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There is consistent debate over whether capacity in working memory (WM) is subject to an item limit, or whether an unlimited number of items can be held in this online memory system. The item limit hypothesis clearly predicts guessing responses when capacity is exceeded, and proponents of this view have highlighted evidence for guessing in visual working memory tasks. Nevertheless, various models that deny item limits can explain the same empirical patterns by asserting extremely low fidelity representations that cannot be distinguished from guesses. To address this ambiguity, we employed a task for which guess responses elicited a qualitatively distinct pattern from low fidelity memories. Inspired by work from Rouder et al. (2014), we employed an orientation WM task that required subjects to recall the precise orientation of each of six memoranda presented 1 s earlier. The orientation stimuli were created by rotating the position of a "clock hand" inside a circular region that was demarcated by four colored quadrants. Critically, when observers guess with these stimuli, the distribution of responses is biased towards the center of these quadrants, creating a "banded" pattern that cannot be explained by a low precision memory. We confirmed the presence of this guessing pattern using formal model comparisons, and we show that the prevalence of this pattern matches observers' own reports of when they thought they were guessing. Thus, these findings provide further evidence for guessing behaviors predicted by item limit models of WM capacity.

The contralateral delay activity (CDA) is an event-related potential component commonly used to examine the online processes of visual working memory. Here, we provide a robust analysis of the statistical power that is needed to achieve reliable and reproducible results with the CDA. Using two very large EEG datasets that examined the contrast between CDA amplitude with set sizes 2 and 6 items and set sizes 2 and 4 items, we present a subsampling analysis that estimates the statistical power achieved with varying numbers of subjects and trials based on the proportion of significant tests in 10,000 iterations. We also generated simulated data using Bayesian multilevel modeling to estimate power beyond the bounds of the original datasets. The number of trials and subjects required depends critically on the effect size. Detecting the presence of the CDA-a reliable difference between contralateral and ipsilateral electrodes during the memory period-required only 30-50 clean trials with a sample of 25 subjects to achieve approximately 80% statistical power. However, for detecting a difference in CDA amplitude between two set sizes, a substantially larger number of trials and subjects were required; approximately 400 clean trials with 25 subjects to achieve 80% power. Thus, to achieve robust tests of how CDA activity differs across conditions, it is essential to be mindful of the estimated effect size. We recommend researchers designing experiments to detect set-size differences in the CDA collect substantially more trials per subject.

Covert spatial attention has a variety of effects on the responses of individual neurons. However, relatively little is known about the net effect of these changes on sensory population codes, even though perception ultimately depends on population activity. Here, we measured the EEG in human observers (male and female), and isolated stimulus-evoked activity that was phase-locked to the onset of attended and ignored visual stimuli. Using an encoding model, we reconstructed spatially selective population tuning functions from the pattern of stimulus-evoked activity across the scalp. Our EEG-based approach allowed us to measure very early visually evoked responses occurring ∼100 ms after stimulus onset. In Experiment 1, we found that covert attention increased the amplitude of spatially tuned population responses at this early stage of sensory processing. In Experiment 2, we parametrically varied stimulus contrast to test how this effect scaled with stimulus contrast. We found that the effect of attention on the amplitude of spatially tuned responses increased with stimulus contrast, and was well described by an increase in response gain (i.e., a multiplicative scaling of the population response). Together, our results show that attention increases the gain of spatial population codes during the first wave of visual processing.SIGNIFICANCE STATEMENT We know relatively little about how attention improves population codes, even though perception is thought to critically depend on population activity. In this study, we used an encoding-model approach to test how attention modulates the spatial tuning of stimulus-evoked population responses measured with EEG. We found that attention multiplicatively scales the amplitude of spatially tuned population responses. Furthermore, this effect was present within 100 ms of stimulus onset. Thus, our results show that attention improves spatial population codes by increasing their gain at this early stage of processing.

Working memory (WM) is an online memory system that is critical for holding information in a rapidly accessible state during ongoing cognitive processing. Thus, there is strong value in methods that provide a temporally resolved index of WM load. While univariate EEG signals have been identified that vary with WM load, recent advances in multivariate analytic approaches suggest that there may be rich sources of information that do not generate reliable univariate signatures. Here, using data from four published studies (n = 286 and >250,000 trials), we demonstrate that multivariate analysis of EEG voltage topography provides a sensitive index of the number of items stored in WM that generalizes to novel human observers. Moreover, multivariate load detection ("mvLoad") can provide robust information at the single-trial level, exceeding the sensitivity of extant univariate approaches. We show that this method tracks WM load in a manner that is (1) independent of the spatial position of the memoranda, (2) precise enough to differentiate item-by-item increments in the number of stored items, (3) generalizable across distinct tasks and stimulus displays, and (4) correlated with individual differences in WM behavior. Thus, this approach provides a powerful complement to univariate analytic approaches, enabling temporally resolved tracking of online memory storage in humans.

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.

Covert spatial attention is essential for humans' ability to direct limited processing resources to the relevant aspects of visual scenes. A growing body of evidence suggests that rhythmic neural activity in the alpha frequency band (8-12 Hz) tracks the spatial locus of covert attention, which suggests that alpha activity is integral to spatial attention. However, extant work has not provided a compelling test of another key prediction: that alpha activity tracks the temporal dynamics of covert spatial orienting. In the current study, we examined the time course of spatially specific alpha activity after central cues and during visual search. Critically, the time course of this activity tracked trial-by-trial variations in orienting latency during visual search. These findings provide important new evidence for the link between rhythmic brain activity and covert spatial attention, and they highlight a powerful approach for tracking the spatial and temporal dynamics of this core cognitive process.

UnlabelledControl over visual selection has long been framed in terms of a dichotomy between "source" and "site," where top-down feedback signals originating in frontoparietal cortical areas modulate or bias sensory processing in posterior visual areas. This distinction is motivated in part by observations that frontoparietal cortical areas encode task-level variables (e.g., what stimulus is currently relevant or what motor outputs are appropriate), while posterior sensory areas encode continuous or analog feature representations. Here, we present evidence that challenges this distinction. We used fMRI, a roving searchlight analysis, and an inverted encoding model to examine representations of an elementary feature property (orientation) across the entire human cortical sheet while participants attended either the orientation or luminance of a peripheral grating. Orientation-selective representations were present in a multitude of visual, parietal, and prefrontal cortical areas, including portions of the medial occipital cortex, the lateral parietal cortex, and the superior precentral sulcus (thought to contain the human homolog of the macaque frontal eye fields). Additionally, representations in many-but not all-of these regions were stronger when participants were instructed to attend orientation relative to luminance. Collectively, these findings challenge models that posit a strict segregation between sources and sites of attentional control on the basis of representational properties by demonstrating that simple feature values are encoded by cortical regions throughout the visual processing hierarchy, and that representations in many of these areas are modulated by attention.Significance statementInfluential models of visual attention posit a distinction between top-down control and bottom-up sensory processing networks. These models are motivated in part by demonstrations showing that frontoparietal cortical areas associated with top-down control represent abstract or categorical stimulus information, while visual areas encode parametric feature information. Here, we show that multivariate activity in human visual, parietal, and frontal cortical areas encode representations of a simple feature property (orientation). Moreover, representations in several (though not all) of these areas were modulated by feature-based attention in a similar fashion. These results provide an important challenge to models that posit dissociable top-down control and sensory processing networks on the basis of representational properties.

We report 4 experiments examining whether associations in visual working memory are subject to proactive interference from long-term memory (LTM). Following a long-term learning phase in which participants learned the colors of 120 unique objects, a working memory (WM) test was administered in which participants recalled the precise colors of 3 concrete objects in an array. Each array in the WM test consisted of 1 old (previously learned) object with a new color (old-mismatch), 1 old object with its old color (old-match), and 1 new object. Experiments 1 to 3 showed that WM performance was better in the old-match condition than in the new condition, reflecting a beneficial contribution from LTM. In the old-mismatch condition, participants sometimes reported colors associated with the relevant shape in LTM, but the probability of successful recall was equivalent to that in the new condition. Thus, information from LTM only intruded in the absence of reportable information in WM. Experiment 4 tested for, and failed to find, proactive interference from the preceding trial in the WM test: Performance in the old-mismatch condition, presenting an object from the preceding trial with a new color, was equal to performance with new objects. Experiment 5 showed that long-term memory for object-color associations is subject to proactive interference. We conclude that the exchange of information between LTM and WM appears to be controlled by a gating mechanism that protects the contents of WM from proactive interference but admits LTM information when it is useful.

The positional-specificity effect refers to enhanced performance in visual short-term memory (VSTM) when the recognition probe is presented at the same location as had been the sample, even though location is irrelevant to the match/nonmatch decision. We investigated the mechanisms underlying this effect with behavioral and fMRI studies of object change-detection performance. To test whether the positional-specificity effect is a direct consequence of active storage in VSTM, we varied memory load, reasoning that it should be observed for all objects presented in a sub-span array of items. The results, however, indicated that although robust with a memory load of 1, the positional-specificity effect was restricted to the second of two sequentially presented sample stimuli in a load-of-2 experiment. An additional behavioral experiment showed that this disruption wasn't due to the increased load per se, because actively processing a second object--in the absence of a storage requirement--also eliminated the effect. These behavioral findings suggest that, during tests of object memory, position-related information is not actively stored in VSTM, but may be retained in a passive tag that marks the most recent site of selection. The fMRI data were consistent with this interpretation, failing to find location-specific bias in sustained delay-period activity, but revealing an enhanced response to recognition probes that matched the location of that trial's sample stimulus.

Perceptual grouping can lead observers to perceive a multielement scene as a smaller number of hierarchical units. Past work has shown that grouping enables more elements to be stored in visual working memory (WM). Although this may appear to contradict so-called discrete resource models that argue for fixed item limits in WM storage, it is also possible that grouping reduces the effective number of "items" in the display. To test this hypothesis, we examined how mnemonic resolution declined as the number of items to be stored increased. Discrete resource models predict that precision will reach a stable plateau at relatively early set sizes, because no further items can be stored once putative item limits are exceeded. Thus, we examined whether the precision by set size function was bilinear when storage was enhanced via perceptual grouping. In line with the hypothesis that each perceptual group counted as a single "item," precision still reached a clear plateau at a set size determined by the number of stored groups. Moreover, the maximum number of elements stored was doubled, and electrophysiological measures showed that selection and storage-related neural responses were the same for a single element and a multielement perceptual group. Thus, perceptual grouping allows more elements to be held in working memory while storage is still constrained by a discrete item limit. (Contains 7 figures.)

Observers can voluntarily select which items are encoded into working memory, and the efficiency of this process strongly predicts memory capacity. Nevertheless, the present work suggests that voluntary intentions do not exclusively determine what is encoded into this online workspace. Observers indicated whether any items from a briefly stored sample display had changed. Unbeknown to observers, these changes were most likely to occur in a specific quadrant of the display (the dominant quadrant). Across 84 subjects and 5 groups of observers, change detection accuracy was significantly higher for items in the dominant quadrant, suggesting that memory encoding was biased towards the dominant quadrant. Only 9 of the 84 subjects were able to correctly specify the dominant quadrant when asked whether any location was more likely to contain the changed item, but more sensitive forced-choice procedures did reveal above-chance discrimination of the dominant quadrant. Nevertheless, because forced choice performance was unrelated to the size of the bias and no observer reported a biased encoding strategy, the bias was unlikely to depend on voluntary encoding strategies. The encoding bias was not due to a reduction in the response threshold for indicating changes in the dominant quadrant (Experiment 2). Finally, separate measures of the number and resolution of the representations in memory suggested that encoding was biased in a discrete slot-based fashion (Experiment 3). That is, although items in the dominant quadrant were more likely to be encoded into memory, mnemonic resolution for the favored items was not affected. (Contains 3 footnotes and 6 figures.)

Are resources in visual working memory allocated in a continuous or a discrete fashion? On one hand, flexible resource models suggest that capacity is determined by a central resource pool that can be flexibly divided such that items of greater complexity receive a larger share of resources. On the other hand, if capacity in working memory is defined in terms of discrete storage "slots," then observers may be able to determine which items are assigned to a slot but not how resources are divided between stored items. To test these predictions, the authors manipulated the relative complexity of the items to be stored while holding the number items constant. Although mnemonic resolution declined when set size increased (Experiment 1), resolution for a given item was unaffected by large variations in the complexity of the other items to be stored when set size was held constant (Experiments 2-4). Thus, resources in visual working memory are distributed in a discrete slot-based fashion, even when interitem variations in complexity motivate an asymmetrical division of resources across items. (Contains 5 footnotes and 8 figures.)

Models of attentional control usually describe online shifts in control settings that accommodate changing task demands. The current studies suggest that online control over distractor exclusion--a core component of visual selection--can be accomplished without online shifts in top-down settings. Measurements of target discrimination accuracy suggested that the degree of distractor exclusion was guided by retinotopic maps of the prior probability of distractor interference at the attended locations. These probability maps can be retrieved via object-based cues, and they interact with shifts of attention to elicit increased levels of distractor exclusion when it is most needed. Thus, static probability maps can provide an internal template that guides the resolution of visual interference as spatial attention traverses the visual field.

When a visual target is identified, there is a period of several hundred milliseconds when the processing of subsequent targets is impaired, a phenomenon labeled the attentional blink (AB). The emerging consensus is that the identification of a visual target temporarily occupies a limited attentional resource that is essential for all visual perception. The present results challenge this view. With the same digit discrimination task that impaired subsequent letter discrimination for several hundred milliseconds, we found no disruption of subsequent face discrimination. These results suggest that all stimuli do not compete for access to a single resource for visual perception. We propose a multi-channel account of interference in the AB paradigm.