Decoding of Working Memory Contents in Auditory Cortex Is Not Distractor-Resistant.

Working memory enables the temporary storage of relevant information in the service of behavior. Neuroimaging studies have suggested that sensory cortex is involved in maintaining contents in working memory. This raised the question of how sensory regions maintain memory representations during the exposure to distracting stimuli. Multivariate pattern analysis of fMRI signals in visual cortex has shown that the contents of visual working memory could be decoded concurrently with passively viewed distractors. The present fMRI study tested whether this finding extends to auditory working memory and to active distractor processing. We asked participants to memorize the pitch of a target sound and to compare it with a probe sound presented after a 13 s delay period. In separate conditions, we compared a blank delay phase (no distraction) with either passive listening to, or active processing of, an auditory distractor presented throughout the memory delay. Consistent with previous reports, pitch-specific memory information could be decoded in auditory cortex during the delay in trials without distraction. In contrast, decoding of target sounds in early auditory cortex dropped to chance level during both passive and active distraction. This was paralleled by memory performance decrements under distraction. Extending the analyses beyond sensory cortex yielded some evidence for memory content-specific activity in inferior frontal and superior parietal cortex during active distraction. In summary, while our findings question the involvement of early auditory cortex in the maintenance of distractor-resistant working memory contents, further research should elucidate the role of hierarchically higher regions.SIGNIFICANCE STATEMENT Information about sensory features held in working memory can be read out from hemodynamic activity recorded in human sensory cortices. Moreover, visual cortex can in parallel store visual content and process newly incoming, task-irrelevant visual input. The present study investigated the role of auditory cortex for working memory maintenance under distraction. While memorized sound frequencies could be decoded in auditory cortex in the absence of distraction, auditory distraction during the delay phase impaired memory performance and prevented decoding of information stored in working memory. Apparently, early auditory cortex is not sufficient to represent working memory contents under distraction that impairs performance. However, exploratory analyses indicated that, under distraction, higher-order frontal and parietal regions might contribute to content-specific working memory storage.

Interference between items stored for distinct tasks in visual working memory.

The action perspective on working memory suggests that memory representations are coded according to their specific temporal and behavioral task demands. This stands in contrast to theories that assume representations are stored in a task-agnostic format within a "common workspace". Here, we tested whether visual items that are memorized for different tasks are stored separately from one another or show evidence of inter-item interference during concurrent maintenance, indicating a common storage. In two experiments, we combined a framing memory task (memorize a motion direction for continuous direction report) with an embedded memory task (memorize a motion direction for a binary direction discrimination) that was placed within the retention period of the framing task. Even though the temporal and action demands were item specific, we observed two types of interference effects between the items: The embedded motion direction was (1) repulsed away and (2) degraded in precision by the motion direction of the item in the framing task. Repulsion and precision degradation increased with item similarity when both items were concurrently held in working memory. In contrast, perceptual and iconic memory control conditions revealed weaker repulsion overall and no interference effect on precision during the stimulus processing stages prior to working memory consolidation. Thus, additional inter-item interference arose uniquely within working memory. Together, our results present evidence that items that are stored for distinct tasks to be performed at distinct points in time, reside in a common workspace in working memory.

Decoding Concurrent Representations of Pitch and Location in Auditory Working Memory.

Multivariate analyses of hemodynamic signals serve to identify the storage of specific stimulus contents in working memory (WM). Representations of visual stimuli have been demonstrated both in sensory regions and in higher cortical areas. While previous research has typically focused on the WM maintenance of a single content feature, it remains unclear whether two separate features of a single object can be decoded concurrently. Also, much less evidence exists for representations of auditory compared with visual stimulus features. To address these issues, human participants had to memorize both pitch and perceived location of one of two sample sounds. After a delay phase, they were asked to reproduce either pitch or location. At recall, both features showed comparable levels of discriminability. Region of interest (ROI)-based decoding of functional magnetic resonance imaging (fMRI) data during the delay phase revealed feature-selective activity for both pitch and location of a memorized sound in auditory cortex and superior parietal lobule. The latter region showed higher decoding accuracy for location than pitch. In addition, location could be decoded from angular and supramarginal gyrus and both superior and inferior frontal gyrus. The latter region also showed a trend for decoding of pitch. We found no region exclusively coding pitch memory information. In summary, the present study yielded evidence for concurrent representations of pitch and location of a single object both in sensory cortex and in hierarchically higher regions, pointing toward representation formats that enable feature integration within the same anatomic brain regions.SIGNIFICANCE STATEMENT Decoding of hemodynamic signals serves to identify brain regions involved in the storage of stimulus-specific information in working memory (WM). While to-be-remembered information typically consists of several features, most previous investigations have focused on the maintenance of one memorized feature belonging to one visual object. The present study assessed the concurrent storage of two features of the same object in auditory WM. We found that both pitch and location of memorized sounds were decodable both in early sensory areas, in higher-level superior parietal cortex and, to a lesser extent, in inferior frontal cortex. While auditory cortex is known to process different features in parallel, their concurrent representation in parietal regions may support the integration of object features in WM.

Decoding Spatial Versus Non-spatial Processing in Auditory Working Memory.

OBJECTIVE: Research on visual working memory has shown that individual stimulus features are processed in both specialized sensory regions and higher cortical areas. Much less evidence exists for auditory working memory. Here, a main distinction has been proposed between the processing of spatial and non-spatial sound features. Our aim was to examine feature-specific activation patterns in auditory working memory. METHODS: We collected fMRI data while 28 healthy adults performed an auditory delayed match-to-sample task. Stimuli were abstract sounds characterized by both spatial and non-spatial information, i.e., interaural time delay and central frequency, respectively. In separate recording blocks, subjects had to memorize either the spatial or non-spatial feature, which had to be compared with a probe sound presented after a short delay. We performed both univariate and multivariate comparisons between spatial and non-spatial task blocks. RESULTS: Processing of spatial sound features elicited a higher activity in a small cluster in the superior parietal lobe than did sound pattern processing, whereas there was no significant activation difference for the opposite contrast. The multivariate analysis was applied using a whole-brain searchlight approach to identify feature-selective processing. The task-relevant auditory feature could be decoded from multiple brain regions including the auditory cortex, posterior temporal cortex, middle occipital gyrus, and extended parietal and frontal regions. CONCLUSION: In summary, the lack of large univariate activation differences between spatial and non-spatial processing could be attributable to the identical stimulation in both tasks. In contrast, the whole-brain multivariate analysis identified feature-specific activation patterns in widespread cortical regions. This suggests that areas beyond the auditory dorsal and ventral streams contribute to working memory processing of auditory stimulus features.

Object-based attention prioritizes working memory contents at a theta rhythm.

Attention selects relevant information regardless of whether it is physically present or internally stored in working memory. Perceptual research has shown that attentional selection of external information is better conceived as rhythmic prioritization than as stable allocation. Here we tested this principle using information processing of internal representations held in working memory. Participants memorized 4 spatial positions that formed the end points of 2 objects. One of the positions was cued for a delayed match-nonmatch test. When uncued positions were probed, participants responded faster to uncued positions located on the same object as the cued position than to those located on the other object, revealing object-based attention in working memory. Manipulating the interval between cue and probe at a high temporal resolution revealed that reaction times oscillated at a theta rhythm of 6 Hz. Moreover, oscillations showed an antiphase relationship between memorized but uncued positions on the same versus other object as the cued position, suggesting that attentional prioritization fluctuated rhythmically in an object-based manner. Our results demonstrate the highly rhythmic nature of attentional selection in working memory. Moreover, the striking similarity between rhythmic attentional selection of mental representations and perceptual information suggests that attentional oscillations are a general mechanism of information processing in human cognition. These findings have important implications for current, attention-based models of working memory. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Context information supports serial dependence of multiple visual objects across memory episodes.

Serial dependence is thought to promote perceptual stability by compensating for small changes of an object's appearance across memory episodes. So far, it has been studied in situations that comprised only a single object. The question of how we selectively create temporal stability of several objects remains unsolved. In a memory task, objects can be differentiated by their to-be-memorized feature (content) as well as accompanying discriminative features (context). We test whether congruent context features, in addition to content similarity, support serial dependence. In four experiments, we observe a stronger serial dependence between objects that share the same context features across trials. Apparently, the binding of content and context features is not erased but rather carried over to the subsequent memory episode. As this reflects temporal dependencies in natural settings, our findings reveal a mechanism that integrates corresponding content and context features to support stable representations of individualized objects over time.

Visual objects interact differently during encoding and memory maintenance.

The storage mechanisms of working memory are the matter of an ongoing debate. The sensory recruitment hypothesis states that memory maintenance and perceptual encoding rely on the same neural substrate. This suggests that the same cortical mechanisms that shape object perception also apply to maintained memory content. We tested this prediction using the Direction Illusion, i.e., the mutual repulsion of two concurrently visible motion directions. Participants memorized the directions of two random dot patterns for later recall. In Experiments 1 and 2, we varied the temporal separation of spatially distinct stimuli to manipulate perceptual concurrency, while keeping concurrency within working memory constant. We observed mutual motion repulsion only under simultaneous stimulus presentation, but proactive repulsion and retroactive attraction under immediate stimulus succession. At inter-stimulus intervals of 0.5 and 2 s, however, proactive repulsion vanished, while the retroactive attraction remained. In Experiment 3, we presented both stimuli at the same spatial position and observed a reappearance of the repulsion effect. Our results indicate that the repulsive mechanisms that shape object perception across space fade during the transition from a perceptual representation to a consolidated memory content. This suggests differences in the underlying structure of perceptual and mnemonic representations. The persistence of local interactions, however, indicates different mechanisms of spatially global and local feature interactions.

Cognitive Performance and Psychological Distress in Breast Cancer Patients at Disease Onset.

OBJECTIVE: Many cancer patients complain about cognitive dysfunction. While cognitive deficits have been attributed to the side effects of chemotherapy, there is evidence for impairment at disease onset, prior to cancer-directed therapy. Further debated issues concern the relationship between self-reported complaints and objective test performance and the role of psychological distress. METHOD: We assessed performance on neuropsychological tests of attention and memory and obtained estimates of subjective distress and quality of life in 27 breast cancer patients and 20 healthy controls. Testing in patients took place shortly after the initial diagnosis, but prior to subsequent therapy. RESULTS: While patients showed elevated distress, cognitive performance differed on a few subtests only. Patients showed slower processing speed and poorer verbal memory than controls. Objective and self-reported cognitive function were unrelated, and psychological distress correlated more strongly with subjective complaints than with neuropsychological test performance. CONCLUSION: This study provides further evidence of limited cognitive deficits in cancer patients prior to the onset of adjuvant therapy. Self-reported cognitive deficits seem more closely related to psychological distress than to objective test performance.

Differential trajectories of memory quality and guessing across sequential reports from working memory.

Working memory enables the storage of few items for a short period of time. Previous research has shown that items in working memory cannot be accessed equally well, indicating that they are held in at least two different states with different capacity limitations. However, it is unclear whether differences between states are due to limitations of the number of items that can be stored, or the quality with which items are stored. We employed a sequential whole-report procedure where participants reported the remembered orientation of each of two or four encoded Gabor patches. In addition, they rated their memory confidence prior to each report. Participants performed 600 trials per condition, allowing us to obtain reliable subjective ratings and estimates of precision, guessing, and misreport using a mixture model, separately for each sequential report. Different measures of memory quality consistently showed discontinuous trajectories across reports with a steep drop from the first to the second remembered item but only slight decreases thereafter. In contrast, both reported and modeled guessing changed continuously across reports. Our results support the notion of two states in working memory and show that they are distinguished by memory quality rather than quantity.

Two types of serial dependence in visual working memory.

Stimulus representations in working memory depend on memory traces of past stimuli both from previous trials and from the current trial. However, it is unclear whether the same or different mechanisms underlie this serial dependence across and within trials. We directly contrasted estimates of bias for pairs of immediately successive stimuli across and within trials. In each trial, participants memorized two consecutive motion direction stimuli (S1 and S2) and after a short delay were cued to report one of them. We found serial dependence across trials: The current S1 was attracted towards the preceding S2 when the latter had been cued for report. In contrast, within the same trial S2 was repulsed from S1. In addition, repulsion within a trial occurred for a broader range of motion direction differences between stimuli than attraction across trials. A second experiment in which 25% of trials did not require a response demonstrated that across-trial attraction did not depend on whether the previous S2 actually had to be reported. Our findings provide evidence for two types of serial dependence operating across and within trials. They support the notion of different mechanisms integrating or segregating current from similar past memory contents depending on their task relevance.

Sequential whole report accesses different states in visual working memory.

Working memory (WM) enables a rapid access to a limited number of items that are no longer physically present. WM studies usually involve the encoding and retention of multiple items, while probing a single item only. Hence, little is known about how well multiple items can be reported from WM. Here we asked participants to successively report each of up to 8 encoded Gabor patches from WM. Recall order was externally cued, and stimulus orientations had to be reproduced on a continuous dimension. Participants were able to sequentially report items from WM with an above-chance precision even at high set sizes. It is important that we observed that precision varied systematically with report order: It dropped steeply from the first to the second report but decreased only slightly thereafter. The observed trajectory of precision decrease across reports was better captured as a discontinuous rather than an exponential function, suggesting that items were reported from different states in visual WM. The following 3 experiments replicated these findings. In particular, they showed that the observed drop could not be explained by a retro-cueing benefit of the first report, a longer delay duration for later reports or a visual interference effect of the first report. Instead, executive interference of the first report reduced precision of subsequent reports. Together, the results show that a sequential whole-report procedure allows the assessment of qualitatively different states in visual WM. (PsycINFO Database Record

Pre-encoding gamma-band activity during auditory working memory.

Previous magnetoencephalography (MEG) studies have revealed gamma-band activity at sensors over parietal and fronto-temporal cortex during the delay phase of auditory spatial and non-spatial match-to-sample tasks, respectively. While this activity was interpreted as reflecting the memory maintenance of sound features, we noted that task-related activation differences might have been present already prior to the onset of the sample stimulus. The present study focused on the interval between a visual cue indicating which sound feature was to be memorized (lateralization or pitch) and sample sound presentation to test for task-related activation differences preceding stimulus encoding. MEG spectral activity was analyzed with cluster randomization tests (N = 15). Whereas there were no differences in frequencies below 40 Hz, gamma-band spectral amplitude (about 50-65 and 90-100 Hz) was higher for the lateralization than the pitch task. This activity was localized at right posterior and central sensors and present for several hundred ms after task cue offset. Activity at 50-65 Hz was also increased throughout the delay phase for the lateralization compared with the pitch task. Apparently cortical networks related to auditory spatial processing were activated after participants had been informed about the task.

Superior Intraparietal Sulcus Controls the Variability of Visual Working Memory Precision.

UNLABELLED: Limitations of working memory (WM) capacity depend strongly on the cognitive resources that are available for maintaining WM contents in an activated state. Increasing the number of items to be maintained in WM was shown to reduce the precision of WM and to increase the variability of WM precision over time. Although WM precision was recently associated with neural codes particularly in early sensory cortex, we have so far no understanding of the neural bases underlying the variability of WM precision, and how WM precision is preserved under high load. To fill this gap, we combined human fMRI with computational modeling of behavioral performance in a delayed color-estimation WM task. Behavioral results replicate a reduction of WM precision and an increase of precision variability under high loads (5 > 3 > 1 colors). Load-dependent BOLD signals in primary visual cortex (V1) and superior intraparietal sulcus (IPS), measured during the WM task at 2-4 s after sample onset, were modulated by individual differences in load-related changes in the variability of WM precision. Although stronger load-related BOLD increase in superior IPS was related to lower increases in precision variability, thus stabilizing WM performance, the reverse was observed for V1. Finally, the detrimental effect of load on behavioral precision and precision variability was accompanied by a load-related decline in the accuracy of decoding the memory stimuli (colors) from left superior IPS. We suggest that the superior IPS may contribute to stabilizing visual WM performance by reducing the variability of memory precision in the face of higher load. SIGNIFICANCE STATEMENT: This study investigates the neural bases of capacity limitations in visual working memory by combining fMRI with cognitive modeling of behavioral performance, in human participants. It provides evidence that the superior intraparietal sulcus (IPS) is a critical brain region that influences the variability of visual working memory precision between and within individuals (Fougnie et al., 2012; van den Berg et al., 2012) under increased memory load, possibly in cooperation with perceptual systems of the occipital cortex. These findings substantially extend our understanding of the nature of capacity limitations in visual working memory and their neural bases. Our work underlines the importance of integrating cognitive modeling with univariate and multivariate methods in fMRI research, thus improving our knowledge of brain-behavior relationships.

Recurrence of task set-related MEG signal patterns during auditory working memory.

Processing of auditory spatial and non-spatial information in working memory has been shown to rely on separate cortical systems. While previous studies have demonstrated differences in spatial versus non-spatial processing from the encoding of to-be-remembered stimuli onwards, here we investigated whether such differences would be detectable already prior to presentation of the sample stimulus. We analyzed broad-band magnetoencephalography data from 15 healthy adults during an auditory working memory paradigm starting with a visual cue indicating the task-relevant stimulus feature for a given trial (lateralization or pitch) and a subsequent 1.5-s pre-encoding phase. This was followed by a sample sound (0.2s), the delay phase (0.8s) and a test stimulus (0.2s) after which participants made a match/non-match decision. Linear discriminant functions were trained to decode task-specific signal patterns throughout the task, and temporal generalization was used to assess whether the neural codes discriminating between the tasks during the pre-encoding phase would recur during later task periods. The spatial versus non-spatial tasks could indeed be discriminated after the onset of the cue onwards, and decoders trained during the pre-encoding phase successfully discriminated the tasks during both sample stimulus encoding and during the delay phase. This demonstrates that task-specific neural codes are established already before the memorandum is presented and that the same patterns are reestablished during stimulus encoding and maintenance. This article is part of a Special Issue entitled SI: Auditory working memory.

Activity in human visual and parietal cortex reveals object-based attention in working memory.

Visual attention enables observers to select behaviorally relevant information based on spatial locations, features, or objects. Attentional selection is not limited to physically present visual information, but can also operate on internal representations maintained in working memory (WM) in service of higher-order cognition. However, only little is known about whether attention to WM contents follows the same principles as attention to sensory stimuli. To address this question, we investigated in humans whether the typically observed effects of object-based attention in perception are also evident for object-based attentional selection of internal object representations in WM. In full accordance with effects in visual perception, the key behavioral and neuronal characteristics of object-based attention were observed in WM. Specifically, we found that reaction times were shorter when shifting attention to memory positions located on the currently attended object compared with equidistant positions on a different object. Furthermore, functional magnetic resonance imaging and multivariate pattern analysis of visuotopic activity in visual (areas V1-V4) and parietal cortex revealed that directing attention to one position of an object held in WM also enhanced brain activation for other positions on the same object, suggesting that attentional selection in WM activates the entire object. This study demonstrated that all characteristic features of object-based attention are present in WM and thus follows the same principles as in perception.

Neural correlates of chemotherapy-related cognitive impairment.

Cancer survivors frequently experience cognitive deficits following chemotherapy. The most commonly affected functions include memory, attention and executive control. The present paper reviews animal research and clinical studies including event-related potential (ERP) and neuroimaging investigations of chemotherapy-related changes of brain structure and function. In rodents, chemotherapeutic substances have been shown to damage neural precursor cells and white matter tracts and are associated with impairments of learning and memory. Structural and functional changes associated with chemotherapy have also been observed in humans. Structural imaging has revealed gray and white matter volume reductions and altered white matter microstructure. Functional studies using either ERPs or hemodynamic imaging have shown that chemotherapy alters the activation patterns of cortical networks involved in higher cognitive functions. Collectively, these findings support the existence of the "chemobrain" phenomenon beyond the patients' subjective reports. However, the rather small number of studies and methodological limitations of some of the pioneering investigations call for further research of high methodological quality, including larger numbers of subjects with appropriate controls to delineate the temporal and spatial pattern of chemotherapy-associated central nervous system (CNS) toxicity. Brain activation studies in humans might systematically vary task difficulty levels to distinguish between compensatory hyper-activations on the one hand and deficient recruitment of resources on the other hand. Integrative functions could be tested by connectivity analyses using both electrophysiological and hemodynamic measures. The ultimate goal should be the development of cognitive-behavioral and pharmacological interventions to reduce the cognitive side effects of the medically indispensable but neurotoxic chemotherapeutic treatments.

fMRI characterization of visual working memory recognition.

Encoding and maintenance of information in visual working memory have been extensively studied, highlighting the crucial and capacity-limiting role of fronto-parietal regions. In contrast, the neural basis of recognition in visual working memory has remained largely unspecified. Cognitive models suggest that recognition relies on a matching process that compares sensory information with the mental representations held in memory. To characterize the neural basis of recognition we varied both the need for recognition and the degree of similarity between the probe item and the memory contents, while independently manipulating memory load to produce load-related fronto-parietal activations. fMRI revealed a fractionation of working memory functions across four distributed networks. First, fronto-parietal regions were activated independent of the need for recognition. Second, anterior parts of load-related parietal regions contributed to recognition but their activations were independent of the difficulty of matching in terms of sample-probe similarity. These results argue against a key role of the fronto-parietal attention network in recognition. Rather the third group of regions including bilateral temporo-parietal junction, posterior cingulate cortex and superior frontal sulcus reflected demands on matching both in terms of sample-probe-similarity and the number of items to be compared. Also, fourth, bilateral motor regions and right superior parietal cortex showed higher activation when matching provided clear evidence for a decision. Together, the segregation between the well-known fronto-parietal activations attributed to attentional operations in working memory from those regions involved in matching supports the theoretical view of separable attentional and mnemonic contributions to working memory. Yet, the close theoretical and empirical correspondence to perceptual decision making may call for an explicit consideration of decision making mechanisms in conceptions of working memory.

Separable neural bases for subprocesses of recognition in working memory.

Working memory supports the recognition of objects in the environment. Memory models have postulated that recognition relies on 2 processes: assessing the degree of similarity between an external stimulus and memory representations and testing the resulting summed-similarity value against a critical level for recognition. Here, we varied the similarity between samples held in working memory and a probe to investigate these 2 processes with magnetoencephalography. Two separable components matched our expectations: First, from 280 ms after probe onset, clearly nonmatching probes differed from both similar nonmatches and matches over left frontal cortex. At 350-400 ms, these signals evolved into a pattern of gradually increasing activation as a function of sample-probe similarity, as expected for a neural representation of summed similarity. Second, a signal potentially reflecting criterion testing was observed at 600-700 ms at right frontotemporal sensors that differentiated between matches and nonmatches without further differences between similar and dissimilar probes. Thus, analysis of the time course of recognition provided strong evidence that similarity summation and criterion testing have separable neural bases. As probably both working and long-term memory recognition draw on these processes, they may be involved in many domains of behavior.

Basic operations in working memory: contributions from functional imaging studies.

Working memory (WM) constitutes a fundamental aspect of human cognition. It refers to the ability to keep information active for further use, while allowing it to be prioritized, modified and protected from interference. Much research has addressed the storage function of WM, however, its 'working' aspect still remains underspecified. Many operations that work on the contents of WM do not appear specific to WM. The present review focuses on those operations that we consider "basic" because they operate in the service of memory itself, by providing its basic functionality of retaining information active, in a stable yet flexible way. Based on current process models of WM we review five strands of research: (1) mnemonic selection of one item amongst others, (2) updating the focus of attention with the selected item, (3) updating the content of visual WM with new item(s), (4) rehearsal of visuospatial information and (5) coping with interference. We discuss the neuronal substrates underlying those operations obtained with functional magnetic resonance imaging and relate them to findings on "executive functions". The presented data support the view that WM emerges from interactions between higher sensory, attentional and mnemonic functions, with separable neural bases. However, interference processing and the representation of rule switching in WM may demand an extension of the current WM models by executive control functions.

What "works" in working memory? Separate systems for selection and updating of critical information.

Cognition depends critically on working memory, the active representation of a limited number of items over short periods of time. In addition to the maintenance of information during the course of cognitive processing, many tasks require that some of the items in working memory become transiently more important than others. Based on cognitive models of working memory, we hypothesized two complementary essential cognitive operations to achieve this: a selection operation that retrieves the most relevant item, and an updating operation that changes the focus of attention onto it. Using functional magnetic resonance imaging, high-resolution oculometry, and behavioral analysis, we demonstrate that these two operations are functionally and neuroanatomically dissociated. Updating the attentional focus elicited transient activation in the caudal superior frontal sulcus and posterior parietal cortex. In contrast, increasing demands on selection selectively modulated activation in rostral superior frontal sulcus and posterior cingulate/precuneus. We conclude that prioritizing one memory item over others invokes independent mechanisms of mnemonic retrieval and attentional focusing, each with its distinct neuroanatomical basis within frontal and parietal regions. These support the developing understanding of working memory as emerging from the interaction between memory and attentional systems.