Investigating the effects of perceptual complexity versus conceptual meaning on the object benefit in visual working memory.

Previous research has demonstrated greater visual working memory (VWM) performance for real-world objects compared with simple features. Greater amplitudes of the contralateral delay activity (CDA)-a sustained event-related potential measured during the delay period of a VWM task-have also been noted for meaningful stimuli, despite being thought of as a neural marker of a fixed working memory capacity. The current study aimed to elucidate the factors underlying improved memory performance for real-world objects by isolating the relative contributions of perceptual complexity (i.e., number of visual features) and conceptual meaning (i.e., availability of semantic, meaningful features). Participants (N = 22) performed a lateralized VWM task to test their memory of intact real-world objects, scrambled real-world objects and colours. The CDA was measured during both encoding and WM retention intervals (600-1000 ms and 1300-1700 ms poststimulus onset, respectively), and behavioural performance was estimated by using d' (memory strength in a two-alternative forced choice task). Behavioural results revealed significantly better performance within-subjects for real-world objects relative to scrambled objects and colours, with no difference between colours and scrambled objects. The amplitude of the CDA was also largest for intact real-world objects, with no difference in magnitude for scrambled objects and colours, during working memory maintenance. However, during memory encoding, both the colours and intact real-world objects had significantly greater amplitudes than scrambled objects and were comparable in magnitude. Overall, findings suggest that conceptual meaning (semantics) supports the memory benefit for real-world objects.

Limitations on flexible allocation of visual short-term memory resources with multiple levels of goal-directed attentional prioritization.

Studies suggest that visual short-term memory (VSTM) is a continuous resource that can be flexibly allocated using probabilistic cues that indicate test likelihood (i.e., goal-directed attentional priority to those items). Previous studies using simultaneous cues have not examined this flexible allocation beyond two distinct levels of priority. Moreover, previous studies have not examined whether there are individual differences in the ability to flexibly allocate VSTM resources, as well as whether this ability benefits from practice. The current study used a continuous report procedure to examine whether participants can use up to three levels of attentional priority to allocate VSTM resources via simultaneous probabilistic spatial cues. Three experiments were performed with differing priority levels, cues, and cue presentation times. Group level analysis demonstrated flexible allocation of VSTM resources; however, there was limited evidence that participants could use three goal-directed priority levels. A temporal analysis suggested that task fatigue, rather than practice effects, may interact with item priority. A Bayesian individual-differences analysis revealed that a minority of participants were using three levels of attentional priority, demonstrating that, while possible, it is not the predominant pattern of behaviour. Thus, we provided evidence that flexible allocation to three attention levels is possible under simultaneous cuing conditions for a minority of participants. Flexible allocation to three categories may be interpreted as a skill of high-performing participants akin to high memory capacity.

Age differences in the use of positive and negative cues to filter distracting information from working memory.

Previous research has demonstrated that as people age, visual working memory (VWM) declines. One potential explanation for this decline is that older adults are less able to ignore irrelevant information, which contributes to VWM filtering deficits. Most research examining age differences in filtering ability has used positive cues (indicating which items to pay attention to), but negative cues (indicating which items to ignore) may be even harder for older adults to implement as some work suggests that negatively cued items are first paid attention to before they are suppressed. The current study aimed to test whether older adults can use negative cues to filter irrelevant information from VWM. Across two experiments, young and older adults were presented with two (Experiment 1) or four (Experiment 2) display items, preceded by a neutral, negative, or positive cue. After a delay, participants reported the target's orientation in a continuous-response task. Results show that both groups benefitted from being provided with a cue (positive or negative) compared to no cue (i.e., neutral condition), but the benefit was smaller for negative cues. Thus, although negative cues aid in filtering of VWM, they are less effective than positive cues, possibly due to residual attention being directed towards distractor items.

Retrospective cue benefits in visual working memory are limited to a single location at a time.

Working memory (WM) performance can be improved by an informative cue presented during storage. This effect, termed a retrocue benefit, can be used to study limits on how human observers prioritize information stored in WM for behavioral output. There is disagreement about whether retrocue benefits extend to multiple WM locations. Here, we hypothesized that multiple retrocues may improve some aspects of memory performance (e.g., a reduction in random guessing) while worsening others (e.g., an increase in the probability of reporting a feature presented at a non-probed location). We tested this possibility in three experiments. Participants remembered arrays of four orientations or colors over a brief delay, and spatial retrocues instructed participants to prioritize zero, one, two, or all four remembered orientations for possible report. At the end of the trial, participants recalled the orientation that appeared at one location. The results of this study revealed that participants' recall errors were lower during cue-one relative to cue-two and cue-four trials, and this benefit was driven primarily by a reduction in random guessing during cue-one trials. We found no evidence suggesting that multiple spatial cues (i.e., during cue-two trials) induced a trade-off between memory precision, random guessing, and non-target reports compared to neutral trials (i.e., cue-zero or cue-four). Thus, cuing participants to prioritize information appearing at multiple unique spatial positions led to no improvement in memory performance compared to neutral or no-cue trials, providing additional support for the view that retrocue benefits on WM performance are limited to a single spatial location at a time.

Sensory Delay Activity: More than an Electrophysiological Index of Working Memory Load.

Sustained contralateral delay activity emerges in the retention period of working memory (WM) tasks and has been commonly interpreted as an electrophysiological index of the number of items held in a discrete-capacity WM resource. More recent findings indicate that these visual and tactile components are sensitive to various cognitive operations beyond the storage of discrete items in WM. In this Perspective, we present recent evidence from unisensory and multisensory visual and tactile WM tasks suggesting that, in addition to memory load, sensory delay activity may also be indicative of attentional and executive processes, as well as reflecting the flexible, rather than discrete, allocation of a continuous WM resource. Together, these findings challenge the traditional model of the functional significance of the contralateral delay activity as a pure measure of item load, and suggest that it may also reflect executive, attentional, and perceptual mechanisms operating in hierarchically organized WM systems.

Drawn to Distraction: Anxiety Impairs Neural Suppression of Known Distractor Features in Visual Search.

When searching for a target, it is possible to suppress the features of a known distractor. This suppression may prevent distractor processing altogether or only after the distractor initially captures attention (i.e., search and destroy). However, suppression may be impaired in individuals with attentional control deficits, such as in high anxiety. In this study (n = 48), we used ERPs to examine the time course of attentional enhancement and suppression when participants were given pretrial information about target or distractor features. Consistent with our hypothesis, we found that individuals with higher levels of anxiety had lower neural measures of suppressing the template-matching distractor, instead showing enhanced processing. These findings indicate that individuals with anxiety are more likely to use a search-and-destroy mechanism of negative templates-highlighting the importance of attentional control abilities in distractor-guided search.

Fear not! Anxiety biases attentional enhancement of threat without impairing working memory filtering.

Individuals with anxiety have attentional biases toward threat-related distractors. This deficit in attentional control has been shown to impact visual working memory (VWM) filtering efficiency, as anxious individuals inappropriately store threatening distractors in VWM. It remains unclear, however, whether this mis-allocation of memory resources is due to inappropriate attentional enhancement of threatening distractors, or to a failure in suppression. Here, we used a systematically lateralized VWM task with fearful and neutral faces to examine event-related potentials related to attentional selection (N2pc), suppression (PD), and working memory maintenance (CDA). We found that state anxiety correlated with attentional enhancement of threat-related distractors, such that more anxious individuals had larger N2pc amplitudes toward fearful distractors than neutral distractors. However, there was no correlation between anxiety and memory storage of fearful distractors (CDA). These findings demonstrate that anxiety biases attention toward fearful distractors, but that this bias does not always guarantee increased memory storage of threat-related distractors.

Reduced Attentional Control in Older Adults Leads to Deficits in Flexible Prioritization of Visual Working Memory.

Visual working memory (VWM) resources have been shown to be flexibly distributed according to item priority. This flexible allocation of resources may depend on attentional control, an executive function known to decline with age. In this study, we sought to determine how age differences in attentional control affect VWM performance when attention is flexibly allocated amongst targets of varying priority. Participants performed a delayed-recall task wherein item priority was varied. Error was modelled using a three-component mixture model to probe different aspects of performance (precision, guess-rate, and non-target errors). The flexible resource model offered a good fit to the data from both age groups, but older adults showed consistently lower precision and higher guess rates. Importantly, when demands on flexible resource allocation were highest, older adults showed more non-target errors, often swapping in the item that had a higher priority at encoding. Taken together, these results suggest that the ability to flexibly allocate attention in VWM is largely maintained with age, but older adults are less precise overall and sometimes swap in salient, but no longer relevant, items possibly due to their lessened ability to inhibit previously attended information.

Electrophysiological correlates of the flexible allocation of visual working memory resources.

Visual working memory is a brief, capacity-limited store of visual information that is involved in a large number of cognitive functions. To guide one's behavior effectively, one must efficiently allocate these limited memory resources across memory items. Previous research has suggested that items are either stored in memory or completely blocked from memory access. However, recent behavioral work proposes that memory resources can be flexibly split across items based on their level of task importance. Here, we investigated the electrophysiological correlates of flexible resource allocation by manipulating the distribution of resources amongst systematically lateralized memory items. We examined the contralateral delay activity (CDA), a waveform typically associated with the number of items held in memory. Across three experiments, we found that, in addition to memory load, the CDA flexibly tracks memory resource allocation. This allocation occurred as early as attentional selection, as indicated by the N2pc. Additionally, CDA amplitude was better-described when fit with a continuous model predicted by load and resources together than when fit with either alone. Our findings show that electrophysiological markers of attentional selection and memory maintenance not only track memory load, but also the proportion of memory resources those items receive.

A daytime nap enhances visual working memory performance and alters event-related delay activity.

Working memory (WM) is impaired following sleep loss and may be improved after a nap. The goal of the current study was to better understand sleep-related WM enhancement by: (1) employing a WM task that assesses the ability to hold and report visual representations as well as the fidelity of the reports on a fine scale, (2) investigating neurophysiological properties of sleep and WM capacity as potential predictors or moderators of sleep-related enhancement, and (3) exploring frontal and occipital event-related delay activity to index the neural processing of stimuli in WM. In a within-subjects design, 36 young adults (Mage = 20, 20 men, 16 women) completed a 300-trial, continuous-report task of visual WM following a 90-min nap opportunity and an equivalent period of wakefulness. Mixed-effect models were used to estimate the odds of successful WM reports and the fidelity of those reports. The odds of a successful report were approximately equal between nap and wake conditions for the start of the task, but by the end, the odds of success were 1.26 times greater in the nap condition. Successful WM reports were more accurate after a nap, independent of the time on task. Neither WM capacity nor any of the sleep variables measured were found to significantly moderate the nap effect on WM. Lastly, napping resulted in amplitude changes for frontal and occipital delay activity relative to the wake condition. The findings are discussed in relation to contemporary models of visual WM and the role of sleep in sustained attention.

Encoding differences affect the number and precision of own-race versus other-race faces stored in visual working memory.

Other-race faces are discriminated and recognized less accurately than own-race faces. Despite a wealth of research characterizing this other-race effect (ORE), little is known about the nature of the representations of own-race versus other-race faces. This is because traditional measures of this ORE provide a binary measure of discrimination or recognition (correct/incorrect), failing to capture potential variation in the quality of face representations. We applied a novel continuous-response paradigm to independently measure the number of own-race and other-race face representations stored in visual working memory (VWM) and the precision with which they are stored. Participants reported target own-race or other-race faces on a circular face space that smoothly varied along the dimension of identity. Using probabilistic mixture modeling, we found that following ample encoding time, the ORE is attributable to differences in the probability of a face being maintained in VWM. Reducing encoding time, a manipulation that is more sensitive to encoding limitations, caused a loss of precision or an increase in variability of VWM for other-race but not own-race faces. These results suggest that the ORE is driven by the inefficiency with which other-race faces are rapidly encoded in VWM and provide novel insights about how perceptual experience influences the representation of own-race and other-race faces in VWM.

ERP evidence for temporal independence of set size and object updating in object substitution masking.

To keep track of dynamically changing objects in one's environment, it is necessary to individuate them from other objects, both temporally and spatially. Spatially, objects can be selected from nearby distractors using selective attention. Temporally, object updating processes incorporate new information into existing representations over time. Both of these processes have been implicated in a type of visual masking called object-substitution masking (OSM). Previous studies have found that the number of distractors (impacting selective attention) interacts with the strength of OSM. However, it has been suggested that this interaction is an artifact of ceiling performance at low set sizes, rather than necessitating a failure of attention during masking. Using event-related potentials (ERPs), we examined whether set size and masking interact as measured by markers of selective attention (N2pc) and visual working memory consolidation/maintenance (SPCN). Set size was found to affect the N2pc (200-350 ms) and late SPCN (500-650 ms), reflecting increased demands on selective attention and unnecessary storage respectively. An early window of the SPCN (350-500 ms) was affected by masking, suggesting that OSM influences object consolidation processes in this window, independent of the number of distractors. Overall, it was found that selective attention and visual awareness are dissociable neural processes in OSM, and that they are independently affected by set size and masking manipulations. Moreover, we found that the early SPCN may reflect disruptions to object consolidation, potentially revealing a neural mechanism supporting an object individuation-through-updating account of OSM.

More than a filter: Feature-based attention regulates the distribution of visual working memory resources.

Across 2 experiments we revisited the filter account of how feature-based attention regulates visual working memory (VWM). Originally drawing from discrete-capacity ("slot") models, the filter account proposes that attention operates like the "bouncer in the brain," preventing distracting information from being encoded so that VWM resources are reserved for relevant information. Given recent challenges to the assumptions of discrete-capacity models, we investigated whether feature-based attention plays a broader role in regulating memory. Both experiments used partial report tasks in which participants memorized the colors of circle and square stimuli, and we provided a feature-based goal by manipulating the likelihood that 1 shape would be probed over the other across a range of probabilities. By decomposing participants' responses using mixture and variable-precision models, we estimated the contributions of guesses, nontarget responses, and imprecise memory representations to their errors. Consistent with the filter account, participants were less likely to guess when the probed memory item matched the feature-based goal. Interestingly, this effect varied with goal strength, even across high probabilities where goal-matching information should always be prioritized, demonstrating strategic control over filter strength. Beyond this effect of attention on which stimuli were encoded, we also observed effects on how they were encoded: Estimates of both memory precision and nontarget errors varied continuously with feature-based attention. The results offer support for an extension to the filter account, where feature-based attention dynamically regulates the distribution of resources within working memory so that the most relevant items are encoded with the greatest precision. (PsycINFO Database Record

Attention mediates the flexible allocation of visual working memory resources.

Though it is clear that it is impossible to store an unlimited amount of information in visual working memory (VWM), the limiting mechanisms remain elusive. While several models of VWM limitations exist, these typically characterize changes in performance as a function of the number of to-be-remembered items. Here, we examine whether changes in spatial attention could better account for VWM performance, independent of load. Across 2 experiments, performance was better predicted by the prioritization of memory items (i.e., attention) than by the number of items to be remembered (i.e., memory load). This relationship followed a power law, and held regardless of whether performance was assessed based on overall precision or any of 3 measures in a mixture model. Moreover, at large set sizes, even minimally attended items could receive a small proportion of resources, without any evidence for a discrete-capacity on the number of items that could be maintained in VWM. Finally, the observed data were best fit by a variable-precision model in which response error was related to the proportion of resources allocated to each item, consistent with a model of VWM in which performance is determined by the continuous allocation of attentional resources during encoding. (PsycINFO Database Record

Within-Category Decoding of Information in Different Attentional States in Short-Term Memory.

A long-standing assumption of cognitive neuroscience has been that working memory (WM) is accomplished by sustained, elevated neural activity. More recently, theories of WM have expanded this view by describing different attentional states in WM with differing activation levels. Several studies have used multivariate pattern analysis (MVPA) of functional magnetic resonance imaging (fMRI) and electroencephalography (EEG) data to study neural activity corresponding to these WM states. Intriguingly, no evidence was found for active neural representations for information held in WM outside the focus of attention ("unattended memory items," UMIs), suggesting that only attended memory items (AMIs) are accompanied by an active trace. However, these results depended on category-level decoding, which lacks sensitivity to neural representations of individual items. Therefore, we employed a WM task in which subjects remembered the directions of motion of two dot arrays, with a retrocue indicating which was relevant for an imminent memory probe (the AMI). This design allowed MVPA decoding of delay-period fMRI signal at the stimulus-item level, affording a more sensitive test of the neural representation of UMIs. Whereas evidence for the AMI was reliably high, evidence for the UMI dropped to baseline, consistent with the notion that different WM attentional states may have qualitatively different mechanisms of retention.

Comparing the Effects of 10-Hz Repetitive TMS on Tasks of Visual STM and Attention.

Numerous studies have demonstrated that visual STM (VSTM) and attention are tightly linked processes that share a number of neuroanatomical substrates. Here, we used repetitive TMS (rTMS) along with simultaneous EEG to examine the causal relationship between intraparietal sulcus functioning and performance on tasks of attention and VSTM. Participants performed two tasks in which they were required to attend to or remember colored items over a brief interval, with 10-Hz rTMS applied on some of the trials. Although no overall behavioral changes were observed across either task, rTMS did affect individual performance on both the attention and VSTM tasks in a manner that was predicted by individual differences in baseline performance. Furthermore, rTMS also affected ongoing oscillations in the alpha and beta bands, and these changes were related to the observed change in behavioral performance. The results reveal a causal relationship between intraparietal sulcus activity and tasks measuring both visual attention and VSTM.

Tuning perception: Visual working memory biases the quality of visual awareness.

Studies of consciousness reveal that it is possible to manipulate subjective awareness of a visual stimulus. For example, items held in visual working memory (VWM) that match target features increase the speed with which the target reaches visual awareness. To examine the effect of VWM on perception, previous studies have mainly used coarse measures of awareness, such as present/absent or forced-choice judgments. These methods can reveal whether or not an individual has seen an item, but they do not provide information about the quality with which the item was seen. Using continuous report methods it has been shown that the fidelity of a perceived item can be affected by whether or not that item is masked. In the present study, we used an object-substitution masking task to examine whether items held in VWM would influence the quality with which a masked target reached awareness, or whether the threshold for awareness was instead affected by stimuli held in memory. We observed that targets matching the contents of VWM were recalled with greater precision compared to items that did not match the contents of VWM. Importantly, this effect occurred without affecting the likelihood of the target being perceived. These results suggest that VWM plays a greater role in modulating the fidelity of perceived representations than in lowering the overall threshold of awareness.

Visual working memory simultaneously guides facilitation and inhibition during visual search.

During visual search, visual working memory (VWM) supports the guidance of attention in two ways: It stores the identity of the search target, facilitating the selection of matching stimuli in the search array, and it maintains a record of the distractors processed during search so that they can be inhibited. In two experiments, we investigated whether the full contents of VWM can be used to support both of these abilities simultaneously. In Experiment 1, participants completed a preview search task in which (a) a subset of search distractors appeared before the remainder of the search items, affording participants the opportunity to inhibit them, and (b) the search target varied from trial to trial, requiring the search target template to be maintained in VWM. We observed the established signature of VWM-based inhibition-reduced ability to ignore previewed distractors when the number of distractors exceeds VWM's capacity-suggesting that VWM can serve this role while also representing the target template. In Experiment 2, we replicated Experiment 1, but added to the search displays a singleton distractor that sometimes matched the color (a task-irrelevant feature) of the search target, to evaluate capture. We again observed the signature of VWM-based preview inhibition along with attentional capture by (and, thus, facilitation of) singletons matching the target template. These findings indicate that more than one VWM representation can bias attention at a time, and that these representations can separately affect selection through either facilitation or inhibition, placing constraints on existing models of the VWM-based guidance of attention.

Re-evaluating the relationships among filtering activity, unnecessary storage, and visual working memory capacity.

The amount of task-irrelevant information encoded in visual working memory (VWM), referred to as unnecessary storage, has been proposed as a potential mechanism underlying individual differences in VWM capacity. In addition, a number of studies have provided evidence for additional activity that initiates the filtering process originating in the frontal cortex and basal ganglia, and is therefore a crucial step in the link between unnecessary storage and VWM capacity. Here, we re-examine data from two prominent studies that identified unnecessary storage activity as a predictor of VWM capacity by directly testing the implied path model linking filtering-related activity, unnecessary storage, and VWM capacity. Across both studies, we found that unnecessary storage was not a significant predictor of individual differences in VWM capacity once activity associated with filtering was accounted for; instead, activity associated with filtering better explained variation in VWM capacity. These findings suggest that unnecessary storage is not a limiting factor in VWM performance, whereas neural activity associated with filtering may play a more central role in determining VWM performance that goes beyond preventing unnecessary storage.

The short- and long-term fates of memory items retained outside the focus of attention.

When a test of working memory (WM) requires the retention of multiple items, a subset of them can be prioritized. Recent studies have shown that, although prioritized (i.e., attended) items are associated with active neural representations, unprioritized (i.e., unattended) memory items can be retained in WM despite the absence of such active representations, and with no decrement in their recognition if they are cued later in the trial. These findings raise two intriguing questions about the nature of the short-term retention of information outside the focus of attention. First, when the focus of attention shifts from items in WM, is there a loss of fidelity for those unattended memory items? Second, could the retention of unattended memory items be accomplished by long-term memory mechanisms? We addressed the first question by comparing the precision of recall of attended versus unattended memory items, and found a significant decrease in precision for unattended memory items, reflecting a degradation in the quality of those representations. We addressed the second question by asking subjects to perform a WM task, followed by a surprise memory test for the items that they had seen in the WM task. Long-term memory for unattended memory items from the WM task was not better than memory for items that had remained selected by the focus of attention in the WM task. These results show that unattended WM representations are degraded in quality and are not preferentially represented in long-term memory, as compared to attended memory items.