Summary An emerging view suggests that spatial position is an integral component of working memory (WM), such that non-spatial features are bound to locations regardless of whether space is relevant [1,2]. For instance, past work has shown that stimulus position is spontaneously remembered when non-spatial features are stored. Item recognition is enhanced when memoranda appear at the same location where they were encoded [3–5], and accessing non-spatial information elicits shifts of spatial attention to the original position of the stimulus [6,7]. However, these findings do not establish that a persistent, active representation of stimulus position is maintained in WM because similar effect have also been documented following storage in long-term memory [8,9]. Here, we show that the spatial position of the memorandum is actively coded by persistent neural activity during a non-spatial WM task. We used a spatial encoding model in conjunction with EEG measurements of oscillatory alpha-band (8–12 Hz) activity to track active representations of spatial position. The position of the stimulus varied trial-to-trial but was wholly irrelevant to the tasks. We nevertheless observed active neural representations of the original stimulus position that persisted throughout the retention interval. Further experiments established that these spatial representations are dependent on the volitional storage of non-spatial features rather than being a lingering effect of sensory energy or initial encoding demands. These findings provide strong evidence that online spatial representations are spontaneously maintained in WM – regardless of task relevance – during the storage of non-spatial features.

Covert spatial attention has long been thought to speed visual processing. Psychophysics studies have shown that target information accrues faster at attended locations than at unattended locations. However, with behavioral evidence alone, it is difficult to determine whether attention speeds visual processing of the target or subsequent postperceptual stages of processing (e.g., converting sensory responses into decision signals). Moreover, although many studies have shown that attention can boost the amplitude of visually evoked neural responses, no robust effect has been observed on the latency of those neural responses. Here, we offer new evidence that may reconcile the neural and behavioral findings. We examined whether covert attention influenced the latency of the N2pc component, an electrophysiological marker of visual selection that has been linked with object individuation-the formation of an object representation that is distinct from the background and from other objects in the scene. To this end, we manipulated whether or not human observers (male and female) covertly attended the location of an impending search target. We found that the target evoked N2pc onset ϳ20 ms earlier when the target location was cued than when it was not cued. In a second experiment, we provided a direct replication of this effect, confirming that the effect of attention on N2pc latency is robust. Thus, although attention may not speed the earliest stages of sensory processing, attention does speed the critical transition between raw sensory encoding and the formation of individuated object representations.

Covert spatial attention has long been thought to speed visual processing. Psychophysics studies have shown that target information accrues faster at attended locations than at unattended locations. However, with behavioral evidence alone, it is difficult to determine whether attention speeds visual processing of the target, or subsequent post-perceptual stages of processing (e.g. decision making and response selection). Moreover, while many studies have shown that that attention can boost the amplitude of visually-evoked neural responses, no effect has been observed on the latency of those neural responses. Here, we offer new evidence that may reconcile the neural and behavioral findings. Our study focused on the N2pc, an EEG marker of visual selection that has been linked with object individuation -the formation of an object representation that is distinct from the background and from other objects. In two experiments, we manipulated whether or not covert attention was precisely deployed to the location of an impending search target. We found that the target-evoked N2pc onset approximately 20 ms earlier when the target location was cued than when it was not cued. Thus, although attention may not speed the earliest stages of sensory processing, attention does speed the critical transition between raw sensory encoding and the formation of individuated object representations. Attention Speeds Target Individuation 3 Significance StatementCovert spatial attention improves processing at attended locations. Past behavioral studies have shown that information about visual targets accrues faster at attended than at unattended locations. However, it has remained unclear whether attention speeds perceptual analysis or subsequent post-perceptual stages of processing. Here we present robust evidence that attention speeds the N2pc, an electrophysiological signal that indexes the formation of individuated object representations. Our findings show that attention speeds a relatively early stage of perceptual processing, while also elucidating the specific perceptual process that is speeded. Attention Speeds Target Individuation

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