Search

We showed to the same observers both dynamic and static 2D patterns that can both evoke distinctive perceptions of motion or optic flow, as if moving in a tunnel or into a dark hole. At all times pupil diameters were monitored with an infrared eye tracker. We found a converging set of results indicating stronger pupil dilations to expansive growth of shapes or optic flows evoking a forward motion into a dark tunnel. Multiple regression analyses showed that the pupil responses to the illusory expanding black holes of static patterns were predicted by the individuals' pupil response to optic flows showing spiraling motion or "free fall" into a black hole. Also, individuals' pupil responses to spiraling motion into dark tunnels predicted the individuals' sense of illusory expansion with the static, illusory expanding, dark holes. This correspondence across individuals between their pupil responses to both dynamic and static, illusory expanding, holes suggests that these percepts reflect a common perceptual mechanism, deriving motion from 2D scenes, and that the observers' pupil adjustments reflect the direction and strength of motion they perceive and the expected outcome of an increase in darkness., Competing Interests: Declaration of Conflicting InterestsThe authors declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.

In humans, the eye pupils respond to both physical light sensed by the retina and mental representations of light produced by the brain. Notably, our pupils constrict when a visual stimulus is illusorily perceived brighter, even if retinal illumination is constant. However, it remains unclear whether such perceptual penetrability of pupil responses is an epiphenomenon unique to humans or whether it represents an adaptive mechanism shared with other animals to anticipate variations in retinal illumination between successive eye fixations. To address this issue, we measured the pupil responses of both humans and macaque monkeys exposed to three chromatic versions (cyan, magenta, and yellow) of the Asahi brightness illusion. We found that the stimuli illusorily perceived brighter or darker trigger differential pupil responses that are very similar in macaques and human participants. Additionally, we show that this phenomenon exhibits an analogous cyan bias in both primate species. Beyond evincing the macaque monkey as a relevant model to study the perceptual penetrability of pupil responses, our results suggest that this phenomenon is tuned to ecological conditions because the exposure to a "bright cyan-bluish sky" may be associated with increased risks of dazzle and retinal damages.

The pupil of the eye provides a rich source of information for cognitive scientists, as it can index a variety of bodily states (e.g., arousal, fatigue) and cognitive processes (e.g., attention, decision-making). As pupillometry becomes a more accessible and popular methodology, researchers have proposed a variety of techniques for analyzing pupil data. Here, we focus on time series-based, signal-to-signal approaches that enable one to relate dynamic changes in pupil size over time with dynamic changes in a stimulus time series, continuous behavioral outcome measures, or other participants' pupil traces. We first introduce pupillometry, its neural underpinnings, and the relation between pupil measurements and other oculomotor behaviors (e.g., blinks, saccades), to stress the importance of understanding what is being measured and what can be inferred from changes in pupillary activity. Next, we discuss possible pre-processing steps, and the contexts in which they may be necessary. Finally, we turn to signal-to-signal analytic techniques, including regression-based approaches, dynamic time-warping, phase clustering, detrended fluctuation analysis, and recurrence quantification analysis. Assumptions of these techniques, and examples of the scientific questions each can address, are outlined, with references to key papers and software packages. Additionally, we provide a detailed code tutorial that steps through the key examples and figures in this paper. Ultimately, we contend that the insights gained from pupillometry are constrained by the analysis techniques used, and that signal-to-signal approaches offer a means to generate novel scientific insights by taking into account understudied spectro-temporal relationships between the pupil signal and other signals of interest., (© 2023. The Author(s).)

Dilating the pupils allow more quanta of light to impact the retina. Consequently, if one pupil is dilated with a pharmacological agent (Tropicamide), the brightness of a surface under observation should increase proportionally to the pupil dilation. Little is known about causal effects of changes in pupil size on perception of an object's brightness. In a psychophysical procedure of brightness adjustment and matching, we presented to one eye geometrical patterns with a central square (the reference pattern) that differed in physical brightness within backgrounds of constant luminance. Subsequently, with the other eye, participants (n = 30) adjusted to the same luminance a similar pattern (target) whose central square luminance was randomly set higher or lower in brightness than the reference. As only one eye was treated with Tropicamide, we assessed whether the subjective brightness of the target square shifted in a consistent direction when viewed with the dilated pupil compared to the untreated (control) eye. We found that, as the pupil increased post drug administration, so significantly did the sense of brightness of the pattern (i.e., higher brightness adjustments followed viewing the reference pattern with the treated (Tropicamide) eye). A reversed effect was observed when the control eye viewed the reference pattern first. The results confirm that even slight pupil dilations can result in an enhanced perceptual experience of brightness of the attended object, corresponding to an average increase of 2.09 cd/m 2 for each 1 mm increase in pupil diameter., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Nicotine has been commonly used in pyschopharmacological studies, showing its benefits as a pharmacological stimulant on cognitive performance. In the current study, we investigated the effects of 2 mg (Experiment 1) and 4 mg (Experiment 2) of nicotine on performance on a multiple-object-tracking task. Participants were young non-smoking adults with no pre-existing attentional deficits. Nicotine and placebo were administered through nicotine and nicotine-free taste-matched chewing gum, respectively. Additionally, we compared pupil size between nicotine and placebo conditions in both experiments. Although we found that pupil size was considerably smaller in the nicotine conditions, nicotine administration did not appear to facilitate behavioural performance. We speculate that nicotine might enhance performance only for certain cognitive functions, and only for specific populations, such as nicotine-deprived smokers., (Copyright © 2020. Published by Elsevier B.V.)

Previous studies on individual differences in pupil size of healthy individuals and their relation to performance have been inconclusive. Using a novel approach, we tested the effect of general cognitive abilities and level of task performance on pretrial baseline and task-evoked pupil (TEP) sizes (N = 116) while we manipulated the level of task demands using a multiple object tracking task. Results did not reveal an effect of general cognitive abilities, estimated by working memory capacity and gF scores, on either baseline or TEP sizes. In contrast, we found an interaction in TEP sizes between level of overall MOT performance and task demands. We propose that individual differences in TEP sizes are related to state-specific level of task performance and task demands, probably in combination with other factors like age, personality traits, and state-specific level of motivation and arousal. We also suggest methodological confounds that may cause the previous inconclusive findings., (Copyright © 2020 The Authors. Published by Elsevier B.V. All rights reserved.)

Measuring task-evoked pupillary (TEP) responses as an index of phasic activity in the locus coeruleus (LC), we examined two competing hypotheses regarding the alerting and orienting mechanisms of attention. According to a dual mechanism account (Fernandez-Duque & Posner, 1997), two separate noradrenergic and cholinergic mechanisms modulate, respectively, the alerting and orienting effects. However, Corbetta and colleagues (2008) proposed that LC phasic activity may also be involved in orienting effect through its functional relationship with the ventral attentional network. We recruited seventy-five healthy Norwegian participants to perform a Posner cueing task. Both behavioral and pupillary responses revealed the alerting effect. Also, both behavioral and pupillary responses indicated that cued attention is affected by age. Behavioral responses also revealed orienting effect However, we found no TEP differences between valid, invalid, and neutral conditions, suggesting that TEP effects were driven by the alerting effect of cue presentation. Moreover, both behavioral and pupillary estimates of alertness and orienting were uncorrelated. Finally, individual differences in general cognitive abilities did not appear to affect the orienting and alerting mechanisms. This pattern of results is consistent with the dual mechanism account of attention. However, the LC involvement in the (re)orienting attention may be driven by state-specific factors., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

We hypothesized that a perceptually ambiguous or bistable object (Necker cube) can be more effectively biased to assume a point of view-from-above (VFA) than from below the object by cueing attention. Participants viewed a Necker cube in which one surface was temporarily shaded so as to prime a specific perspective on the cube. Subsequently, the standard (wireframe) Necker cube was viewed for 3 seconds, and participants reported what perspective they had seen initially and whether their perception shifted to the alternative perspective during the brief viewing. Concomitantly, pupil size was monitored with an eye-tracker to obtain an index of cognitive effort. There were two conditions: passive viewing and forced attention to sustain the initially primed perspective. We confirmed the presence of a VFA bias with forced attention, which was accompanied by reduced attentional effort, as indexed by a reduced pupil diameter, compared with the view-from-below. Participants showed no bias during passive viewing. We suggest that the level of intensive attention, when retrieving and maintaining a specific view from memory, is mirrored in the size of the eye pupils and may reflect ecological constraints on visual perception.

We hypothesized that pupil constrictions to the glare illusion, where converging luminance gradients subjectively enhance the perception of brightness, would be stronger for 'blue' than for other colors. Such an expectation was based on reflections about the ecology of vision, where the experience of dazzling light is common when one happens to look directly at sunlight through some occluders. Thus, we hypothesized that pupil constrictions to 'blue' reflect an ecologically-based expectation of the visual system from the experience of sky's light and color, which also leads to interpret the blue gradients of illusory glare to act as effective cues to impending probable intense light. We therefore manipulated the gradients color of glare illusions and measured changes in subjective brightness of identical shape stimuli. We confirmed that the blue resulted in what was subjectively evaluated as the brightest condition, despite all colored stimuli were equiluminant. This enhanced brightness effect was observed both in a psychophysical adjustment task and in changes in pupil size, where the maximum pupil constriction peak was observed with the 'blue' converging gradients over and above to the pupil response to blue in other conditions (i.e., diverging gradients and homogeneous patches). Moreover, glare-related pupil constrictions for each participant were correlated to each individual's subjective brightness adjustments. Homogenous blue hues also constricted the pupil more than other hues, which represents a pupillometric analog of the Helmholtz-Kohlrausch effect on brightness perception. Together, these findings show that pupillometry constitutes an easy tool to assess individual differences in color brightness perception., (Copyright © 2019 The Authors. Published by Elsevier B.V. All rights reserved.)

Dynamic non-luminance-mediated changes in pupil diameter have frequently been shown to be a reliable index for the level of arousal, mental effort, and activity in the locus coeruleus, the brainstem's noradrenergic arousal center. While pupillometry has most commonly been used to assess the level of arousal in particular psychological states or the level of engagement in cognitive tasks, some recent studies have found a relationship between average resting-state (i.e. baseline) pupil sizes and individuals' working memory capacity (WMC), indicating that individuals with higher WMC on average have larger pupils than individuals with relatively lower WMC. In the present study, we measured pupil size continuously in 212 participants during rest (i.e. while fixating) and estimated WMC in all participants by administering the Letter-Number Sequencing (LNS) task from WAIS-III. We were unable to replicate the relation between average pupil size and WMC. However, the novel finding was that higher WMC was associated with higher variability in resting-state pupil size. The present results are relevant for the current debate on the role of noradrenergic activity on working memory capacity., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Pupillary responses and associated vergence eye movements were monitored during imagery of objects of differing sizes ("large" or "small") from varying distances ("near" or "far"). Objects' imagined size and distance affected oculomotor behavior. Objects visualized as "far" resulted in the larger pupil dilations and smaller visual angle, while small objects imagined "near" were associated with smaller pupils in contrast to relatively larger pupils when imagined as "far" away. Furthermore, near objects resulted in larger visual angle, and particularly, vergence adjustments were observed that were dependent on imagined size and distance of the small objects. The findings are consistent with the idea that imagery emulates perception, also at the embodied level of oculomotor behaviors., (© 2018 Cognitive Science Society, Inc.)

During the execution of a cognitive task, the brain maintains contextual information to guide behavior and achieve desired goals. The AX-Continuous Performance Task is used to study proactive versus reactive cognitive control. Young adults tend to behave proactively in standard testing conditions. However, it remains unclear how interindividual variability (e.g., in cognitive and motivational factors) may drive people into more reactive or proactive control under the same task demands. We investigated the use of control strategies in a large population of healthy young adults. We computed the proactive behavioral index and consequently divided participants into proactive, reactive, and intermediate groups. We found that reactive participants were generally slower, presented lower context sensitivity, and larger response variability. Pupillary changes and blink rate index cognitive effort allocation. We measured, concomitantly to the task, the pupil size and frequency of blinks associated with the cue maintenance and response intervals. During the cue period, nonfrequent, nontarget cues led to increased pupil dilation and number of blinks in all participants. During the response interval, we found more errors and increased pupil dilation to the probe when all participants had to overcome a response bias generated by the frequent cue. Only reactive participants showed larger response-related pupil when they had to overcome a response bias related to the frequent probe. Contrary to expectations, groups did not differ in ocular measures in the cue period. In conclusion, interindividual differences in cognitive control between healthy adults can be mapped onto different patterns of effort allocation indexed by the pupil.

In an internet-based, forced-choice, test of the 'face race lightness illusion', the majority of respondents, regardless of their ethnicity, reported perceiving the African face as darker in skin tone than the European face, despite the mean luminance, contrast and numbers of pixels of the images were identical. In the laboratory, using eye tracking, it was found that eye fixations were distributed differently on the African face and European face, so that gaze dwelled relatively longer onto the locally brighter regions of the African face and, in turn, mean pupil diameters were smaller than for the European face. There was no relationship between pupils' size and implicit social attitude (IAT) scores. In another experiment, the faces were presented either tachistoscopically (140 ms) or longer (2500 ms) so that, when gaze was prevented from looking directly at the faces in the former condition, the tendency to report the African face as "dark" disappeared, but it was present when gaze was free to move for just a few seconds. We conclude that the presence of the illusion depends on oculomotor behavior and we also propose a novel account based on a predictive strategy of sensory acquisition. Specifically, by differentially directing gaze towards to facial regions that are locally different in luminance, the resulting changes in retinal illuminance yield respectively darker or brighter percepts while attending to each face, hence minimizing the mismatch between visual input and the learned perceptual prototypes of ethnic categories., Competing Interests: The authors have declared that no competing interests exist.

Introduction: In early multiple sclerosis (MS) patients, cognitive changes and fatigue are frequent and troublesome symptoms, probably related to both structural and functional brain changes. Whether there is a common cause of these symptoms in MS is unknown. In theory, an altered regulation of central neuropeptides can lead to changes in regulation of autonomic function, cognitive difficulties, and fatigue. Direct measurements of central neuropeptides are difficult to perform, but measurements of the eye pupil can be used as a reliable proxy of function., Methods: This study assesses pupil size during problem-solving in early MS patients versus controls. A difference in pupil size to a cognitive challenge could signal altered activity within the autonomic system because of early functional brain changes associated with cognitive load. We recruited MS patients (mean disease duration: 2.6 years, N  = 41) and age-matched healthy controls ( N  = 43) without eye pathology. Neurological impairment, magnetic resonance imaging, visual evoked potentials, depression, and fatigue were assessed in all of the patients. In both groups, we assessed processing speed and retinal imaging. Pupil size was recorded with an eye-tracker during playback of multiplication tasks., Results: Both groups performed well on the cognitive test. The groups showed similar pupillary responses with a mean of 0.55 mm dilation in patients and 0.54 mm dilation in controls for all the tasks collapsed together. However, controls ( N  = 9) with low cognitive scores (LCS) had an increased pupillary response to cognitive tasks, whereas LCS MS patients ( N  = 6) did not ( p  < .05). There was a tendency toward a smaller pupillary response in patients with fatigue., Conclusions: This is the first study to investigate pupillary responses to cognitive tasks in MS patients. Our results suggest that MS-related changes in cognition and fatigue may be associated with changes in arousal and the autonomic regulation of task-related pupillary responses. This supports the theory of a link between cognition and fatigue in MS.

Recent evidence showed that pupil dilation (PD) reflects modulations in the magnitude of the Simon interference effect due to correspondence sequence. In the present study we used this measure to assess whether these modulations, thought to result from cognitive control mechanisms, are influenced by prior practice with an incompatible stimulus-response (S-R) mapping. To this end, PD and reaction times (RTs) were recorded while participants performed a Simon task before and after executing a spatially incompatible practice. The sequential analysis revealed that PD mirrored the conflict-adaptation pattern observed in RTs. Crucially, sequential modulations were not affected by prior practice. These findings support the view that the modulations of the Simon effect due to prior practice and those due to correspondence sequence result from two different mechanisms, and suggest that PD can help to better understand the mechanisms underlying response selection and cognitive control in the Simon task., (Copyright © 2016 Elsevier B.V. All rights reserved.)

This study evaluated whether music-induced aesthetic "chill" responses, which typically correspond to peak emotional experiences, can be objectively monitored by degree of pupillary dilation. Participants listened to self-chosen songs versus control songs chosen by other participants. The experiment included an active condition where participants made key presses to indicate when experiencing chills and a passive condition (without key presses). Chills were reported more frequently for self-selected songs than control songs. Pupil diameter was concurrently measured by an eye-tracker while participants listened to each of the songs. Pupil size was larger within specific time-windows around the chill events, as monitored by key responses, than in comparison to pupil size observed during 'passive' song listening. In addition, there was a clear relationship between pupil diameter within the chills-related time-windows during both active and passive conditions, thus ruling out the possibility that chills-related pupil dilations were an artifact of making a manual response. These findings strongly suggest that music chills can be visible in the moment-to-moment changes in the size of pupillary responses and that a neuromodulatory role of the central norepinephrine system is thereby implicated in this phenomenon., (Copyright © 2016 The Authors. Published by Elsevier Inc. All rights reserved.)

Attentional effort relates to the allocation of limited-capacity attentional resources to meet current task demands and involves the activation of top-down attentional systems in the brain. Pupillometry is a sensitive measure of this intensity aspect of top-down attentional control. Studies relate pupillary changes in response to cognitive processing to activity in the locus coeruleus (LC), which is the main hub of the brain's noradrenergic system and it is thought to modulate the operations of the brain's attentional systems. In the present study, participants performed a visual divided attention task known as multiple object tracking (MOT) while their pupil sizes were recorded by use of an infrared eye tracker and then were tested again with the same paradigm while brain activity was recorded using fMRI. We hypothesized that the individual pupil dilations, as an index of individual differences in mental effort, as originally proposed by Kahneman (1973), would be a better predictor of LC activity than the number of tracked objects during MOT. The current results support our hypothesis, since we observed pupil-related activity in the LC. Moreover, the changes in the pupil correlated with activity in the superior colliculus and the right thalamus, as well as cortical activity in the dorsal attention network, which previous studies have shown to be strongly activated during visual tracking of multiple targets. Follow-up pupillometric analyses of the MOT task in the same individuals also revealed that individual differences to cognitive load can be remarkably stable over a lag of several years. To our knowledge this is the first study using pupil dilations as an index of attentional effort in the MOT task and also relating these to functional changes in the brain that directly implicate the LC-NE system in the allocation of processing resources.

If a mental image is a rerepresentation of a perception, then properties such as luminance or brightness should also be conjured up in the image. We monitored pupil diameters with an infrared eye tracker while participants first saw and then generated mental images of shapes that varied in luminance or complexity, while looking at an empty gray background. Participants also imagined familiar scenarios (e.g., a "sunny sky" or a "dark room") while looking at the same neutral screen. In all experiments, participants' eye pupils dilated or constricted, respectively, in response to dark and bright imagined objects and scenarios. Shape complexity increased mental effort and pupillary sizes independently of shapes' luminance. Because the participants were unable to voluntarily constrict their eyes' pupils, the observed pupillary adjustments to imaginary light present a strong case for accounts of mental imagery as a process based on brain states similar to those that arise during perception.

Sensing others' emotions through subtle facial expressions is a highly important social skill. We investigated the effects of intranasal oxytocin treatment on the evaluation of explicit and 'hidden' emotional expressions and related the results to individual differences in sensitivity to others' subtle expressions of anger and happiness. Forty healthy volunteers participated in this double-blind, placebo-controlled crossover study, which shows that a single dose of intranasal oxytocin (40 IU) enhanced or 'sharpened' evaluative processing of others' positive and negative facial expression for both explicit and hidden emotional information. Our results point to mechanisms that could underpin oxytocin's prosocial effects in humans. Importantly, individual differences in baseline emotional sensitivity predicted oxytocin's effects on the ability to sense differences between faces with hidden emotional information. Participants with low emotional sensitivity showed greater oxytocin-induced improvement. These participants also showed larger task-related pupil dilation, suggesting that they also allocated the most attentional resources to the task. Overall, oxytocin treatment enhanced stimulus-induced pupil dilation, consistent with oxytocin enhancement of attention towards socially relevant stimuli. Since pupil dilation can be associated with increased attractiveness and approach behaviour, this effect could also represent a mechanism by which oxytocin increases human affiliation.

We recorded by use of an infrared eye-tracker the pupil diameters of participants while they observed visual illusions of lightness or brightness. Four original illusions {based on Gaetano Kanisza's [Kanizsa G (1976) Subjective contours. Sci Am 234:48-52] and Akiyoshi Kitaoka's [Kitaoka A. (2005) Trick Eyes (Barnes & Noble, New Providence, NJ).] examples} were manipulated to obtain control conditions in which the perceived illusory luminance was either eliminated or reduced. All stimuli were equiluminant so that constrictions in pupillary size could not be ascribed to changes in light energy. We found that the pupillary diameter rapidly varied according to perceived brightness and lightness strength. Differences in local contrast information could be ruled out as an explanation because, in a second experiment, the observers maintained eye fixation in the center of the display; thus, differential stimulation of the fovea by local contrast changes could not be responsible for the pupillary differences. Hence, the most parsimonious explanation for the present findings is that pupillary responses to ambient light reflect the perceived brightness or lightness of the scene and not simply the amount of physical light energy entering the eye. Thus, the pupillary physiological response reflects the subjective perception of light and supports the idea that the brain's visual circuitry is shaped by visual experience with images and their possible sources.

People are generally better at remembering faces of their own race than faces of a different race, and this effect is known as the own-race bias (ORB) effect. We used eye-tracking and pupillometry to investigate whether Caucasian and Asian face stimuli elicited different-looking patterns in Caucasian participants in a face-memory task. Consistent with the ORB effect, we found better recognition performance for own-race faces than other-race faces, and shorter response times. In addition, at encoding, eye movements and pupillary responses to Asian faces (i.e., the other race) were different from those to Caucasian faces (i.e., the own race). Processing of own-race faces was characterized by more active scanning, with a larger number of shorter fixations, and more frequent saccades. Moreover, pupillary diameters were larger when viewing other-race than own-race faces, suggesting a greater cognitive effort when encoding other-race faces.

We recorded the pupil diameters of participants performing the words' color-naming Stroop task (i.e., naming the color of a word that names a color). Non-color words were used as baseline to firmly establish the effects of semantic relatedness induced by color word distractors. We replicated the classic Stroop effects of color congruency and color incongruency with pupillary diameter recordings: relative to non-color words, pupil diameters increased for color distractors that differed from color responses, while they reduced for color distractors that were identical to color responses. Analyses of the time courses of pupil responses revealed further differences between color-congruent and color-incongruent distractors, with the latter inducing a steep increase of pupil size and the former a relatively lower increase. Consistent with previous findings that have demonstrated that pupil size increases as task demands rise, the present results indicate that pupillometry is a robust measure of Stroop interference, and it represents a valuable addition to the cognitive scientist's toolbox.

Women's sexual preferences can change over the hormonal cycle, as several studies, based on responses to questionnaires, diaries, and ratings of photographs, have indicated increased sexual interests around the time of ovulation. However, fewer studies have measured changes in attention or interest to sexually significant stimuli in terms of physiological responses that are not under voluntary control and measure sexual interest indirectly (i.e., without mention of sexual feelings or activities). In the present study, we indexed changes in sexual interest in terms of changes in the eye pupil's size. Pupillary diameter is known to have a proportional relation to the observer's level of interest and attention to a visual stimulus as well as to physical pleasure. Fourteen women (7 being "pill" users) viewed photos on a computer screen while their pupil diameters were recorded using an infrared eye-tracking device. Three measures were taken for each participant during three time windows that estimated the ovulatory, luteal, and menstrual phase of the cycle. We found an increase in mean pupil diameter for sexually significant stimuli during the fertile phase and this pupillary change was also specific to pictures of the participants' actual sexual partners. Moreover, this effect was only seen for women who did not use oral contraceptives. These findings confirm that women's attention for sexually significant stimuli is higher during their fertile phase of the menstrual cycle, and that changes in sexual interest are implicitly measurable using pupillometry.