Eye movements support behavioral pattern completion.

The ability to recall a detailed event from a simple reminder is supported by pattern completion, a cognitive operation performed by the hippocampus wherein existing mnemonic representations are retrieved from incomplete input. In behavioral studies, pattern completion is often inferred through the false endorsement of lure (i.e., similar) items as old. However, evidence that such a response is due to the specific retrieval of a similar, previously encoded item is severely lacking. We used eye movement (EM) monitoring during a partial-cue recognition memory task to index reinstatement of lure images behaviorally via the recapitulation of encoding-related EMs or gaze reinstatement. Participants reinstated encoding-related EMs following degraded retrieval cues and this reinstatement was negatively correlated with accuracy for lure images, suggesting that retrieval of existing representations (i.e., pattern completion) underlies lure false alarms. Our findings provide evidence linking gaze reinstatement and pattern completion and advance a functional role for EMs in memory retrieval.

Neural reactivation and judgements of vividness reveal separable contributions to mnemonic representation.

Mnemonic representations vary in fidelity, sharpness, and strength-qualities that can be examined using both introspective judgements of mental states and objective measures of brain activity. Subjective and objective measures are both valid ways of "reading out" the content of someone's internal mnemonic states, each with different strengths and weaknesses. St-Laurent and colleagues (2015) compared the neural correlates of memory vividness ratings with patterns of neural reactivation evoked during memory recall and found considerable overlap between the two, suggesting a common neural basis underlying these different markers of representational quality. Here we extended this work with meta-analytic methods by pooling together four neuroimaging datasets in order to contrast the neural substrates of neural reactivation and those of vividness judgements. While reactivation and vividness judgements correlated positively with one another and were associated with common univariate activity in the dorsal attention network and anterior hippocampus, some notable differences were also observed. Vividness judgments were tied to stronger activation in the striatum and dorsal attention network, together with activity suppression in default mode network nodes. We also observed a trend for reactivation to be more closely associated with early visual cortex activity. A mediation analysis found support for the hypothesis that neural reactivation is necessary for memory vividness, with activity in the anterior hippocampus associated with greater reactivation. Our results suggest that neural reactivation and vividness judgements reflect common mnemonic processes but differ in the extent to which they engage effortful, attentional processes. Additionally, the similarity between reactivation and vividness appears to arise, partly, through hippocampal engagement during memory retrieval.

Effects of temporal order and intentionality on reflective attention to words in noise.

Speech perception in noise is a cognitively demanding process that challenges not only the auditory sensory system, but also cognitive networks involved in attention. The predictive coding theory has been influential in characterizing the influence of prior context on processing incoming auditory stimuli, with comparatively less research dedicated to "postdictive" processes and subsequent context effects on speech perception. Effects of subsequent semantic context were evaluated while manipulating the relationship of three target words presented in noise and the temporal position of targets compared to the subsequent contextual cue, demonstrating that subsequent context benefits were present regardless of whether the targets were related to each other and did not depend on the position of the target. However, participants instructed to focus on the relation between target and cue performed worse than those who did not receive this instruction, suggesting a disruption of a natural process of continuous speech recognition. We discuss these findings in relation to lexical commitment and stimulus-driven attention to short-term memory as mechanisms of subsequent context integration.

Holding On to the Past: Older Adults Show Lingering Neural Activation of No-Longer-Relevant Items in Working Memory.

Goal-relevant information can be maintained in working memory over a brief delay interval to guide an upcoming decision. There is also evidence suggesting the existence of a complementary process: namely, the ability to suppress information that is no longer relevant to ongoing task goals. Moreover, this ability to suppress or inhibit irrelevant information appears to decline with age. In this study, we compared younger and older adults undergoing fMRI on a working memory task designed to address whether the modulation of neural representations of relevant and no-longer-relevant items during a delay interval is related to age and overall task performance. Following from the theoretical predictions of the inhibitory deficit hypothesis of aging, we hypothesized that older adults would show higher activation of no-longer-relevant items during a retention delay compared to young adults and that higher activation of these no-longer-relevant items would predict worse recognition memory accuracy for relevant items. Our results support this prediction and more generally demonstrate the importance of goal-driven modulation of neural activity in successful working memory maintenance. Furthermore, we showed that the largest age differences in the regulation of category-specific pattern activity during working memory maintenance were seen throughout the medial temporal lobe and prominently in the hippocampus, further establishing the importance of "long-term memory" retrieval mechanisms in the context of high-load working memory tasks that place large demands on attentional selection mechanisms.

Eye Movement Reinstatement and Neural Reactivation During Mental Imagery.

Half a century ago, Donald Hebb posited that mental imagery is a constructive process that emulates perception. Specifically, Hebb claimed that visual imagery results from the reactivation of neural activity associated with viewing images. He also argued that neural reactivation and imagery benefit from the re-enactment of eye movement patterns that first occurred at viewing (fixation reinstatement). To investigate these claims, we applied multivariate pattern analyses to functional MRI (fMRI) and eye tracking data collected while healthy human participants repeatedly viewed and visualized complex images. We observed that the specificity of neural reactivation correlated positively with vivid imagery and with memory for stimulus image details. Moreover, neural reactivation correlated positively with fixation reinstatement, meaning that image-specific eye movements accompanied image-specific patterns of brain activity during visualization. These findings support the conception of mental imagery as a simulation of perception, and provide evidence consistent with the supportive role of eye movement in neural reactivation.

The Primacy of Spatial Context in the Neural Representation of Events.

Some theories of episodic memory hypothesize that spatial context plays a fundamental role in episodic memory, acting as a scaffold on which episodes are constructed. A prediction based on this hypothesis is that spatial context should play a primary role in the neural representation of an event. To test this hypothesis in humans, male and female participants imagined events, composed of familiar locations, people, and objects, during an fMRI scan. We used multivoxel pattern analysis to determine the neural areas in which events could be discriminated based on each feature. We found that events could be discriminated according to their location in areas throughout the autobiographical memory network, including the parahippocampal cortex and posterior hippocampus, retrosplenial cortex, posterior cingulate cortex, precuneus, and medial prefrontal cortex. Events were also discriminable based on person and object features, but in fewer regions. Comparing classifier performance in regions involved in memory for scenes and events demonstrated that the location of an event was more accurately classified than the person or object involved. These results support theories that suggest that spatial context is a prominent defining feature of episodic memory.SIGNIFICANCE STATEMENT Remembered and imagined events are complex, consisting of many elements, including people, objects, and locations. In this study, we sought to determine how these types of elements differentially contribute to how the brain represents an event. Participants imagined events consisting of familiar locations, people, and objects (e.g., kitchen, mom, umbrella) while their brain activity was recorded with fMRI. We found that the neural patterns of activity in brain regions associated with spatial and episodic memory could distinguish events based on their location, and to some extent, based on the people and objects involved. These results suggest that the spatial context of an event plays an important role in how an event is represented in the brain.

Noise differentially impacts phoneme representations in the auditory and speech motor systems.

Although it is well accepted that the speech motor system (SMS) is activated during speech perception, the functional role of this activation remains unclear. Here we test the hypothesis that the redundant motor activation contributes to categorical speech perception under adverse listening conditions. In this functional magnetic resonance imaging study, participants identified one of four phoneme tokens (/ba/, /ma/, /da/, or /ta/) under one of six signal-to-noise ratio (SNR) levels (-12, -9, -6, -2, 8 dB, and no noise). Univariate and multivariate pattern analyses were used to determine the role of the SMS during perception of noise-impoverished phonemes. Results revealed a negative correlation between neural activity and perceptual accuracy in the left ventral premotor cortex and Broca's area. More importantly, multivoxel patterns of activity in the left ventral premotor cortex and Broca's area exhibited effective phoneme categorization when SNR ≥ -6 dB. This is in sharp contrast with phoneme discriminability in bilateral auditory cortices and sensorimotor interface areas (e.g., left posterior superior temporal gyrus), which was reliable only when the noise was extremely weak (SNR > 8 dB). Our findings provide strong neuroimaging evidence for a greater robustness of the SMS than auditory regions for categorical speech perception in noise. Under adverse listening conditions, better discriminative activity in the SMS may compensate for loss of specificity in the auditory system via sensorimotor integration.

Distributed neural representations of phonological features during speech perception.

A fundamental goal of the human auditory system is to map complex acoustic signals onto stable internal representations of the basic sound patterns of speech. Phonemes and the distinctive features that they comprise constitute the basic building blocks from which higher-level linguistic representations, such as words and sentences, are formed. Although the neural structures underlying phonemic representations have been well studied, there is considerable debate regarding frontal-motor cortical contributions to speech as well as the extent of lateralization of phonological representations within auditory cortex. Here we used functional magnetic resonance imaging (fMRI) and multivoxel pattern analysis to investigate the distributed patterns of activation that are associated with the categorical and perceptual similarity structure of 16 consonant exemplars in the English language used in Miller and Nicely's (1955) classic study of acoustic confusability. Participants performed an incidental task while listening to phonemes in the MRI scanner. Neural activity in bilateral anterior superior temporal gyrus and supratemporal plane was correlated with the first two components derived from a multidimensional scaling analysis of a behaviorally derived confusability matrix. We further showed that neural representations corresponding to the categorical features of voicing, manner of articulation, and place of articulation were widely distributed throughout bilateral primary, secondary, and association areas of the superior temporal cortex, but not motor cortex. Although classification of phonological features was generally bilateral, we found that multivariate pattern information was moderately stronger in the left compared with the right hemisphere for place but not for voicing or manner of articulation.

Recency, repetition, and the multidimensional basis of recognition memory.

Recency and repetition are two factors that have large effects on human memory performance. One way of viewing the beneficial impact of these variables on recognition memory is to assume that both factors modulate a unidimensional memory trace strength. Although previous functional neuroimaging studies have indicated that recency and repetition may modulate similar brain structures, particularly in the region of the inferior parietal cortex, there is extensive behavioral evidence that human subjects can make independent and accurate recognition memory judgments about both an item's recency and its frequency. In the present study, we used fMRI to examine patterns of brain activity during recognition memory for auditory-verbal stimuli that were parametrically and orthogonally manipulated in terms of recency and number of repetitions. We found in a continuous recognition paradigm that the lateral inferior parietal cortex, a region that has previously been associated with recollective forms of memory, is highly sensitive to recency but not repetition. In a multivariate analysis of whole-brain activation patterns, we found orthogonal components that dissociated recency and repetition variables, indicating largely independent neural bases underlying these two factors. The results demonstrate that although both recency and repetition dramatically improve recognition memory performance, the neural bases for this improvement are dissociable, and thus are difficult to explain in terms of access to a unitary memory trace.

Memory reactivation in healthy aging: evidence of stimulus-specific dedifferentiation.

We investigated how aging affects the neural specificity of mental replay, the act of conjuring up past experiences in one's mind. We used functional magnetic resonance imaging (fMRI) and multivariate pattern analysis to quantify the similarity between brain activity elicited by the perception and memory of complex multimodal stimuli. Young and older human adults viewed and mentally replayed short videos from long-term memory while undergoing fMRI. We identified a wide array of cortical regions involved in visual, auditory, and spatial processing that supported stimulus-specific representation at perception as well as during mental replay. Evidence of age-related dedifferentiation was subtle at perception but more salient during mental replay, and age differences at perception could not account for older adults' reduced neural reactivation specificity. Performance on a post-scan recognition task for video details correlated with neural reactivation in young but not in older adults, indicating that in-scan reactivation benefited post-scan recognition in young adults, but that some older adults may have benefited from alternative rehearsal strategies. Although young adults recalled more details about the video stimuli than older adults on a post-scan recall task, patterns of neural reactivation correlated with post-scan recall in both age groups. These results demonstrate that the mechanisms supporting recall and recollection are linked to accurate neural reactivation in both young and older adults, but that age affects how efficiently these mechanisms can support memory's representational specificity in a way that cannot simply be accounted for by degraded sensory processes.

Levels of processing in working memory: differential involvement of frontotemporal networks.

How does the brain maintain to-be-remembered information in working memory (WM), particularly when the focus of attention is drawn to processing other information? Cognitive models of WM propose that when items are displaced from focal attention recall involves retrieval from long-term memory (LTM). In this fMRI study, we tried to clarify the role of LTM in performance on a WM task and the type of representation that is used to maintain an item in WM during rehearsal-filled versus distractor-filled delays. Participants made a deep or shallow levels-of-processing (LOP) decision about a single word at encoding and tried to recall the word after a delay filled with either rehearsal of the word or a distracting math task. Recalling one word after 10 sec of distraction demonstrated behavioral and neural indices of retrieval from LTM (i.e., LOP effects and medial-temporal lobe activity). In contrast, recall after rehearsal activated cortical areas that reflected reporting the word from focal attention. In addition, areas that showed an LOP effect at encoding (e.g., left ventrolateral VLPFC and the anterior temporal lobes [ATLs]) were reactivated at recall, especially when recall followed distraction. Moreover, activity in left VLPFC during encoding, left ATL during the delay, and left hippocampus during retrieval predicted recall success after distraction. Whereas shallow LOP and rehearsal-related areas supported active maintenance of one item in focal attention, the behavioral processes and neural substrates that support LTM supported recall of one item after it was displaced from focal attention.

Distributed Patterns of Reactivation Predict Vividness of Recollection.

According to the principle of reactivation, memory retrieval evokes patterns of brain activity that resemble those instantiated when an event was first experienced. Intuitively, one would expect neural reactivation to contribute to recollection (i.e., the vivid impression of reliving past events), but evidence of a direct relationship between the subjective quality of recollection and multiregional reactivation of item-specific neural patterns is lacking. The current study assessed this relationship using fMRI to measure brain activity as participants viewed and mentally replayed a set of short videos. We used multivoxel pattern analysis to train a classifier to identify individual videos based on brain activity evoked during perception and tested how accurately the classifier could distinguish among videos during mental replay. Classification accuracy correlated positively with memory vividness, indicating that the specificity of multivariate brain patterns observed during memory retrieval was related to the subjective quality of a memory. In addition, we identified a set of brain regions whose univariate activity during retrieval predicted both memory vividness and the strength of the classifier's prediction irrespective of the particular video that was retrieved. Our results establish distributed patterns of neural reactivation as a valid and objective marker of the quality of recollection.

The effect of rehearsal rate and memory load on verbal working memory.

While many neuroimaging studies have investigated verbal working memory (WM) by manipulating memory load, the subvocal rehearsal rate at these various memory loads has generally been left uncontrolled. Therefore, the goal of this study was to investigate how mnemonic load and the rate of subvocal rehearsal modulate patterns of activity in the core neural circuits underlying verbal working memory. Using fMRI in healthy subjects, we orthogonally manipulated subvocal rehearsal rate and memory load in a verbal WM task with long 45-s delay periods. We found that middle frontal gyrus (MFG) and superior parietal lobule (SPL) exhibited memory load effects primarily early in the delay period and did not exhibit rehearsal rate effects. In contrast, we found that inferior frontal gyrus (IFG), premotor cortex (PM) and Sylvian-parietal-temporal region (area Spt) exhibited approximately linear memory load and rehearsal rate effects, with rehearsal rate effects lasting through the entire delay period. These results indicate that IFG, PM and area Spt comprise the core articulatory rehearsal areas involved in verbal WM, while MFG and SPL are recruited in a general supervisory role once a memory load threshold in the core rehearsal network has been exceeded.

Short-term retention of a single word relies on retrieval from long-term memory when both rehearsal and refreshing are disrupted.

Many working memory (WM) models propose that the focus of attention (or primary memory) has a capacity limit of one to four items, and therefore, that performance on WM tasks involves retrieving some items from long-term (or secondary) memory (LTM). In the present study, we present evidence suggesting that recall of even one item on a WM task can involve retrieving it from LTM. The WM task required participants to make a deep (living/nonliving) or shallow ("e"/no "e") level-of-processing (LOP) judgment on one word and to recall the word after a 10-s delay on each trial. During the delay, participants either rehearsed the word or performed an easy or a hard math task. When the to-be-remembered item could be rehearsed, recall was fast and accurate. When it was followed by a math task, recall was slower, error-prone, and benefited from a deeper LOP at encoding, especially for the hard math condition. The authors suggest that a covert-retrieval mechanism may have refreshed the item during easy math, and that the hard math condition shows that even a single item cannot be reliably held in WM during a sufficiently distracting task--therefore, recalling the item involved retrieving it from LTM. Additionally, performance on a final free recall (LTM) test was better for items recalled following math than following rehearsal, suggesting that initial recall following math involved elaborative retrieval from LTM, whereas rehearsal did not. The authors suggest that the extent to which performance on WM tasks involves retrieval from LTM depends on the amounts of disruption to both rehearsal and covert-retrieval/refreshing maintenance mechanisms.

The relationship between delay period eye movements and visuospatial memory.

We investigated whether overt shifts of attention were associated with visuospatial memory performance. Participants were required to study the locations of a set of visual objects and subsequently detect changes to the spatial location of one of the objects following a brief delay period. Relational information regarding the locations among all of the objects could be used to support performance on the task (Experiment 1) or relational information was removed during test and location manipulation judgments had to be made for a singly presented target item (Experiment 2). We computed the similarity of the fixation patterns in space during the study phase to the fixations made during the delay period. Greater fixation pattern similarity across participants was associated with higher accuracy when relational information was available at test (Experiment 1); however, this association was not observed when the target item was presented in isolation during the test display (Experiment 2). Similarly, increased fixation pattern similarity on a given trial (within participants) was associated with successful task performance when the relations among studied items could be used for comparison (Experiment 1), but not when memory for absolute spatial location was assessed (Experiment 2). This pattern of behavior and performance on the two tasks suggested that eye movements facilitated memory for the relationships among objects. Shifts of attention through eye movements may provide a mechanism for the maintenance of relational visuospatial memory.

The neural basis of vivid memory is patterned on perception.

When we have a rich and vivid memory for a past experience, it often feels like we are transported back in time to witness once again this event. Indeed, a perfect memory would exactly mimic the experiential quality of direct sensory perception. We used fMRI and multivoxel pattern analysis to map and quantify the similarity between patterns of activation evoked by direct perception of a diverse set of short video clips and the vivid remembering, with closed eyes, of these clips. We found that the patterns of distributed brain activation during vivid memory mimicked the patterns evoked during sensory perception. Using whole-brain patterns of activation evoked by perception of the videos, we were able to accurately classify brain patterns that were elicited when participants tried to vividly recall those same videos. A discriminant analysis of the activation patterns associated with each video revealed a high degree (explaining over 80% of the variance) of shared representational similarity between perception and memory. These results show that complex, multifeatured memory involves a partial reinstatement of the whole pattern of brain activity that is evoked during initial perception of the stimulus.

Serotonin transporter binding after recovery from bulimia nervosa.

OBJECTIVE: Physiological and pharmacological studies indicate that altered brain serotonin (5-HT) activity could contribute to a susceptibility to develop appetitive and behavioral alterations that are characteristic of bulimia nervosa (BN). METHOD: Eight individuals recovered from BN (REC BN) and eight healthy control women were scanned with [11C]DASB and positron emission tomography imaging of the 5-HT transporter (5-HTT). Logan graphical analysis was applied, and parametric binding potential (BP(nondisplaceable (ND)) ) images were generated. Voxel-by-voxel t-tests and a region of interest (ROI) analysis were conducted. RESULTS: REC BN had significantly lower [11C]DASB BP(ND) in midbrain, superior and inferior cingulate and significantly higher [11C]DASB BP(ND) in anterior cingulate and superior temporal gyrus in the voxel-based analysis. ROI analysis indicated lower [11C]DASB BP(ND) in midbrain (p = .07), containing the dorsal raphe, in REC BN, consistent with our earlier studies. DISCUSSION: These preliminary findings of a small-scale study confirm and extend previous data suggesting that ill and recovered BN have altered 5-HTT measures, which potentially contribute to BN symptomatology and/or differential responses to medication.

The influence of long-term memory on working memory: Age-differences in proactive facilitation and interference.

Prior learning can hinder subsequent memory, especially when there is conflict between old and new information. The ability to handle this proactive interference is an important source of differences in memory performance between younger and older adults. In younger participants, Oberauer et al. (2017, Journal of Experimental Psychology: Learning, Memory, and Cognition, 43[1], 1) report evidence of proactive facilitation from previously learned information in a working memory task in the absence of proactive interference between long-term and working memory. In the present work, we examine the generality of these findings to different stimulus materials and to older adults. Participants first learned image-word associations and then completed an image-word working memory task. Some pairs were the same as those initially learned, for which we expected facilitation relative to previously unencountered pairs. Other pairs were made up of previously learned elements in different combinations, for which we might expect interference. Younger and older participants showed similar levels of facilitation from previously learned associations relative to new pairs. In addition, older participants exhibited proactive interference from long-term to working memory, whereas younger participants exhibited facilitation, even for pairings that conflicted with those learned earlier in the experiment. These findings confirm older adults' greater susceptibility to proactive interference and we discuss the theoretical implications of younger adults' apparent immunity to interference.

Absence of a mere-exposure effect in older and younger adults.

The mere-exposure effect, in which repeated stimuli are liked more than novel stimuli, is a well-known effect. However, little research has studied adult age differences in mere-exposure effects, despite possible applications in helping older adults transition to new living environments. Here, we report four experiments assessing mere-exposure to neutral-face stimuli in groups of older and younger adult participants tested online. In each experiment, repeated face exposure did not increase liking within either age group; rather, Bayesian evidence favored the null hypothesis of no effect. Older adults reported higher overall liking ratings relative to younger adults, and both groups preferred younger faces, though this tendency was stronger in the younger group. Further exploratory analysis considering factors such as gender or race of the faces and participants did not reveal any consistent results for the mere-exposure effect. We discuss these findings in relation to other recent studies reporting mixed evidence for mere-exposure effects. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Four-modality imaging of unmedicated subjects with schizophrenia: 18F-fluorodeoxyglucose and 18F-fallypride PET, diffusion tensor imaging, and MRI.

Diminished prefrontal function, dopaminergic abnormalities in the striatum and thalamus, reductions in white matter integrity and frontotemporal gray matter deficits are the most replicated findings in schizophrenia. We used four imaging modalities (18F-fluorodeoxyglucose and 18F-fallypride PET, diffusion tensor imaging, structural MRI) in 19 healthy and 25 schizophrenia subjects to assess the relationship between functional (dopamine D2/D3 receptor binding potential, glucose metabolic rate) and structural (fractional anisotropy, MRI) correlates of schizophrenia and their additive diagnostic prediction potential. Multivariate ANOVA was used to compare structural and functional image sets for identification of schizophrenia. Integration of data from all four modalities yielded better predictive power than less inclusive combinations, specifically in the thalamus, left dorsolateral prefrontal and temporal regions. Among the modalities, fractional anisotropy showed highest discrimination in white matter whereas 18F-fallypride binding showed highest discrimination in gray matter. Structural and functional modalities displayed comparable discriminative power but different topography, with higher sensitivity of structural modalities in the left prefrontal region. Combination of functional and structural imaging modalities with inclusion of both gray and white matter appears most effective in diagnostic discrimination. The highest sensitivity of 18F-fallypride PET to gray matter changes in schizophrenia supports the primacy of dopaminergic abnormalities in its pathophysiology.