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Awh, E.

Publications and source records attributed to Awh, E..

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Item-specific delay activity demonstrates concurrent storage of multiple items in working memory

AbstractA longstanding view holds that information is maintained in working memory (WM) via persistent neural activity that encodes the content of WM. Recent work, however, has challenged the view that all items stored in WM are actively maintained. Instead, \"activity-silent\" models propose that items can be maintained in WM without the need for persistent neural activity, raising the possibility that only a subset of items - perhaps just a single item - may be actively represented at a given time. While past studies have successfully decoded multiple items stored in WM, these studies cannot rule out an active switching account in which only a single item is actively represented at a time. Here, we directly tested whether multiple representations can be held concurrently in an active state. We tracked spatial representations in WM using alpha-band (8-12 Hz) activity, which encodes spatial positions held in WM. Human observers (male and female) remembered one or two positions over a short delay while we recorded EEG. Using a spatial encoding model, we reconstructed stimulus-specific working memory representations (channel tuning functions, CTFs) from the scalp distribution of alphaband power. Consistent with past work, we found the selectivity of spatial CTFs was lower when two items were stored than when one item was stored. Critically, data-driven simulations revealed that the selectivity of spatial representations in the two-item condition could not be explained by models restricting storage to a single item at a time. Thus, our findings provide robust evidence for the concurrent storage of multiple items in visual working memory.\n\nAuthor SummaryWorking memory (WM) is a mental workspace where we temporarily hold information \"online\" in pursuit of our current goals. However, recent activity-silent models of WM have challenged the view that all items are held in an \"online\" state, instead proposing that only a subset of representations in WM - perhaps just one item - are represented by persistent activity at a time. To directly test a single-item model of persistent activity, we used a spatial encoding model to read out the strength of two representations from alpha-band (8-12 Hz) power in the human EEG signal. We provide direct evidence that both locations were maintained concurrently, ruling out the possibility that declines in stimulus-specific activity are due to storing one of two items in an activity-silent state.

neuroscience

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 online representations in working memory (WM) said to reside in the focus of attention, and passive offline representations of related information. Here, we dissect the focus of attention by showing that distinct neural signals index the online storage of objects and sustained spatial attention. We recorded EEG activity during two tasks that employed identical stimulus displays while the relative demands for object storage and spatial attention varied. We found distinct delay-period signatures for an attention task (which only required spatial attention) and 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.

neuroscience

Alpha-band oscillations track the retrieval of precise spatial representations from long-term memory

A hallmark of episodic memory is the phenomenon of mentally re-experiencing the details of past events, and a well-established concept is that the neuronal activity that mediates encoding is reinstated at retrieval. Evidence for reinstatement has come from multiple modalities, including functional Magnetic Resonance Imaging (fMRI) and electroencephalography (EEG). These EEG studies have shed light on the time-course of reinstatement, but have been limited to distinguishing between a few categories and/or limited measures of memory strength. The goal of this work was to investigate whether recently developed experimental and technical approaches, namely an inverted encoding model applied to alpha oscillatory power in conjunction with sensitive tests of memory retrieval in a continuous space, can track and reconstruct memory retrieval of specific spatial locations. In Experiment 1, we establish that an inverted encoding model applied to multivariate alpha topography can track retrieval of precise spatial memories. In Experiment 2, we demonstrate that the pattern of multivariate alpha activity at study is similar to the pattern observed during retrieval. Finally, we observe that these encoding models predict memory retrieval behavior, including the accuracy and latency of recall. These findings highlight the broad potential for using encoding models to characterize long-term memory retrieval.

neuroscience

Working memory implements distinct maintenance mechanisms depending on task goals

Working memory is the function by which we temporarily maintain information to achieve current task goals. Models of working memory typically debate where this information is stored, rather than how it is stored. Here we ask instead what neural mechanisms are involved in storage, and how these mechanisms change as a function of task goals. Participants either had to reproduce the orientation of a memorized bar (continuous recall task), or identify the memorized bar in a search array (visual search task). The sensory input and retention interval were identical in both tasks. Next, we used decoding and forward modeling on multivariate electroencephalogram (EEG) and time-frequency decomposed EEG to investigate which neural signals carry more informational content during the retention interval. In the continuous recall task, working memory content was preferentially carried by induced oscillatory alpha-band power, while in the visual search task it was more strongly carried by the distribution of evoked (consistently elevated and non-oscillatory) EEG activity. To show the independence of these two signals, we were able to remove informational content from one signal without affecting informational content in the other. Finally, we show that the tuning characteristics of both signals change in opposite directions depending on the current task goal. We propose that these signals reflect oscillatory and elevated firing-rate mechanisms that respectively support location-based and object-based maintenance. Together, these data challenge current models of working memory that place storage in particular regions, but rather emphasize the importance of different distributed maintenance signals depending on task goals.\n\nSignificance statement (120 words)Without realizing, we are constantly moving things in and out of our minds eye, an ability also referred to as working memory. Where did I put my screwdriver? Do we still have milk in the fridge? A central question in working memory research is how the brain maintains this information temporarily. Here we show that different neural mechanisms are involved in working memory depending on what the memory is used for. For example, remembering what a bottle of milk looks like invokes a different neural mechanism from remembering how much milk it contains: the first one primarily involved in being able to find the object, and the other one involving spatial position, such as the milk level in the bottle.

neuroscience