Modulation of the pupillary response by the content of visual working memory

Studies of selective attention during perception have revealed modulation of the pupillary response according to the brightness of task-relevant (attended) vs. -irrelevant (unattended) stimuli within a visual display. As a strong test of top-down modulation of the pupil response by selective attention, we asked whether changes in pupil diameter follow internal shifts of attention to memoranda of visual stimuli of different brightness maintained in working memory, in the absence of any visual stimulation. Across 3 studies, we reveal dilation of the pupil when participants orient attention to the memorandum of a dark grating relative to that of a bright grating. The effect occurs even when the attention-orienting cue is independent of stimulus brightness, and even when stimulus brightness is merely incidental and not required for the working-memory task of judging stimulus orientation. Furthermore, relative dilation and constriction of the pupil occurred dynamically and followed the changing temporal expectation that 1 or the other stimulus would be probed across the retention delay. The results provide surprising and consistent evidence that pupil responses are under top-down control by cognitive factors, even when there is no direct adaptive gain for such modulation, since no visual stimuli were presented or anticipated. The results also strengthen the view of sensory recruitment during working memory, suggesting even activation of sensory receptors. The thought-provoking corollary to our findings is that the pupils provide a reliable measure of what is in the focus of mind, thus giving a different meaning to old proverbs about the eyes being a window to the mind.

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Early Top–Down Control of Visual Processing Predicts Working Memory Performance

Journal of Cognitive Neuroscience ◽

10.1162/jocn.2009.21257 ◽

2010 ◽

Vol 22 (6) ◽

pp. 1224-1234 ◽

Cited By ~ 83

Author(s):

Aaron M. Rutman ◽

Wesley C. Clapp ◽

James Z. Chadick ◽

Adam Gazzaley

Keyword(s):

Working Memory ◽

Selective Attention ◽

Visual Processing ◽

Temporal Dynamics ◽

Memory Performance ◽

Event Related Potential ◽

Natural Scenes ◽

Attention And Memory ◽

Top Down ◽

Sensory Neural

Selective attention confers a behavioral benefit on both perceptual and working memory (WM) performance, often attributed to top–down modulation of sensory neural processing. However, the direct relationship between early activity modulation in sensory cortices during selective encoding and subsequent WM performance has not been established. To explore the influence of selective attention on WM recognition, we used electroencephalography to study the temporal dynamics of top–down modulation in a selective, delayed-recognition paradigm. Participants were presented with overlapped, “double-exposed” images of faces and natural scenes, and were instructed to either remember the face or the scene while simultaneously ignoring the other stimulus. Here, we present evidence that the degree to which participants modulate the early P100 (97–129 msec) event-related potential during selective stimulus encoding significantly correlates with their subsequent WM recognition. These results contribute to our evolving understanding of the mechanistic overlap between attention and memory.

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Electroencephalographic Evidence of Altered Top–Down Attentional Modulation in Fibromyalgia Patients During a Working Memory Task

Brain Topography ◽

10.1007/s10548-017-0561-3 ◽

2017 ◽

Vol 30 (4) ◽

pp. 539-547 ◽

Cited By ~ 5

Author(s):

Alberto J. González-Villar ◽

Marina Pidal-Miranda ◽

Manuel Arias ◽

Dolores Rodríguez-Salgado ◽

María T. Carrillo-de-la-Peña

Keyword(s):

Working Memory ◽

Memory Task ◽

Top Down ◽

Attentional Modulation

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Neural markers of category-based selective working memory in aging

10.1101/435388 ◽

2018 ◽

Author(s):

Robert M Mok ◽

M. Clare O'Donoghue ◽

Nicholas E Myers ◽

Erin H.S. Drazich ◽

Anna Christina Nobre

Keyword(s):

Older Adults ◽

Working Memory ◽

Selective Attention ◽

Memory Task ◽

Relevant Information ◽

Anterior Cingulate ◽

Memory Processing ◽

Normal Cognitive Function ◽

Psychophysiological Interaction ◽

Relevant Category

Working memory (WM) is essential for normal cognitive function, but shows marked decline in aging. Studies have shown that the ability to attend selectively to relevant information amongst competing distractors is related to WM capacity. The extent to which WM deficits in aging are related to impairments in selective attention is unclear. To investigate the neural mechanisms supporting selective attention in WM in aging, we tested a large group of older adults using functional magnetic resonance imaging whilst they performed a category-based (faces/houses) selective-WM task. Older adults were able to use attention to encode targets and suppress distractors to reach high levels of task performance. A subsequent, surprise recognition-memory task showed strong consequences of selective attention. Attended items in the relevant category were recognised significantly better than items in the ignored category. Neural measures also showed reliable markers of selective attention during WM. Purported control regions including the dorsolateral and inferior prefrontal and anterior cingulate cortex were reliably recruited for attention to both categories. Activation levels in category-sensitive visual cortex showed reliable modulation according to attentional demands, and positively correlated with subsequent memory measures of attention and WM span. Psychophysiological interaction analyses showed that activity in category-sensitive areas were coupled with non-sensory cortex known to be involved in cognitive control and memory processing, including regions in the PFC and hippocampus. In summary, we found that brain mechanisms of attention for selective WM are relatively preserved in aging, and individual differences in these abilities corresponded to the degree of attention-related modulation in the brain.

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Anticipatory cue‐related alpha desynchronization reflects top‐down disinhibitory control during verbal working memory task

Alzheimer s & Dementia ◽

10.1002/alz.052888 ◽

2021 ◽

Vol 17 (S5) ◽

Author(s):

Mina Mirjalili ◽

Reza Zomorrodi ◽

Zafiris J. Daskalakis ◽

Sean Hill ◽

Tarek K. Rajji

Keyword(s):

Working Memory ◽

Memory Task ◽

Verbal Working Memory ◽

Top Down

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Population Dynamics of Early Visual Cortex during Working Memory

Journal of Cognitive Neuroscience ◽

10.1162/jocn_a_01196 ◽

2018 ◽

Vol 30 (2) ◽

pp. 219-233 ◽

Cited By ~ 4

Author(s):

Masih Rahmati ◽

Golbarg T. Saber ◽

Clayton E. Curtis

Keyword(s):

Working Memory ◽

Population Dynamics ◽

Visual Cortex ◽

Parietal Cortex ◽

Visual Stimulus ◽

Visual Stimulation ◽

Brain Activity ◽

Top Down ◽

Population Activity ◽

Early Visual Cortex

Although the content of working memory (WM) can be decoded from the spatial patterns of brain activity in early visual cortex, how populations encode WM representations remains unclear. Here, we address this limitation by using a model-based approach that reconstructs the feature encoded by population activity measured with fMRI. Using this approach, we could successfully reconstruct the locations of memory-guided saccade goals based on the pattern of activity in visual cortex during a memory delay. We could reconstruct the saccade goal even when we dissociated the visual stimulus from the saccade goal using a memory-guided antisaccade procedure. By comparing the spatiotemporal population dynamics, we find that the representations in visual cortex are stable but can also evolve from a representation of a remembered visual stimulus to a prospective goal. Moreover, because the representation of the antisaccade goal cannot be the result of bottom–up visual stimulation, it must be evoked by top–down signals presumably originating from frontal and/or parietal cortex. Indeed, we find that trial-by-trial fluctuations in delay period activity in frontal and parietal cortex correlate with the precision with which our model reconstructed the maintained saccade goal based on the pattern of activity in visual cortex. Therefore, the population dynamics in visual cortex encode WM representations, and these representations can be sculpted by top–down signals from frontal and parietal cortex.

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Top-down modulation: bridging selective attention and working memory

Trends in Cognitive Sciences ◽

10.1016/j.tics.2011.11.014 ◽

2012 ◽

Vol 16 (2) ◽

pp. 129-135 ◽

Cited By ~ 600

Author(s):

Adam Gazzaley ◽

Anna C. Nobre

Keyword(s):

Working Memory ◽

Selective Attention ◽

Top Down

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Layer-specificity in the effects of attention and working memory on activity in primary visual cortex

Nature Communications ◽

10.1038/ncomms13804 ◽

2017 ◽

Vol 8 (1) ◽

Cited By ~ 61

Author(s):

Timo van Kerkoerle ◽

Matthew W. Self ◽

Pieter R. Roelfsema

Keyword(s):

Working Memory ◽

Visual Cortex ◽

Primary Visual Cortex ◽

Neuronal Activity ◽

Current Source ◽

Memory Task ◽

Neuronal Firing ◽

Top Down ◽

Attention Task ◽

Early Visual Cortex

Abstract Neuronal activity in early visual cortex depends on attention shifts but the contribution to working memory has remained unclear. Here, we examine neuronal activity in the different layers of the primary visual cortex (V1) in an attention-demanding and a working memory task. A current-source density analysis reveales top-down inputs in the superficial layers and layer 5, and an increase in neuronal firing rates most pronounced in the superficial and deep layers and weaker in input layer 4. This increased activity is strongest in the attention task but it is also highly reliable during working memory delays. A visual mask erases the V1 memory activity, but it reappeares at a later point in time. These results provide new insights in the laminar circuits involved in the top-down modulation of activity in early visual cortex in the presence and absence of visual stimuli.

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Interactions Between Visual Working Memory and Selective Attention

Psychological Science ◽

10.1111/1467-9280.00290 ◽

2000 ◽

Vol 11 (6) ◽

pp. 467-473 ◽

Cited By ~ 350

Author(s):

Paul E. Downing

Keyword(s):

Working Memory ◽

Selective Attention ◽

Visual Working Memory ◽

Memory Task ◽

Relevant Information ◽

Opposite Pattern ◽

Sample Object ◽

Probe Task ◽

Specific Prediction ◽

Memory Contents

The relationship between working memory and selective attention has traditionally been discussed as operating in one direction: Attention filters incoming information, allowing only relevant information into short-term processing stores. This study tested the prediction that the contents of visual working memory also influence the guidance of selective attention. Participants held a sample object in working memory on each trial. Two objects, one matching the sample and the other novel, were then presented simultaneously. As measured by a probe task, attention shifted to the object matching the sample. This effect generalized across object type, attentional-probe task, and working memory task. In contrast, a matched task with no memory requirement showed the opposite pattern, demonstrating that this effect is not simply due to exposure to the sample. These results confirm a specific prediction about the influence of working memory contents on the guidance of attention.

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The Role of Spatial Selective Attention in Working Memory for Locations: Evidence from Event-Related Potentials

Journal of Cognitive Neuroscience ◽

10.1162/089892900562444 ◽

2000 ◽

Vol 12 (5) ◽

pp. 840-847 ◽

Cited By ~ 131

Author(s):

Edward Awh ◽

Lourdes Anllo-Vento ◽

Steven A. Hillyard

Keyword(s):

Working Memory ◽

Selective Attention ◽

Visual Processing ◽

Spatial Working Memory ◽

Memory Task ◽

Stimulus Display ◽

Event Related Potentials ◽

Related Potentials ◽

On Line ◽

Multichannel Recordings

We investigated the hypothesis that the covert focusing of spatial attention mediates the on-line maintenance of location information in spatial working memory. During the delay period of a spatial working-memory task, behaviorally irrelevant probe stimuli were flashed at both memorized and nonmemorized locations. Multichannel recordings of event-related potentials (ERPs) were used to assess visual processing of the probes at the different locations. Consistent with the hypothesis of attention-based rehearsal, early ERP components were enlarged in response to probes that appeared at memorized locations. These visual modulations were similar in latency and topography to those observed after explicit manipulations of spatial selective attention in a parallel experimental condition that employed an identical stimulus display.

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