Encoded and updated spatial working memories share a common representational format in alpha activity.

Working memory (WM) flexibly updates information to adapt to the dynamic environment. Here, we used alpha-band activity in the EEG to reconstruct the content of dynamic WM updates and compared this representational format to static WM content. An inverted encoding model using alpha activity precisely tracked both the initially encoded position and the updated position following an auditory cue signaling mental updating. The timing of the update, as tracked in the EEG, correlated with reaction times and saccade latency. Finally, cross-training analyses revealed a robust generalization of alpha-band reconstruction of WM contents before and after updating. These findings demonstrate that alpha activity tracks the dynamic updates to spatial WM and that the format of this activity is preserved across the encoded and updated representations. Thus, our results highlight a new approach for measuring updates to WM and show common representational formats during dynamic mental updating and static storage.

Prediction error and event segmentation in episodic memory.

Organizing the continuous flow of experiences into meaningful events is a crucial prerequisite for episodic memory. Prediction error and event segmentation both play important roles in supporting the genesis of meaningful mnemonic representations of events. We review theoretical contributions discussing the relationship between prediction error and event segmentation, as well as literature on episodic memory related to prediction error and event segmentation. We discuss the extent of overlap of mechanisms underlying memory emergence through prediction error and event segmentation, with a specific focus on attention and working memory. Finally, we identify areas in research that are currently developing and suggest future directions. We provide an overview of mechanisms underlying memory formation through predictions, violations of predictions, and event segmentation.

The reactivation of task rules triggers the reactivation of task-relevant items.

Working memory (WM) describes the temporary storage of task-relevant items and procedural rules to guide action. Despite its central importance for goal-directed behavior, the interplay between WM and long-term memory (LTM) remains poorly understood. Recent studies have shown that repeated use of the same task-relevant item in WM results in a hand-off of the storage of that item to LTM, and switching to a new item reactivates WM. To further elucidate the rules governing WM-LTM interactions, we here planned to probe whether a change in task rules, independent of a switch in task-relevant items, would also lead to WM reactivation of maintained items. To this end, we used scalp-recorded electroencephalogram (EEG) data, specifically the contralateral delay activity (CDA), to track WM item storage while manipulating repetitions and changes in task rules and task-relevant items across trials in a visual WM task. We tested two rival hypotheses: If changes in task rules result in a reactivation of the target item representation, then the CDA should increase when a task change is cued even when the same target has been repeated across trials. However, if the reactivation of a task-relevant item only depends on the mnemonic availability of the item itself instead of the task it is used for, then only the changes in task-relevant items should reactivate the representations. Accordingly, the CDA amplitude should decrease for repeated task-relevant items independently of a task change. We found a larger CDA on task-switch compared to task-repeat trials, suggesting that the reactivation of task rules triggers the reactivation of task-relevant items in WM. By demonstrating that WM reactivation of LTM is interdependent for task rules and task-relevant items, this study informs our understanding of visual WM and its interplay with LTM. PREREGISTERED STAGE 1 PROTOCOL: https://osf.io/zp9e8 (date of in-principle acceptance: 19/12/2021).

Internal attention is the only retroactive mechanism for controlling precision in working memory.

Recent research has suggested that humans can assert control over the precision of working memory (WM) items. However, the mechanisms that enable this control are unclear. While some studies suggest that internal attention improves precision, it may not be the only factor, as previous work also demonstrated that WM storage is disentangled from attention. To test whether there is a precision control mechanism beyond internal attention, we contrasted internal attention and precision requirements within the same trial in three experiments. In every trial, participants memorized two items briefly. Before the test, a retro-cue indicated which item would be tested first, thus should be attended. Importantly, we encouraged participants to store the unattended item with higher precision by testing it using more similar lure colors at the probe display. Accuracy was analyzed on a small proportion of trials where the target-lure similarity, hence the task difficulty, was equal for attended and unattended items. Experiments 2 and 3 controlled for output interference by the first test and involuntary precision boost by the retro-cue, respectively. In all experiments, the unattended item had lower accuracy than the attended item, suggesting that individuals were not able to remember it more precisely than the attended item. Thus, we conclude that there is no precision control mechanism beyond internal attention, highlighting the close relationship between attentional and qualitative prioritization within WM. We discuss the important implications of these findings for our understanding of the fundamentals of WM and WM-driven behaviors.

Preparation for upcoming attentional states in the hippocampus and medial prefrontal cortex.

Goal-directed attention is usually studied by providing individuals with explicit instructions on what they should attend to. But in daily life, we often use past experiences to guide our attentional states. Given the importance of memory for predicting upcoming events, we hypothesized that memory-guided attention is supported by neural preparation for anticipated attentional states. We examined preparatory coding in the human hippocampus and mPFC, two regions that are important for memory-guided behaviors, in two tasks: one where attention was guided by memory and another in which attention was explicitly instructed. Hippocampus and mPFC exhibited higher activity for memory-guided vs. explicitly instructed attention. Furthermore, representations in both regions contained information about upcoming attentional states. In the hippocampus, this preparation was stronger for memory-guided attention, and occurred alongside stronger coupling with visual cortex during attentional guidance. These results highlight the mechanisms by which memories are used to prepare for upcoming attentional goals.

At any given moment, humans are bombarded with a constant stream of new information. But the brain can take in only a fraction of that information at once. So how does the brain decide what to pay attention to and what to ignore? Many laboratory studies of attention avoid this issue by simply telling participants what to attend to. But in daily life, people rarely receive instructions like that. Instead people must often rely on past experiences to guide their attention. When cycling close to home, for example, a person knows to watch out for the blind junction at the top of the hill and for the large pothole just around the corner. Günseli and Aly set out to bridge the gap between laboratory studies of attention and real-world experience by asking healthy volunteers to perform two versions of a task while lying inside a brain scanner. The task involved looking at pictures of rooms with different shapes. Each room also contained a different painting. In one version of the task, the volunteers were told to pay attention to either the paintings or to the room shapes. In the other version, the volunteers had to use previously memorized cues to work out for themselves whether they should focus on the paintings or on the shapes. The brain scans showed that two areas of the brain with roles in memory ­ the hippocampus and the prefrontal cortex ­ were involved in the task. Notably, both areas increased their activity when the volunteers used memory to guide their attention, compared to when they received instructions telling them what to focus on. Moreover, patterns of activity within the hippocampus and prefrontal cortex contained information about what the participants were about to focus on next ­ even before volunteers saw the particular picture that they were supposed to pay attention to. In the hippocampus, this was particularly the case when the volunteers based their decisions on memory. These results reveal a key way in which humans leverage memories of past experiences to help optimize future behavior. Understanding this process could shed light on why memory impairments make it harder for people to adjust their behavior to achieve specific goals.

Dynamic internal states shape memory retrieval.

Why do we sometimes easily retrieve memories, but other times appear to forget them? We often look to our external environment for retrieval cues, but another way to optimize memory retrieval is to be in a mental state, or mode, that prioritizes access to our internal representation of the world. Such a 'retrieval mode' was proposed by Endel Tulving (1983), who considered it a neurocognitive state in which one keeps the goal of memory retrieval in mind. Building on Tulving's proposal, we review converging evidence from multiple lines of research that emphasize the importance of internal states in the instantiation of retrieval modes that optimize successful remembering. We identify three key factors that contribute to a retrieval mode by modulating either the likelihood or the content of retrieval: (1) an intention to remember or forget (either in the present or the future), (2) attentional selection of goal-relevant memories and suppression of distractors, and (3) fluctuating levels of acetylcholine in the hippocampus. We discuss empirical evidence that these internal states individually influence memory retrieval and propose how they may interact synergistically. Characterizing these dynamic internal factors is an important key for unlocking our understanding of the organization and accessibility of our memories.

EEG dynamics reveal a dissociation between storage and selective attention within working memory.

Selective attention plays a prominent role in prioritizing information in working memory (WM), improving performance for attended representations. However, it remains unclear whether unattended WM representations suffer from information loss. Here we tested the hypothesis that within WM, selectively attending to an item and stopping storing other items are independent mechanisms. We recorded EEG while participants performed a WM recall task in which the item most likely to be tested was cued retrospectively during retention. By manipulating retro-cue reliability (i.e., the ratio of valid to invalid cue trials), we varied the incentive to retain non-cued items. Storage and selective attention in WM were measured during the retention interval by contralateral delay activity (CDA) and contralateral alpha power suppression, respectively. Soon after highly reliable cues, the cued item was attended, and non-cued items suffered information loss. However, for less reliable cues, initially the cued item was attended, but unattended items were kept in WM. Later during the delay, previously unattended items suffered information loss despite now attention being reallocated to their locations, presumably to strengthen their weakening traces. These results show that storage and attention in WM are distinct processes that can behave differently depending on the relative importance of representations.

Dissecting the Neural Focus of Attention Reveals Distinct Processes for Spatial Attention and Object-Based Storage in Visual Working Memory.

Complex cognition relies on both on-line representations in working memory (WM), said to reside in the focus of attention, and passive off-line representations of related information. Here, we dissected the focus of attention by showing that distinct neural signals index the on-line storage of objects and sustained spatial attention. We recorded electroencephalogram (EEG) activity during two tasks that employed identical stimulus displays but varied the relative demands for object storage and spatial attention. We found distinct delay-period signatures for an attention task (which required only spatial attention) and a WM task (which invoked both spatial attention and object storage). Although both tasks required active maintenance of spatial information, only the WM task elicited robust contralateral delay activity that was sensitive to mnemonic load. Thus, we argue that the focus of attention is maintained via a collaboration between distinct processes for covert spatial orienting and object-based storage.

Local and interregional alpha EEG dynamics dissociate between memory for search and memory for recognition.

Attention during visual search is thought to be guided by an active visual working memory (VWM) representation of the search target. We tested the hypothesis that a VWM representation used for searching a target among competing information (a "search template") is distinct from VWM representations used for simple recognition tasks, without competition. We analyzed EEG from 20 human participants while they performed three different VWM-based visual detection tasks. All tasks started with identical lateralized VWM cues, but differed with respect to the presence and nature of competing distractors during the target display at test, where participants performed a simple recognition task without distractors, or visual search in pop-out (distinct) and serial (non-distinct) search displays. Performance was worst for non-distinct search, and best for simple recognition. During the one second delay period between cue and test, we observed robust suppression of EEG dynamics in the alpha (8-14Hz) band over parieto-occipital sites contralateral to the relevant VWM item, both in terms of local power as well as interregional phase synchrony within a posterior-parietal network. Importantly, these lateralization dynamics were more strongly expressed prior to search compared to simple recognition. Furthermore, before the VWM cue, alpha phase synchrony between prefrontal and mid-posterior-parietal sites was strongest for non-distinct search, reflecting enhanced anticipatory control prior to VWM encoding. Directional connectivity analyses confirmed this effect to be in an anterior-to-posterior direction. Together, these results provide evidence for frontally mediated top-down control of VWM in preparation of visual search.

Task-irrelevant memories rapidly gain attentional control with learning.

Although many of our perceptual biases stem from long-term, repeated exposure, current theories of visual search assume a central role for visual working memory (VWM) in guiding attention to target information. Crucially, whether a VWM representation guides attention depends on the relative priority that the memory has within VWM. Here, in a combined visual search/VWM task, we used attentional guidance by irrelevant memories to measure how long a target representation remains prioritized in VWM when observers repeatedly search for the same target. Irrelevant memories started guiding attention already when the target was repeated once, indicating that the target representation rapidly lost priority within VWM as it moved to long-term memory. By showing that training can lead to interference from irrelevant memories, the findings resolve a long-standing paradox on why VWM appears central to, yet at the same time not sufficient nor necessary for attentional guidance.

The reliability of retro-cues determines the fate of noncued visual working memory representations.

Retrospectively cueing an item retained in visual working memory during maintenance is known to improve its retention. However, studies have provided conflicting results regarding the costs of such retro-cues for the noncued items, leading to different theories on the mechanisms behind visual working memory maintenance and retro-cueing. Here we tested an alternative explanation of the conflicting results regarding retro-cue costs-namely, that they are caused at least partly by differences in retro-cue reliability. We manipulated the ratio of valid-cue trials to invalid-cue trials within blocks. We used a continuous-report procedure that allowed fitting a model that provided recall probability and precision estimates for the memory representations. Reconciling previous contradictory findings, benefits for valid cues were observed in all conditions, but invalid cueing costs were found only when the retro-cue had a high reliability (i.e., was 80 % valid), but not when it had a lower reliability (i.e., 50 % valid). This was found for both the recall probability and the precision of visual working memory representations. Our results suggest that the cognitive mechanisms underlying retro-cue effects are strategically adjusted by participants, depending on the perceived retro-cue reliability.

Is a search template an ordinary working memory? Comparing electrophysiological markers of working memory maintenance for visual search and recognition.

Visual search requires the maintenance of a search template in visual working memory in order to guide attention towards the target. This raises the question whether a search template is essentially the same as a visual working memory representation used in tasks that do not require attentional guidance, or whether it is a qualitatively different representation. Two experiments tested this by comparing electrophysiological markers of visual working memory maintenance between simple recognition and search tasks. For both experiments, responses were less rapid and less accurate in search task than in simple recognition. Nevertheless, the contralateral delay activity (CDA), an index of quantity and quality of visual working memory representations, was equal across tasks. On the other hand, the late positive complex (LPC), which is sensitive to the effort invested in visual working memory maintenance, was greater for the search task than the recognition task. Additionally, when the same target cue was repeated across trials (Experiment 2), the amplitude of visual working memory markers (both CDA and LPC) decreased, demonstrating learning of the target at an equal rate for both tasks. Our results suggest that a search template is qualitatively the same as a representation used for simple recognition, but greater effort is invested in its maintenance.

Effects of search difficulty on the selection, maintenance, and learning of attentional templates.

Prominent theories of attention claim that visual search is guided through attentional templates stored in working memory. Recently, the contralateral delay activity (CDA), an electrophysiological index of working memory storage, has been found to rapidly decrease when participants repeatedly search for the same target, suggesting that, with learning, the template moves out of working memory. However, this has only been investigated with pop-out search for distinct targets, for which a strong attentional template may not be necessary. More effortful search tasks might rely more on an active attentional template in working memory, leading to a slower handoff to long-term memory and thus a slower decline of the CDA. Using ERPs, we compared the rate of learning of attentional templates in pop-out and effortful search tasks. In two experiments, the rate of decrease in the CDA was the same for both search tasks. Similar results were found for a second component indexing working memory effort, the late positive complex. However, the late positive complex was also sensitive to anticipated search difficulty, as was expressed in a greater amplitude before the harder search task. We conclude that the amount of working memory effort invested in maintaining an attentional template, but not the rate of learning, depends on search difficulty.

The time course of protecting a visual memory representation from perceptual interference.

Cueing a remembered item during the delay of a visual memory task leads to enhanced recall of the cued item compared to when an item is not cued. This cueing benefit has been proposed to reflect attention within visual memory being shifted from a distributed mode to a focused mode, thus protecting the cued item against perceptual interference. Here we investigated the dynamics of building up this mnemonic protection against visual interference by systematically varying the stimulus onset asynchrony (SOA) between cue onset and a subsequent visual mask in an orientation memory task. Experiment 1 showed that a cue counteracted the deteriorating effect of pattern masks. Experiment 2 demonstrated that building up this protection is a continuous process that is completed in approximately half a second after cue onset. The similarities between shifting attention in perceptual and remembered space are discussed.