The Pupillary Light Response Reflects Visual Working Memory Content

Hustá, Cecília; Dalmaijer, Edwin S.; Belopolsky, Artem V.; Mathôt, Sebastiaan

doi:10.1101/477562

Cited by 5 publications

(4 citation statements)

References 35 publications

(31 reference statements)

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“…Visual working memory-Pupil size also reflects the contents maintained in working memory. This has been shown in several studies (Blom et al, 2016;Fabius et al, 2017;Hustá et al, 2019;Unsworth & Robison, 2017;Zokaei et al, 2019); for example, in a study by Hustá and colleagues (2019), participants were presented with dark and bright stimuli (Fig. 1d,e).…”

Section: Functional Benefits Of Cognitively Driven Pupil-size Changessupporting

confidence: 64%

Functional benefits of cognitively driven pupil-size changes

Vilotijević,

Mathot

2023

Preprint

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Pupil-size changes are typically associated with the pupil light response (PLR), wherethey are driven by the physical entry of light into the eye. However, pupil-size changes are also influenced by various cognitive processes, where they are driven byhigher-level cognition. For example, the strength of the PLR is not solely affected byphysical properties of the light but also by cognitive factors, such as whether thesource of light is attended or not, which results in an increase or decrease in thestrength of the PLR. Surprisingly, although cognitively driven pupil-size changes havebeen the focus of extensive research, their possible functions are rarely discussed.Here we consider the relative (dis)advantages of small versus large pupils in different situations from a theoretical point of view, and compare these to empirical results showing how pupil size actually changes in these situations. Based on this, we suggest that cognitively driven pupil-size changes optimize vision either through preparation, embodied representations, or a differential emphasis on central or peripheral vision. More generally, we argue that cognitively driven pupil-size changes are a form of sensory tuning: a subtle adjustment of the eyes to optimize vision for the current situation and the immediate future.

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Section: Functional Benefits Of Cognitively Driven Pupil-size Changessupporting

confidence: 64%

Functional benefits of cognitively driven pupil-size changes

Vilotijević,

Mathot

2023

Preprint

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“…Thus, across our 3 experiments, we gathered the first set of clear evidence substantiating the proposal that pupillary responses adjust according to the brightness of items in working memory. Unlike previous studies addressing similar issues (22, 23, 25), our study carefully excludes any possible contamination by presentation or anticipation of visual stimuli of different brightness, in order to rule out explanations based on perceptual attention mechanisms related to selective encoding or anticipation. Our striking observations suggest that attention in working memory draws on similar sensory-orienting mechanisms as external attention (16, 17) to prioritize items.…”

Section: Discussionmentioning

confidence: 99%

“…Another recent preprint (25) reports a study specially aimed at testing pupil responses linked to shifts of selective attention within working memory, but in this case it is not possible to separate attention-related modulation of working-memory memoranda from preparatory attention to anticipated probe stimuli. In this case, the brightness of the probe stimuli matched that of the cued memorandum.…”

mentioning

confidence: 99%

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

Zokaei

Board

Manohar

et al. 2019

Proc. Natl. Acad. Sci. U.S.A.

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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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“…In recent years, studies have shown that the PLR is modulated by a wide range of cognitive factors: eye-movement preparation, such that a pupil constriction is prepared along with an eye movement towards a bright object (Ebitz, Pearson, & Platt, 2014;Mathôt, van der Linden, Grainger, & Vitu, 2015); covert visual attention, such that the PLR is strongest to stimuli presented within the focus of attention (Binda, Pereverzeva, & Murray, 2013a;Mathôt, van der Linden, Grainger, & Vitu, 2013;Naber, Alvarez, & Nakayama, 2013) ; visual working memory (vWM), such that the Manuscript in preparation [v1.0.2; 3 of 19 pupil is smaller when a bright, as compared to a dark, stimulus is maintained in vWM (Husta, Dalmaijer, Belopolsky, & Mathot, 2018) ; subjective interpretation, such that the pupil is smaller when people interpret a picture as depicting something bright, such as a sun, regardless of the actual brightness of the picture (Binda, Pereverzeva, & Murray, 2013b;Laeng & Endestad, 2012;; and mental imagery, such that the pupil constricts when people merely imagine a bright object or scene (Laeng & Sulutvedt, 2014). Taken together, these studies show clearly that the PLR, while not being under direct voluntary control, is modulated by higher-level cognition.…”

Section: The Effect Of Semantic Brightness On Pupil Sizementioning

confidence: 99%

The effect of semantic brightness on pupil size: A replication with Dutch words

Mathôt

Sundermann

Rijn

2019

Preprint

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Theories of embodied language hold that word processing is automatically accompanied by sensory and motor simulations. For example, when you read the word 'sun', a sensory simulation of brightness as well as a motor simulation of pupil constriction would be automatically triggered.Consistent with this notion, Mathôt, Grainger, and Strijkers (2017) found that the eye's pupil was slightly smaller after reading single words that were associated with brightness (e.g. 'sun') as compared to darkness (e.g. 'night' ); that is, the pupil light response was modulated by the semantic brightness of words. However, (other) key findings within the field of embodied language have proven difficult to replicate, and we therefore felt that it was crucial to replicate the effect of semantic brightness on pupil size. To this end, we conducted a close-but-non-identical replication of two key experiments from Mathôt, Grainger, and Strijkers (2017): one experiment with visually presented words, and one experiment with spoken words. Both experiments were successfully replicated. We propose that cognitive modulations of the pupil light response reflect activity in visual brain areas; therefore, the effect of semantic brightness on pupil size can be used as a marker for the involvement of visual brain areas in language processing, and thus to address a wide variety of key questions within psycholinguistics.

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The Pupillary Light Response Reflects Visual Working Memory Content

Abstract: 2Recent studies have shown that the pupillary light response (PLR) is modulated by 3 higher cognitive functions, presumably through activity in visual sensory brain areas. 4

Cited by 5 publications

References 35 publications

Functional benefits of cognitively driven pupil-size changes

Functional benefits of cognitively driven pupil-size changes

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

The effect of semantic brightness on pupil size: A replication with Dutch words

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