Stephen M. Emrich

Distributed patterns of activity in sensory cortex reflect the precision of multiple items maintained in visual short-term memory.

Traditionally, load sensitivity of sustained, elevated activity has been taken as an index of storage for a limited number of items in visual short-term memory (VSTM). Recently, studies have demonstrated that the contents of a single item held in VSTM can be decoded from early visual cortex, despite the fact that these areas do not exhibit elevated, sustained activity. It is unknown, however, whether the patterns of neural activity decoded from sensory cortex change as a function of load, as one would expect from a region storing multiple representations. Here, we use multivoxel pattern analysis to examine the neural representations of VSTM in humans across multiple memory loads. In an important extension of previous findings, our results demonstrate that the contents of VSTM can be decoded from areas that exhibit a transient response to visual stimuli, but not from regions that exhibit elevated, sustained load-sensitive delay-period activity. Moreover, the neural information present in these transiently activated areas decreases significantly with increasing load, indicating load sensitivity of the patterns of activity that support VSTM maintenance. Importantly, the decrease in classification performance as a function of load is correlated with within-subject changes in mnemonic resolution. These findings indicate that distributed patterns of neural activity in putatively sensory visual cortex support the representation and precision of information in VSTM.

Strengthened Effective Connectivity Underlies Transfer of Working Memory Training to Tests of Short-Term Memory and Attention

Although long considered a natively endowed and fixed trait, working memory (WM) ability has recently been shown to improve with intensive training. What remains controversial and poorly understood, however, are the neural bases of these training effects and the extent to which WM training gains transfer to other cognitive tasks. Here we present evidence from human electrophysiology (EEG) and simultaneous transcranial magnetic stimulation and EEG that the transfer of WM training to other cognitive tasks is supported by changes in task-related effective connectivity in frontoparietal and parieto-occipital networks that are engaged by both the trained and transfer tasks. One consequence of this effect is greater efficiency of stimulus processing, as evidenced by changes in EEG indices of individual differences in short-term memory capacity and in visual search performance. Transfer to search-related activity provides evidence that something more fundamental than task-specific strategy or stimulus-specific representations has been learned. Furthermore, these patterns of training and transfer highlight the role of common neural systems in determining individual differences in aspects of visuospatial cognition.

Visual Search Elicits the Electrophysiological Marker of Visual Working Memory

Background Although limited in capacity, visual working memory (VWM) plays an important role in many aspects of visually-guided behavior. Recent experiments have demonstrated an electrophysiological marker of VWM encoding and maintenance, the contralateral delay activity (CDA), which has been shown in multiple tasks that have both explicit and implicit memory demands. Here, we investigate whether the CDA is evident during visual search, a thoroughly-researched task that is a hallmark of visual attention but has no explicit memory requirements. Methodology/Principal Findings The results demonstrate that the CDA is present during a lateralized search task, and that it is similar in amplitude to the CDA observed in a change-detection task, but peaks slightly later. The changes in CDA amplitude during search were strongly correlated with VWM capacity, as well as with search efficiency. These results were paralleled by behavioral findings showing a strong correlation between VWM capacity and search efficiency. Conclusions/Significance We conclude that the activity observed during visual search was generated by the same neural resources that subserve VWM, and that this activity reflects the maintenance of previously searched distractors.

Competition increases binding errors in visual working memory

When faced with maintaining multiple objects in visual working memory, item information must be bound to the correct object in order to be correctly recalled. Sometimes, however, binding errors occur, and participants report the feature (e.g., color) of an unprobed, non-target item. In the present study, we examine whether the configuration of sample stimuli affects the proportion of these binding errors. The results demonstrate that participants mistakenly report the identity of the unprobed item (i.e., they make a non-target response) when sample items are presented close together in space, suggesting that binding errors can increase independent of increases in memory load. Moreover, the proportion of these non-target responses is linearly related to the distance between sample items, suggesting that these errors are spatially specific. Finally, presenting sample items sequentially decreases non-target responses, suggesting that reducing competition between sample stimuli reduces the number of binding errors. Importantly, these effects all occurred without increases in the amount of error in the memory representation. These results suggest that competition during encoding can account for some of the binding errors made during VWM recall.

Finding memory in search: The effect of visual working memory load on visual search

There is now substantial evidence that during visual search, previously searched distractors are stored in memory to prevent them from being reselected. Studies examining which memory resources are involved in this process have indicated that while a concurrent spatial working memory task does affect search slopes, depleting visual working memory (VWM) resources does not. In the present study, we confirm that VWM load indeed has no effect on the search slope; however, there is an increase in overall reaction times that is directly related to the number of items held in VWM. Importantly, this effect on search time increases proportionally with the memory load until the capacity of VWM is reached. Furthermore, the search task interfered with the number of items stored in VWM during the concurrent change-detection task. These findings suggest that VWM plays a role in the inhibition of previously searched distractors.

Visual Working Memory Supports the Inhibition of Previously Processed Information: Evidence From Preview Search

In four experiments we assessed whether visual working memory (VWM) maintains a record of previously processed visual information, allowing old information to be inhibited, and new information to be prioritized. Specifically, we evaluated whether VWM contributes to the inhibition (i.e., visual marking) of previewed distractors in a preview search. We evaluated this proposal by testing three predictions. First, Experiments 1 and 2 demonstrate that preview inhibition is more effective when the number of previewed distractors is below VWM capacity than above; an effect that can only be observed at small preview set sizes (Experiment 2A) and when observers are allowed to move their eyes freely (Experiment 2B). Second, Experiment 3 shows that, when quantified as the number of inhibited distractors, the magnitude of the preview effect is stable across different search difficulties. Third, Experiment 4 demonstrates that individual differences in preview inhibition are correlated with individual differences in VWM capacity. These findings provide converging evidence that VWM supports the inhibition of previewed distractors. More generally, these findings demonstrate how VWM contributes to the efficiency of human visual information processing--VWM prioritizes new information by inhibiting old information from being reselected for attention.

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.

Out with the old: inhibition of old items in a preview search is limited.

If some of the distractors in a visual search task are previewed prior to the presentation of the remaining distractors and the target, search time is reduced relative to when all of the items are displayed simultaneously. Here, we tested whether the ability to preferentially search new items during such a preview search is limited. We confirmed previous studies: The proportion of fixations on old items was significantly less than chance. However, the probability of fixating old locations was negatively affected by increasing the number of previewed distractors, suggesting that inhibition is limited to a small number of old items. Furthermore, the ability to inhibit old locations was limited to the first four fixations, indicating that by the fifth fixation, the resources required to sustain inhibition had been depleted. Together, these findings suggest that inhibition of old items in a preview search is a top-down mediated process dependent on capacity-limited cognitive resources.

Impairment in long-term retention but not short-term performance on a water maze reversal task following hippocampal or mediodorsal striatal N-methyl-D-aspartate receptor blockade.

Male Long-Evans rats were injected with 32 ng/mul of the N-methyl-D-aspartate (NMDA) receptor antagonist 3-(2-carboxypiperazin-4-yl) propyl-1-phosphonic acid (CPP) or vehicle and trained to locate a hidden platform in a different location (reversal training) than used on the initial 4 days of training. Rats treated with vehicle or CPP into the dorsal hippocampus, basolateral amygdala, or mediodorsal striatum had similar latencies to locate the platform on the reversal day. Rats infused with CPP into the dorsal hippocampus or mediodorsal striatum failed to search preferentially in the novel location during a 24-hr, drug-free retention test, whereas all other groups searched preferentially in this location. Therefore, blocking dorsal hippocampal or mediodorsal striatal NMDA receptors selectively blocked long-term spatial retention without producing short-term performance deficits.

Maintaining the ties that bind: The role of an intermediate visual memory store in the persistence of awareness

Segregation and feature binding are essential to the perception and awareness of objects in a visual scene. When a fragmented line-drawing of an object moves relative to a background of randomly oriented lines, the previously hidden object is segregated from the background and consequently enters awareness. Interestingly, in such shape-from-motion displays, the percept of the object persists briefly when the motion stops, suggesting that the segregated and bound representation of the object is maintained in awareness. Here, we tested whether this persistence effect is mediated by capacity-limited working-memory processes, or by the amount of object-related information available. The experiments demonstrate that persistence is affected mainly by the proportion of object information available and is independent of working-memory limits. We suggest that this persistence effect can be seen as evidence for an intermediate, form-based memory store mediating between sensory and working memory.