Modulations Of Pupil Size Research Articles

The effect of covert visual attention on pupil size during perceptual fading

Pupil size is modulated by various cognitive factors such as attention, working memory, mental imagery, and subjective perception. Previous studies examining cognitive effects on pupil size mainly focused on inducing or enhancing a subjective experience of brightness or darkness (for example by asking participants to attend to/ memorize a bright or dark stimulus), and then showing that this affects pupil size. Surprisingly, the inverse has never been done; that is, it is still unknown what happens when a subjective experience of brightness or darkness is eliminated or strongly reduced even though bright or dark stimuli are physically present. Here, we aim to answer this question by using perceptual fading, a phenomenon where a visual stimulus gradually fades from visual awareness despite its continuous presentation. The study contains two blocks: Fading and Non-Fading. In the Fading block, participants were presented with black and white patches with a fuzzy outline that were presented at the same location throughout the block, thus inducing strong perceptual fading. In contrast, in the Non-Fading block, the patches switched sides on each trial, thus preventing perceptual fading. Participants covertly attended to one of the two patches, indicated by a cue, and reported the offset of one of a set of circles that are displayed on top. We hypothesized that pupil size will be modulated by covert visual attention in the Non-Fading block, but that this effect will not (or to a lesser extent) arise in the Fading block. We found that covert visual attention to bright/ dark does modulate pupil size even during perceptual fading (Fading block), but to a lesser extent than when the perceptual experience of brightness/ darkness is preserved (Non-Fading block). This implies that pupil size is always modulated by covert attention, but that the effect decreases as subjective experience of brightness or darkness decreases. In broader terms, this suggests that cognitive modulations of pupil size reflect a mixture of high-level and lower-level visual processing.

Pupillometry indexes ocular dominance plasticity

Short-term monocular deprivation in normally sighted adult humans produces a transient shift of ocular dominance, boosting the deprived eye. This effect has been documented with both perceptual tests and through physiological recordings, but no previous study simultaneously measured physiological responses and the perceptual effects of deprivation. Here we propose an integrated experimental paradigm that combines binocular rivalry with pupillometry, to introduce an objective physiological index of ocular dominance plasticity, acquired concurrently with perceptual testing. Ten participants reported the perceptual dynamics of binocular rivalry, while we measured pupil diameter. Stimuli were a white and a black disk, each presented monocularly. Rivalry dynamics and pupil-size traces were compared before and after 2 h of monocular deprivation, achieved by applying a translucent patch over the dominant eye. Consistent with prior research, we observed that monocular deprivation boosts the deprived-eye signal and consequently increases ocular dominance. In line with previous studies, we also observed subtle but systematic modulations of pupil size that tracked alternations between exclusive dominance phases of the black or white disk. Following monocular deprivation, the amplitude of these pupil-size modulations increased, which is consistent with the post-deprivation boost of the deprived eye and the increase of ocular dominance. This provides evidence that deprivation impacts the effective strength of monocular visual stimuli, coherently affecting perceptual reports and the automatic and unconscious regulation of pupil diameter. Our results show that a combined paradigm of binocular rivalry and pupillometry gives new insights into the physiological mechanisms underlying deprivation effects.

Impact of light on task-evoked pupil responses during cognitive tasks.

Light has many non-image-forming functions including modulation of pupil size and stimulation of alertness and cognition. Part of these non-image-forming effects may be mediated by the brainstem locus coeruleus. The processing of sensory inputs can be associated with a transient pupil dilation that is likely driven in part by the phasic activity of the locus coeruleus. In the present study, we aimed to characterise the task-evoked pupil response associated with auditory inputs under different light levels and across two cognitive tasks. We continuously monitored the pupil of 20 young healthy participants (mean [SD] 24.05 [4.0] years; 14 women) whilst they completed an attentional and an emotional auditory task whilst exposed to repeated 30-40-s blocks of light interleaved with darkness periods. Blocks could either consist of monochromatic orange light (0.16 melanopic equivalent daylight illuminance (EDI) lux) or blue-enriched white light of three different levels [37, 92, 190 melanopic EDI lux; 6500 K]. For the analysis, 15 and then 14 participants were included in the attentional and emotional tasks, respectively. Generalised linear mixed models showed a significant main effect of light level on the task-evoked pupil responses triggered by the attentional and emotional tasks (p ≤ 0.0001). The impact of light was different for the target versus non-target stimulus of the attentional task but was not different for the emotional and neutral stimulus of the emotional task. There is a smaller sustained pupil size during brighter light blocks but, a higher light level triggers a stronger task-evoked pupil response to auditory stimulation, presumably through the recruitment of the locus coeruleus.

Pupil Size Is Sensitive to Low-Level Stimulus Features, Independent of Arousal-Related Modulation.

Similar to a camera aperture, pupil size adjusts to the surrounding luminance. Unlike a camera, pupil size is additionally modulated both by stimulus properties and by cognitive processes, including attention and arousal, though the interdependence of these factors is unclear. We hypothesized that different stimulus properties interact to jointly modulate pupil size while remaining independent from the impact of arousal. We measured pupil responses from human observers to equiluminant stimuli during a demanding rapid serial visual presentation (RSVP) task at fixation and tested how response amplitude depends on contrast, spatial frequency, and reward level. We found that under constant luminance, unattended stimuli evoke responses that are separable from changes caused by general arousal or attention. We further uncovered a double-dissociation between task-related responses and stimulus-evoked responses, suggesting that different sources of pupil size modulation are independent of one another. Our results shed light on neural pathways underlying pupillary response.

Data-Driven Pupil Response Profiles as Transdiagnostic Readouts for the Detection of Neurocognitive Functioning in Affective and Anxiety Disorders

BackgroundNeurocognitive functioning is a relevant transdiagnostic dimension in psychiatry. As pupil size dynamics track cognitive load during a working memory task, we aimed to explore if this parameter allows identification of psychophysiological subtypes in healthy participants and patients with affective and anxiety disorders. MethodsOur sample consisted of 226 participants who completed the n-back task during simultaneous functional magnetic resonance imaging and pupillometry measurements. We used latent class growth modeling to identify clusters based on pupil size in response to cognitive load. In a second step, these clusters were compared on affective and anxiety symptom levels, performance in neurocognitive tests, and functional magnetic resonance imaging activity. ResultsThe clustering analysis resulted in two distinct pupil response profiles: one with a stepwise increasing pupil size with increasing cognitive load (reactive group) and one with a constant pupil size across conditions (nonreactive group). A larger increase in pupil size was significantly associated with better performance in neurocognitive tests in executive functioning and sustained attention. Statistical maps of parametric modulation of pupil size during the n-back task showed the frontoparietal network in the positive contrast and the default mode network in the negative contrast. The pupil response profile of the reactive group was associated with more thalamic activity, likely reflecting better arousal upregulation and less deactivation of the limbic system. ConclusionsPupil measurements have the potential to serve as a highly sensitive psychophysiological readout for detection of neurocognitive deficits in the core domain of executive functioning, adding to the development of valid transdiagnostic constructs in psychiatry.

Early detection of cognitive decline in Alzheimer's disease using eye tracking.

Patients with Alzheimer's disease (AD) are known to exhibit visuospatial processing impairment, as reflected in eye movements from the early stages of the disease. We investigated whether the pattern of gaze exploration during visual tasks could be useful for detecting cognitive decline at the earliest stage. Sixteen AD patients (age: 79.1 ± 7.9 years, Mini Mental State Examination [MMSE] score: 17.7 ± 5.3, mean ± standard deviation) and 16 control subjects (age: 79.4 ± 4.6, MMSE score: 26.9 ± 2.4) participated. In the visual memory task, subjects memorized presented line drawings for later recall. In the visual search tasks, they searched for a target Landolt ring of specific orientation (serial search task) or color (pop-out task) embedded among arrays of distractors. Using video-oculography, saccade parameters, patterns of gaze exploration, and pupil size change during task performance were recorded and compared between AD and control subjects. In the visual memory task, the number of informative regions of interest (ROIs) fixated was significantly reduced in AD patients compared to control subjects. In the visual search task, AD patients took a significantly longer time and more saccades to detect the target in the serial but not in pop-out search. In both tasks, there was no significant difference in the saccade frequency and amplitude between groups. On-task pupil modulation during the serial search task was decreased in AD. The number of ROIs fixated in the visual memory task and search time and saccade numbers in the serial search task differentiated both groups of subjects with high sensitivity, whereas saccade parameters of pupil size modulation were effective in confirming normal cognition from cognitive decline with high specificity. Reduced fixation on informative ROIs reflected impaired attentional allocation. Increased search time and saccade numbers in the visual search task indicated inefficient visual processing. Decreased on-task pupil size during visual search suggested decreased pupil modulation with cognitive load in AD patients, reflecting impaired function of the locus coeruleus. When patients perform the combination of these tasks to visualize multiple aspects of visuospatial processing, cognitive decline can be detected at an early stage with high sensitivity and specificity and its progression be evaluated.

Value-driven modulation of visual perception by visual and auditory reward cues: The role of performance-contingent delivery of reward.

Perception is modulated by reward value, an effect elicited not only by stimuli that are predictive of performance-contingent delivery of reward (PC) but also by stimuli that were previously rewarded (PR). PC and PR cues may engage different mechanisms relying on goal-driven versus stimulus-driven prioritization of high value stimuli, respectively. However, these two modes of reward modulation have not been systematically compared against each other. This study employed a behavioral paradigm where participants' visual orientation discrimination was tested in the presence of task-irrelevant visual or auditory reward cues. In the first phase (PC), correct performance led to a high or low monetary reward dependent on the identity of visual or auditory cues. In the subsequent phase (PR), visual or auditory cues were not followed by reward delivery anymore. We hypothesized that PC cues have a stronger modulatory effect on visual discrimination and pupil responses compared to PR cues. We found an overall larger task-evoked pupil dilation in PC compared to PR phase. Whereas PC and PR cues both increased the accuracy of visual discrimination, value-driven acceleration of reaction times (RTs) and pupillary responses only occurred for PC cues. The modulation of pupil size by high reward PC cues was strongly correlated with the modulation of a combined measure of speed and accuracy. These results indicate that although value-driven modulation of perception can occur even when reward delivery is halted, stronger goal-driven control elicited by PC reward cues additionally results in a more efficient balance between accuracy and speed of perceptual choices.

Rule-based and stimulus-based cues bias auditory decisions via different computational and physiological mechanisms.

Expectations, such as those arising from either learned rules or recent stimulus regularities, can bias subsequent auditory perception in diverse ways. However, it is not well understood if and how these diverse effects depend on the source of the expectations. Further, it is unknown whether different sources of bias use the same or different computational and physiological mechanisms. We examined how rule-based and stimulus-based expectations influenced behavior and pupil-linked arousal, a marker of certain forms of expectation-based processing, of human subjects performing an auditory frequency-discrimination task. Rule-based cues consistently biased choices and response times (RTs) toward the more-probable stimulus. In contrast, stimulus-based cues had a complex combination of effects, including choice and RT biases toward and away from the frequency of recently presented stimuli. These different behavioral patterns also had: 1) distinct computational signatures, including different modulations of key components of a novel form of a drift-diffusion decision model and 2) distinct physiological signatures, including substantial bias-dependent modulations of pupil size in response to rule-based but not stimulus-based cues. These results imply that different sources of expectations can modulate auditory processing via distinct mechanisms: one that uses arousal-linked, rule-based information and another that uses arousal-independent, stimulus-based information to bias the speed and accuracy of auditory perceptual decisions.

Blunted pupil light reflex is associated with negative symptoms and working memory in individuals with schizophrenia

Two largely separate lines of research have documented altered pupillary dynamics in individuals diagnosed with schizophrenia. An older set of studies has demonstrated reductions in the pupillary light reflex (PLR) in individuals with schizophrenia; however, clinical and cognitive correlates of this blunted PLR have been relatively unexplored. More recently, a large body of work has demonstrated reductions in pupillary dilation in response to cognitive demands in individuals with schizophrenia, and the degree of this blunted pupil dilation has been related to more severe cognitive deficits and motivational negative symptoms. These clinically relevant alterations in the cognitive modulation of pupil size have been interpreted as reflecting insufficient information processing resources or inappropriate effort allocation. To begin to bridge these two lines of work, we investigated the PLR in 34 individuals with schizophrenia and 30 healthy controls and related the amplitude of the PLR to motivational negative symptoms and cognitive performance. Consistent with prior work, we found that the PLR was reduced in individuals with schizophrenia, and furthermore, that these measurements were highly reliable across individuals. Blunted constriction was associated with more severe motivational negative symptoms and poorer working memory among individuals with schizophrenia. These observed correlates provide a bridge between older literature documenting an altered PLR and more recent work reporting associations between negative symptoms, cognition, and blunted pupillary dilation in response to cognitive demands in individuals with schizophrenia. We provide possible mechanistic interpretations of our data and consider a parsimonious explanation for reduced cognitive- and light-related modulation of pupil size.

Pupil size as an indicator of cognitive activity in mild Alzheimer's disease.

It is well established that pupil activity indexes cognitive processing. For instance, research has consistently demonstrated that the pupil reacts to working memory span task performance. However, little is known about pupil reaction to cognitive processing in Alzheimer's Disease (AD). We thus investigated whether span tasks can modulate pupil size in patients with AD. We invited 24 patients with AD and 24 healthy older adults to perform backward and forward spans, as well as to count aloud in a control condition, while their pupil activity was recorded with eye tracking glasses. In patients with AD, analysis demonstrated larger pupil size during backward spans (M = 2.12, SD = .39) than during forward spans (M = 1.98, SD = .36) [t(23) = 3.22, p = .004], larger pupil size during forward spans than during counting (M = 1.67, SD = .33) [t(23) = 4.75, p < .001], as well as larger pupil size during backward spans than during counting [t(23) = 10.60, p < .001]. In control participants, analysis demonstrated larger pupil size during backward spans (M = 3.36, SD = .49) than during forward spans (M = 2.85, SD = .68) [t(23) = 5.82, p < .001], larger pupil size during forward spans than during counting (M = 2.09, SD = .62) [t(23) = 5.42, < .001], as well as larger pupil size during backward spans than during counting [t(23) = 9.70, p < .001]. Results also demonstrated a significant interaction effect between groups and conditions [F(2,92) = 16.63, p < .001]; in other words, patients with AD have shown fewer variations on the pupil size across the conditions compared to the control participants. The larger pupil size during backward spans, compared with forward spans or counting, can be attributed to the high cognitive load of backward spans. The modulation of pupil size, as observed across backward/forward spans and counting, can possibly be attributed to sympathetic/adrenergic and parasympathetic/cholinergic activities. Our study demonstrates the value of pupillometry as a potential biomarker of cognitive processing in AD.

Pupil constriction via the parasympathetic pathway precedes perceptual switch of ambiguous stimuli

Perceptual rivalry of ambiguous stimuli reflects the interaction of neural activity among multiple cortical regions. However, it remains unclear what drives a spontaneous perceptual alteration. We hypothesized that increased fluctuations in spontaneous neural activity due to arousal reduction drive the perceptual switch. Here, we show that the pupils shrank a few seconds prior to the onset of the spontaneous perceptual switch. Such pupil constriction was not observed before the exogenous perceptual switch. Pharmacological experiments confirmed that the pupil constriction disappeared when the peripheral parasympathetic pathway (pupil sphincter muscle) was blocked, but it remained intact when the peripheral sympathetic pathway (pupil dilator muscle) was manipulated. Furthermore, rapid pupil dilations with behavioral response are also mediated by the peripheral parasympathetic pathway. The present findings suggested that transient arousal drops, as denoted by the autonomic nervous modulation of pupil size, are involved in inducing the spontaneous perceptual switch of bistable stimuli.

Multisensory signals inhibit pupillary light reflex: Evidence from pupil oscillation.

Multisensory integration, which enhances stimulus saliency at the early stage of the processing hierarchy, has been recently shown to produce a larger pupil size than its unisensory constituents. Theoretically, any modulation on pupil size ought to be associated with the sympathetic and parasympathetic pathways that are sensitive to light. But it remains poorly understood how the pupillary light reflex is changed in a multisensory context. The present study evoked an oscillation of the pupillary light reflex by periodically changing the luminance of a visual stimulus at 1.25Hz. It was found that such induced pupil size oscillation was substantially attenuated when the bright but not the dark phase of the visual flicker was periodically and synchronously presented with a burst of tones. This inhibition effect persisted when the visual flicker was task-irrelevant and out of attentional focus, but disappeared when the visual flicker was moved from the central field to the periphery. These findings not only offer a comprehensive characterization of the multisensory impact on pupil response to light, but also provide valuable clues about the individual contributions of the sympathetic and parasympathetic pathways to multisensory modulation of pupil size.

Objective pupillometry shows that perceptual styles covary with autistic-like personality traits

We measured the modulation of pupil size (in constant lighting) elicited by observing transparent surfaces of black and white moving dots, perceived as a cylinder rotating about its vertical axis. The direction of rotation was swapped periodically by flipping stereo-depth of the two surfaces. Pupil size modulated in synchrony with the changes in front-surface color (dilating when black). The magnitude of pupillary modulation was larger for human participants with higher Autism-Spectrum Quotient (AQ), consistent with a local perceptual style, with attention focused on the front surface. The modulation with surface color, and its correlation with AQ, was equally strong when participants passively viewed the stimulus. No other indicator, including involuntary pursuit eye movements, covaried with AQ. These results reinforce our previous report with a similar bistable stimulus (Turi, Burr, & Binda, 2018), and go on to show that bistable illusory motion is not necessary for the effect, or its dependence on AQ.

Factors Influencing Saccadic Reaction Time: Effect of Task Modality, Stimulus Saliency, Spatial Congruency of Stimuli, and Pupil Size.

It is often assumed that the reaction time of a saccade toward visual and/or auditory stimuli reflects the sensitivities of our oculomotor-orienting system to stimulus saliency. Endogenous factors, as well as stimulus-related factors, would also affect the saccadic reaction time (SRT). However, it was not clear how these factors interact and to what extent visual and auditory-targeting saccades are accounted for by common mechanisms. The present study examined the effect of, and the interaction between, stimulus saliency and audiovisual spatial congruency on the SRT for visual- and for auditory-target conditions. We also analyzed pre-target pupil size to examine the relationship between saccade preparation and pupil size. Pupil size is considered to reflect arousal states coupling with locus-coeruleus (LC) activity during a cognitive task. The main findings were that (1) the pattern of the examined effects on the SRT varied between visual- and auditory-auditory target conditions, (2) the effect of stimulus saliency was significant for the visual-target condition, but not significant for the auditory-target condition, (3) Pupil velocity, not absolute pupil size, was sensitive to task set (i.e., visual-targeting saccade vs. auditory-targeting saccade), and (4) there was a significant correlation between the pre-saccade absolute pupil size and the SRTs for the visual-target condition but not for the auditory-target condition. The discrepancy between target modalities for the effect of pupil velocity and between the absolute pupil size and pupil velocity for the correlation with SRT may imply that the pupil effect for the visual-target condition was caused by a modality-specific link between pupil size modulation and the SC rather than by the LC-NE (locus coeruleus-norepinephrine) system. These results support the idea that different threshold mechanisms in the SC may be involved in the initiation of saccades toward visual and auditory targets.

Phasic Activation of Dorsal Raphe Serotonergic Neurons Increases Pupil Size

SummaryTransient variations in pupil size (PS) under constant luminance are coupled to rapid changes in arousal state,1, 2, 3 which have been interpreted as vigilance,4 salience,5 or a surprise signal.6, 7, 8 Neural control of such fluctuations presumably involves multiple brain regions5,9, 10, 11 and neuromodulatory systems,3,12,13 but it is often associated with phasic activity of the noradrenergic system.9,12,14,15 Serotonin (5-HT), a neuromodulator also implicated in aspects of arousal16 such as sleep-wake transitions,17 motivational state regulation,18 and signaling of unexpected events,19 seems to affect PS,20, 21, 22, 23, 24 but these effects have not been investigated in detail. Here we show that phasic 5-HT neuron stimulation causes transient PS changes. We used optogenetic activation of 5-HT neurons in the dorsal raphe nucleus (DRN) of head-fixed mice performing a foraging task. 5-HT-driven modulations of PS were maintained throughout the photostimulation period and sustained for a few seconds after the end of stimulation. We found no evidence that the increase in PS with activation of 5-HT neurons resulted from interactions of photostimulation with behavioral variables, such as locomotion or licking. Furthermore, we observed that the effect of 5-HT on PS depended on the level of environmental uncertainty, consistent with the idea that 5-HT could report a surprise signal.19 These results advance our understanding of the neuromodulatory control of PS, revealing a tight relationship between phasic activation of 5-HT neurons and changes in PS.

Speed-accuracy tradeoff of visual search: Modulation of pupil size

Speed-accuracy tradeoff of visual search: Modulation of pupil size

Risk prediction error signaling: A two-component response?

Organisms use rewards to navigate and adapt to (uncertain) environments. Error-based learning about rewards is supported by the dopaminergic system, which is thought to signal reward prediction errors to make adjustments to past predictions. More recently, the phasic dopamine response was suggested to have two components: the first rapid component is thought to signal the detection of a potentially rewarding stimulus; the second, slightly later component characterizes the stimulus by its reward prediction error. Error-based learning signals have also been found for risk. However, whether the neural generators of these signals employ a two-component coding scheme like the dopaminergic system is unknown. Here, using human high density EEG, we ask whether risk learning, or more generally speaking surprise-based learning under uncertainty, is similarly comprised of two temporally dissociable components. Using a simple card game, we show that the risk prediction error is reflected in the amplitude of the P3b component. This P3b modulation is preceded by an earlier component, that is modulated by the stimulus salience. Source analyses are compatible with the idea that both the early salience signal and the later risk prediction error signal are generated in insular, frontal, and temporal cortex. The identified sources are parts of the risk processing network that receives input from noradrenergic cells in the locus coeruleus. Finally, the P3b amplitude modulation is mirrored by an analogous modulation of pupil size, which is consistent with the idea that both the P3b and pupil size indirectly reflect locus coeruleus activity.

No modulation of pupil size and event-related pupil response by transcutaneous auricular vagus nerve stimulation (taVNS)

Transcutaneous auricular vagus nerve stimulation (taVNS) bears therapeutic potential for a wide range of medical conditions. However, previous studies have found substantial interindividual variability in responsiveness to taVNS, and no reliable predictive biomarker for stimulation success has been developed so far. In this study, we investigate pupil size and event-related pupil response as candidate biomarkers. Both measures have a direct physiological link to the activity of the locus coeruleus (LC), a brainstem structure and the main source of norepinephrine in the brain. LC activation is considered one of the key mechanisms of action of taVNS, therefore, we expected a clear increase of the pupillary measures under taVNS compared to sham (placebo) stimulation, such that it could serve as a prospective predictor for individual clinical and physiological taVNS effects in future studies. We studied resting pupil size and pupillary responses to target stimuli in an auditory oddball task in 33 healthy young volunteers. We observed stronger pupil responses to target than to standard stimuli. However, and contrary to our hypothesis, neither pupil size nor the event-related pupil response nor behavioral performance were modulated by taVNS. We discuss potential explanations for this negative finding and its implications for future clinical investigation and development of taVNS.

Melanopsin and Cone Photoreceptor Inputs to the Afferent Pupil Light Response.

Background: Retinal photoreceptors provide the main stage in the mammalian eye for regulating the retinal illumination through changes in pupil diameter, with a small population of melanopsin-expressing intrinsically photosensitive retinal ganglion cells (ipRGCs) forming the primary afferent pathway for this response. The purpose of this study is to determine how melanopsin interacts with the three cone photoreceptor classes in the human eye to modulate the light-adapted pupil response.Methods: We investigated the independent and combined contributions of the inner and outer retinal photoreceptor inputs to the afferent pupil pathway in participants with trichromatic color vision using a method to independently control the excitations of ipRGCs, cones and rods in the retina.Results: We show that melanopsin-directed stimuli cause a transient pupil constriction generated by cones in the shadow of retinal blood vessels; desensitizing these penumbral cone signals uncovers a signature melanopsin pupil response that includes a longer latency (292 ms) and slower time (4.1x) and velocity (7.7x) to constriction than for cone-directed stimuli, and which remains sustained post-stimulus offset. Compared to melanopsin-mediated pupil responses, the cone photoreceptor-initiated pupil responses are more transient with faster constriction latencies, higher velocities and a secondary constriction at light offset. The combined pupil responses reveal that melanopsin signals are additive with the cone signals.Conclusions: The visual system uses the L–, M–, and S–cone photoreceptor inputs to the afferent pupil pathway to accomplish the tonic modulations of pupil size to changes in image contrast. The inner retinal melanopsin-expressing ipRGCs mediate the longer-term, sustained pupil constriction to set the light-adapted pupil diameter during extended light exposures.

Both a Gauge and a Filter: Cognitive Modulations of Pupil Size.

Over 50 years of research have established that cognitive processes influence pupil size. This has led to the widespread use of pupil size as a peripheral measure of cortical processing in psychology and neuroscience. However, the function of cortical control over the pupil remains poorly understood. Why does visual attention change the pupil light reflex? Why do mental effort and surprise cause pupil dilation? Here, we consider these functional questions as we review and synthesize two literatures on cognitive effects on the pupil: how cognition affects pupil light response and how cognition affects pupil size under constant luminance. We propose that cognition may have co-opted control of the pupil in order to filter incoming visual information to optimize it for particular goals. This could complement other cortical mechanisms through which cognition shapes visual perception.