day. Multi-voxel pattern analyses assessed the sensitivity of two brain networks of interest-as identified by a meta-analysis of prior autobiographical and laboratory-based memory retrieval studies-to the original source of the photographs (own life or other's life) and their experiential history as stimuli (previewed or non-previewed). The classification analyses revealed a striking dissociation: activity patterns within the autobiographical memory network were significantly more diagnostic than those within the laboratory-based network as to whether photographs depicted one's own personal experience (regardless of whether they had been previously seen), whereas activity patterns within the laboratory-based memory network were significantly more diagnostic than those within the autobiographical memory network as to whether photographs had been previewed (regardless of whether they were from the participant's own life). These results, also apparent in whole-brain searchlight classifications, provide evidence for dissociable patterns of activation across two putative memory networks as a function of whether real-world photographs trigger the retrieval of firsthand experiences or secondhand event knowledge.

Dissociable profiles of generalization/discrimination in the human hippocampus during associative retrieval.

When encountering stimuli that vary slightly from previous experiences, neural signals within the CA3 and dentate gyrus (CA3 DG) hippocampal subfields are thought to facilitate mnemonic discrimination, whereas CA1 may be less sensitive to minor stimulus changes, allowing for generalization across similar events. Studies have also posited a critical role for CA1 in the comparison of events to memory-derived expectations, but the degree to which these processes are impacted by explicit retrieval demands is yet unclear. To evaluate extant accounts of hippocampal subfield function, we acquired high-resolution fMRI data as participants performed a task in which famous names were used to cue the retrieval of previously paired images. Although both left CA3 DG and CA1 showed match enhancement effects, responding more to original paired images (targets) than to never-before-seen images (novels), the sensitivity of these subfields to stimulus changes and task demands diverged. CA3 DG showed a goal-independent, yet highly specific, preference for previously encountered stimuli, responding equally strongly to targets and mispaired associates, while showing equally weak responses to close lures and novels. In contrast, recognition signals in CA1 were goal-dependent (i.e., not evoked by mispaired associates), yet accommodating of subtle stimulus differences, such that close lures evoked comparable activity as targets. © 2016 Wiley Periodicals, Inc.

Goal-Directed Modulation of Neural Memory Patterns: Implications for fMRI-Based Memory Detection.

Remembering a past event elicits distributed neural patterns that can be distinguished from patterns elicited when encountering novel information. These differing patterns can be decoded with relatively high diagnostic accuracy for individual memories using multivoxel pattern analysis (MVPA) of fMRI data. Brain-based memory detection--if valid and reliable--would have clear utility beyond the domain of cognitive neuroscience, in the realm of law, marketing, and beyond. However, a significant boundary condition on memory decoding validity may be the deployment of "countermeasures": strategies used to mask memory signals. Here we tested the vulnerability of fMRI-based memory detection to countermeasures, using a paradigm that bears resemblance to eyewitness identification. Participants were scanned while performing two tasks on previously studied and novel faces: (1) a standard recognition memory task; and (2) a task wherein they attempted to conceal their true memory state. Univariate analyses revealed that participants were able to strategically modulate neural responses, averaged across trials, in regions implicated in memory retrieval, including the hippocampus and angular gyrus. Moreover, regions associated with goal-directed shifts of attention and thought substitution supported memory concealment, and those associated with memory generation supported novelty concealment. Critically, whereas MVPA enabled reliable classification of memory states when participants reported memory truthfully, the ability to decode memory on individual trials was compromised, even reversing, during attempts to conceal memory. Together, these findings demonstrate that strategic goal states can be deployed to mask memory-related neural patterns and foil memory decoding technology, placing a significant boundary condition on their real-world utility.

Decoding fMRI Signatures of Real-world Autobiographical Memory Retrieval.

Extant neuroimaging data implicate frontoparietal and medial-temporal lobe regions in episodic retrieval, and the specific pattern of activity within and across these regions is diagnostic of an individual's subjective mnemonic experience. For example, in laboratory-based paradigms, memories for recently encoded faces can be accurately decoded from single-trial fMRI patterns [Uncapher, M. R., Boyd-Meredith, J. T., Chow, T. E., Rissman, J., & Wagner, A. D. Goal-directed modulation of neural memory patterns: Implications for fMRI-based memory detection. Journal of Neuroscience, 35, 8531-8545, 2015; Rissman, J., Greely, H. T., & Wagner, A. D. Detecting individual memories through the neural decoding of memory states and past experience. Proceedings of the National Academy of Sciences, U.S.A., 107, 9849-9854, 2010]. Here, we investigated the neural patterns underlying memory for real-world autobiographical events, probed at 1- to 3-week retention intervals as well as whether distinct patterns are associated with different subjective memory states. For 3 weeks, participants (n = 16) wore digital cameras that captured photographs of their daily activities. One week later, they were scanned while making memory judgments about sequences of photos depicting events from their own lives or events captured by the cameras of others. Whole-brain multivoxel pattern analysis achieved near-perfect accuracy at distinguishing correctly recognized events from correctly rejected novel events, and decoding performance did not significantly vary with retention interval. Multivoxel pattern classifiers also differentiated recollection from familiarity and reliably decoded the subjective strength of recollection, of familiarity, or of novelty. Classification-based brain maps revealed dissociable neural signatures of these mnemonic states, with activity patterns in hippocampus, medial PFC, and ventral parietal cortex being particularly diagnostic of recollection. Finally, a classifier trained on previously acquired laboratory-based memory data achieved reliable decoding of autobiographical memory states. We discuss the implications for neuroscientific accounts of episodic retrieval and comment on the potential forensic use of fMRI for probing experiential knowledge.

Effects of aging on value-directed modulation of semantic network activity during verbal learning.

While impairments in memory recall are apparent in aging, older adults show a remarkably preserved ability to selectively remember information deemed valuable. Here, we use fMRI to compare brain activation in healthy older and younger adults during encoding of high and low value words to determine whether there are differences in how older adults achieve value-directed memory selectivity. We find that memory selectivity in older adults is associated with value-related changes in activation during word presentation in left hemisphere regions that are involved in semantic processing, similar to young adults. However, highly selective young adults show a relatively greater increase in semantic network activity during encoding of high-value items, whereas highly selective older adults show relatively diminished activity during encoding of low-value items. Additionally, only younger adults showed value-related increases in activity in semantic and reward processing regions during presentation of the value cue preceding each to-be-remembered word. Young adults therefore respond to cue value more proactively than do older adults, yet the magnitude of value-related differences in cue period brain activity did not predict individual differences in memory selectivity. Thus, our data also show that age-related reductions in prestimulus activity do not always lead to inefficient performance.

Shared and distinct contributions of rostrolateral prefrontal cortex to analogical reasoning and episodic memory retrieval.

Rostrolateral prefrontal cortex (RLPFC) is widely appreciated to support higher cognitive functions, including analogical reasoning and episodic memory retrieval. However, these tasks have typically been studied in isolation, and thus it is unclear whether they involve common or distinct RLPFC mechanisms. Here, we introduce a novel functional magnetic resonance imaging (fMRI) task paradigm to compare brain activity during reasoning and memory tasks while holding bottom-up perceptual stimulation and response demands constant. Univariate analyses on fMRI data from twenty participants identified a large swath of left lateral prefrontal cortex, including RLPFC, that showed common engagement on reasoning trials with valid analogies and memory trials with accurately retrieved source details. Despite broadly overlapping recruitment, multi-voxel activity patterns within left RLPFC reliably differentiated these two trial types, highlighting the presence of at least partially distinct information processing modes. Functional connectivity analyses demonstrated that while left RLPFC showed consistent coupling with the fronto-parietal control network across tasks, its coupling with other cortical areas varied in a task-dependent manner. During the memory task, this region strengthened its connectivity with the default mode and memory retrieval networks, whereas during the reasoning task it coupled more strongly with a nearby left prefrontal region (BA 45) associated with semantic processing, as well as with a superior parietal region associated with visuospatial processing. Taken together, these data suggest a domain-general role for left RLPFC in monitoring and/or integrating task-relevant knowledge representations and showcase how its function cannot solely be attributed to episodic memory or analogical reasoning computations.

Frontoparietal representations of task context support the flexible control of goal-directed cognition.

Cognitive control allows stimulus-response processing to be aligned with internal goals and is thus central to intelligent, purposeful behavior. Control is thought to depend in part on the active representation of task information in prefrontal cortex (PFC), which provides a source of contextual bias on perception, decision making, and action. In the present study, we investigated the organization, influences, and consequences of context representation as human subjects performed a cued sorting task that required them to flexibly judge the relationship between pairs of multivalent stimuli. Using a connectivity-based parcellation of PFC and multivariate decoding analyses, we determined that context is specifically and transiently represented in a region spanning the inferior frontal sulcus during context-dependent decision making. We also found strong evidence that decision context is represented within the intraparietal sulcus, an area previously shown to be functionally networked with the inferior frontal sulcus at rest and during task performance. Rule-guided allocation of attention to different stimulus dimensions produced discriminable patterns of activation in visual cortex, providing a signature of top-down bias over perception. Furthermore, demands on cognitive control arising from the task structure modulated context representation, which was found to be strongest after a shift in task rules. When context representation in frontoparietal areas increased in strength, as measured by the discriminability of high-dimensional activation patterns, the bias on attended stimulus features was enhanced. These results provide novel evidence that illuminates the mechanisms by which humans flexibly guide behavior in complex environments.

Cortical reinstatement mediates the relationship between content-specific encoding activity and subsequent recollection decisions.

Episodic recollection entails the conscious remembrance of event details associated with previously encountered stimuli. Recollection depends on both the establishment of cortical representations of event features during stimulus encoding and the cortical reinstatement of these representations at retrieval. Here, we used multivoxel pattern analyses of functional magnetic resonance imaging data to examine how cortical and hippocampal activity at encoding and retrieval drive recollective memory decisions. During encoding, words were associated with face or scene source contexts. At retrieval, subjects were cued to recollect the source associate of each presented word. Neurally derived estimates of encoding strength and pattern reinstatement in occipitotemporal cortex were computed for each encoding and retrieval trial, respectively. Analyses demonstrated that (1) cortical encoding strength predicted subsequent memory accuracy and reaction time, (2) encoding strength predicted encoding-phase hippocampal activity, and (3) encoding strength and retrieval-phase hippocampal activity predicted the magnitude of cortical reinstatement. Path analyses further indicated that cortical reinstatement partially mediated both the effect of cortical encoding strength and the effect of retrieval-phase hippocampal activity on subsequent source memory performance. Taken together, these results indicate that memory-guided decisions are driven in part by a pathway leading from hippocampally linked cortical encoding of event attributes to hippocampally linked cortical reinstatement at retrieval.

Value-based modulation of memory encoding involves strategic engagement of fronto-temporal semantic processing regions.

A number of prior fMRI studies have focused on the ways in which the midbrain dopaminergic reward system coactivates with hippocampus to potentiate memory for valuable items. However, another means by which people could selectively remember more valuable to-be-remembered items is to be selective in their use of effective but effortful encoding strategies. To broadly examine the neural mechanisms of value on subsequent memory, we used fMRI to assess how differences in brain activity at encoding as a function of value relate to subsequent free recall for words. Each word was preceded by an arbitrarily assigned point value, and participants went through multiple study-test cycles with feedback on their point total at the end of each list, allowing for sculpting of cognitive strategies. We examined the correlation between value-related modulation of brain activity and participants' selectivity index, which measures how close participants were to their optimal point total, given the number of items recalled. Greater selectivity scores were associated with greater differences in the activation of semantic processing regions, including left inferior frontal gyrus and left posterior lateral temporal cortex, during the encoding of high-value words relative to low-value words. Although we also observed value-related modulation within midbrain and ventral striatal reward regions, our fronto-temporal findings suggest that strategic engagement of deep semantic processing may be an important mechanism for selectively encoding valuable items.

Fidelity of neural reactivation reveals competition between memories.

Remembering an event from the past is often complicated by the fact that our memories are cluttered with similar events. Though competition is a fundamental part of remembering, there is little evidence of how mnemonic competition is neurally represented. Here, we assessed whether competition between visual memories is captured in the relative degree to which target vs. competing memories are reactivated within the ventral occipitotemporal cortex (VOTC). To assess reactivation, we used multivoxel pattern analysis of fMRI data, quantifying the degree to which retrieval events elicited patterns of neural activity that matched those elicited during encoding. Consistent with recent evidence, we found that retrieval of visual memories was associated with robust VOTC reactivation and that the degree of reactivation scaled with behavioral expressions of target memory retrieval. Critically, competitive remembering was associated with more ambiguous patterns of VOTC reactivation, putatively reflecting simultaneous reactivation of target and competing memories. Indeed, the more weakly that target memories were reactivated, the more likely that competing memories were later remembered. Moreover, when VOTC reactivation indicated that conflict between target and competing memories was high, frontoparietal mechanisms were markedly engaged, revealing specific neural mechanisms that tracked competing mnemonic evidence. Together, these findings provide unique evidence that neural reactivation captures competition between individual memories, providing insight into how well target memories are retrieved in the present and how likely competing memories will be remembered in the future.

Distributed representations in memory: insights from functional brain imaging.

Forging new memories for facts and events, holding critical details in mind on a moment-to-moment basis, and retrieving knowledge in the service of current goals all depend on a complex interplay between neural ensembles throughout the brain. Over the past decade, researchers have increasingly utilized powerful analytical tools (e.g., multivoxel pattern analysis) to decode the information represented within distributed functional magnetic resonance imaging activity patterns. In this review, we discuss how these methods can sensitively index neural representations of perceptual and semantic content and how leverage on the engagement of distributed representations provides unique insights into distinct aspects of memory-guided behavior. We emphasize that, in addition to characterizing the contents of memories, analyses of distributed patterns shed light on the processes that influence how information is encoded, maintained, or retrieved, and thus inform memory theory. We conclude by highlighting open questions about memory that can be addressed through distributed pattern analyses.

Detecting individual memories through the neural decoding of memory states and past experience.

A wealth of neuroscientific evidence indicates that our brains respond differently to previously encountered than to novel stimuli. There has been an upswell of interest in the prospect that functional MRI (fMRI), when coupled with multivariate data analysis techniques, might allow the presence or absence of individual memories to be detected from brain activity patterns. This could have profound implications for forensic investigations and legal proceedings, and thus the merits and limitations of such an approach are in critical need of empirical evaluation. We conducted two experiments to investigate whether neural signatures of recognition memory can be reliably decoded from fMRI data. In Exp. 1, participants were scanned while making explicit recognition judgments for studied and novel faces. Multivoxel pattern analysis (MVPA) revealed a robust ability to classify whether a given face was subjectively experienced as old or new, as well as whether recognition was accompanied by recollection, strong familiarity, or weak familiarity. Moreover, a participant's subjective mnemonic experiences could be reliably decoded even when the classifier was trained on the brain data from other individuals. In contrast, the ability to classify a face's objective old/new status, when holding subjective status constant, was severely limited. This important boundary condition was further evidenced in Exp. 2, which demonstrated that mnemonic decoding is poor when memory is indirectly (implicitly) probed. Thus, although subjective memory states can be decoded quite accurately under controlled experimental conditions, fMRI has uncertain utility for objectively detecting an individual's past experiences.

Behavioral representational similarity analysis reveals how episodic learning is influenced by and reshapes semantic memory.

While semantic and episodic memory have been shown to influence each other, uncertainty remains as to how this interplay occurs. We introduce a behavioral representational similarity analysis approach to assess whether semantic space can be subtly re-sculpted by episodic learning. Eighty participants learned word pairs that varied in semantic relatedness, and learning was bolstered via either testing or restudying. Next-day recall is superior for semantically related pairs, but there is a larger benefit of testing for unrelated pairs. Analyses of representational change reveal that successful recall is accompanied by a pulling together of paired associates, with cue words in semantically related (but not unrelated) pairs changing more across learning than target words. Our findings show that episodic learning is associated with systematic and asymmetrical distortions of semantic space which improve later recall by making cues more predictive of targets, reducing interference from potential lures, and establishing novel connections within pairs.

Establishing a causal role for left ventrolateral prefrontal cortex in value-directed memory encoding with high-definition transcranial direct current stimulation.

One critical approach for promoting the efficiency of memory is to adopt selective encoding strategies to prioritize more valuable information. Past neuroimaging studies have shown that value-directed modulation of verbal memory depends heavily on the engagement of left-lateralized semantic processing regions, particularly in the ventrolateral prefrontal cortex (VLPFC). In the present study, we used high-definition transcranial direct current stimulation (HD-tDCS) to seek evidence for a causal role of left VLPFC in supporting the memory advantage for high-value items. Three groups of healthy young adult participants were presented with lists of words to remember, with each word accompanied by an arbitrarily assigned point value. During the first session, all participants received sham stimulation as they encoded five lists of 30 words each. Two of these lists were immediately tested with free recall, with feedback given to allow participants to develop metacognitive insight and strategies to maximize their point total. The second session had the exact same structure as the first, but the groups differed in whether they received continued sham stimulation (N = 22) or anodal stimulation of the left VLPFC (N = 21) or right VLPFC (N = 20). Those lists not tested with immediate recall were tested with recognition judgments after a one-day delay. Since no brain stimulation was applied during this Day 2 test, any performance differences can be attributed to the effects of stimulation on Day 1 encoding processes. Anodal stimulation of left VLPFC significantly boosted participants' memory encoding selectivity. In comparison, no such effect was seen in participants who received right VLPFC or sham stimulation. Estimates of recollection- and familiarity-based responding revealed that left VLPFC stimulation specifically amplified the effects of item value on recollection. These results demonstrate a causal role for left VLPFC in the implementation of selective value-directed encoding strategies, putatively by boosting deep semantic processing of high-value words. Our findings also provide further evidence on the hemispheric lateralization of value-directed verbal memory encoding.

Enhancing learning and retention with distinctive virtual reality environments and mental context reinstatement.

Memory is inherently context-dependent: internal and environmental cues become bound to learnt information, and the later absence of these cues can impair recall. Here, we developed an approach to leverage context-dependence to optimise learning of challenging, interference-prone material. While navigating through desktop virtual reality (VR) contexts, participants learnt 80 foreign words in two phonetically similar languages. Those participants who learnt each language in its own unique context showed reduced interference and improved one-week retention (92%), relative to those who learnt the languages in the same context (76%)-however, this advantage was only apparent if participants subjectively experienced VR-based contexts as "real" environments. A follow-up fMRI experiment confirmed that reinstatement of brain activity patterns associated with the original encoding context during word retrieval was associated with improved recall performance. These findings establish that context-dependence can be harnessed with VR to optimise learning and showcase the important role of mental context reinstatement.

Modulation of inferotemporal cortex activation during verbal working memory maintenance.

Regions of the left inferotemporal cortex are involved in visual word recognition and semantics. We utilized functional magnetic resonance imaging to localize an inferotemporal language area and to demonstrate that this area is involved in the active maintenance of visually presented words in working memory. Maintenance activity in this inferotemporal area showed an effect of memory load for words, but not pseudowords. In the absence of visual input, the selective modulation of this language-related inferotemporal area for the maintenance of words is accompanied by an increased functional connectivity with left prefrontal cortex. These results demonstrate an involvement of inferotemporal cortex in verbal working memory and provide neurophysiological support for the notion that nonphonological language representations can be recruited in the service of verbal working memory. More generally, they suggest that verbal working memory should be conceptualized as the frontally guided, sustained activation of pre-existing cortical language representations.