Pupils Dilated Research Articles

Anticipating noxious stimulation rather than afferent nociceptive input may evoke pupil asymmetry

Unilateral nociceptive stimulation is associated with subtle signs of pupil asymmetry that may reflect lateralized activity in the locus coeruleus. To explore drivers of this pupil asymmetry, electrical stimuli, delivered alone or 200 ms before or after an acoustic startle stimulus, were administered to one ankle under four experimental conditions: with or without a 1.6 s anticipatory period, or while the forearm ipsilateral or contralateral to the electrical stimulus was heated tonically to induce moderate pain (15 healthy participants in each condition). Pupil diameter was measured at the start of each trial, at stimulus delivery, and each second for 5 s after stimulus delivery. At the start of the first trial, the pupil ipsilateral to the side on which electric shocks were later delivered was larger than the contralateral pupil. Both pupils dilated robustly during the anticipatory period and dilated further during single- and dual-stimulus trials. However, pupil asymmetry persisted throughout the experiment. Tonically-applied forearm heat-pain modulated the pupillary response to phasic electrical stimuli, with a slight trend for dilatation to be greater contralateral to the forearm being heated. Together, these findings suggest that focusing anxiously on the expected site of noxious stimulation was associated with dilatation of the ipsilateral pupil whereas phasic nociceptive stimuli and psychological arousal triggered bilateral pupillary dilatation. It was concluded that preparatory cognitive activity rather than phasic afferent nociceptive input is associated with pupillary signs of lateralized activity in the locus coeruleus.

Pupils Dilate More to Harder Vocabulary Words than Easier Ones.

Understanding cognitive effort expended during assessments is essential to improving efficiency, accuracy, and accessibility within these assessments. Pupil dilation is commonly used as a psychophysiological measure of cognitive effort, yet research on its relationship with effort expended specifically during language processing is limited. The present study adds to and expands on this literature by investigating the relationships among pupil dilation, trial difficulty, and accuracy during a vocabulary test. Participants (n = 63, Mage = 19.25) completed a subset of trials from the Peabody Picture Vocabulary Test while seated at an eye-tracker monitor. During each trial, four colored images were presented on the monitor while a word was presented via audio recording. Participants verbally indicated which image they thought represented the target word. Words were categorized into Easy, Medium, and Hard difficulty. Pupil dilation during the Medium and Hard trials was significantly greater than during the Easy trials, though the Medium and Hard trials did not significantly differ from each other. Pupil dilation in comparison to trial accuracy presented a more complex pattern, with comparisons between accurate and inaccurate trials differing depending on the timing of the stimulus presentation. These results present further evidence that pupil dilation increases with cognitive effort associated with vocabulary tests, providing insights that could help refine vocabulary assessments and other related tests of language processing.

Pupil dilation reveals the intensity of touch.

Touch is important for many aspects of our daily activities. One of the most important tactile characteristics is its perceived intensity. However, quantifying the intensity of perceived tactile stimulation is not always possible using overt responses. Here, we show that pupil responses can objectively index the intensity of tactile stimulation in the absence of overt participant responses. In Experiment 1 (n = 32), we stimulated three reportedly differentially sensitive body locations (finger, forearm, and calf) with a single tap of a tactor while tracking pupil responses. Tactile stimulation resulted in greater pupil dilation than a baseline without stimulation. Furthermore, pupils dilated more for the more sensitive location (finger) than for the less sensitive location (forearm and calf). In Experiment 2 (n = 20) we extended these findings by manipulating the intensity of the stimulation with three different intensities, here a short vibration, always at the little finger. Again, pupils dilated more when being stimulated at higher intensities as compared to lower intensities. In summary, pupils dilated more for more sensitive parts of the body at constant stimulation intensity and for more intense stimulation at constant location. Taken together, the results show that the intensity of perceived tactile stimulation can be objectively measured with pupil responses - and that such responses are a versatile marker for touch research. Our findings may pave the way for previously impossible objective tests of tactile sensitivity, for example in minimally conscious state patients.

Emphasis on peripheral vision is accompanied by pupil dilation

People are best able to detect stimuli in peripheral vision when their pupils are large, and best able to discriminate stimuli in central vision when their pupils are small. However, it is unclear whether our visual system makes use of this by dilating the pupils when attention is directed towards peripheral vision. Therefore, throughout three experiments (N = 100), we tested whether pupil size adapts to the “breadth” of attention. We found that pupils dilate with increasing attentional breadth, both when attention is diffusely spread and when attention is directed at specific locations in peripheral vision. Based on our results and others, we propose that cognitively driven pupil dilation is not an epiphenomenal marker of locus coeruleus activity, as is often assumed, but rather is an adaptive response that reflects an emphasis on peripheral vision.

Individual differences in pupil dilation to others’ emotional and neutral eyes with varying pupil sizes

ABSTRACT Sensitivity to others’ emotional signals is an important factor for social interaction. While many studies of emotional reactivity focus on facial emotional expressions, signals such as pupil dilation which can indicate arousal, may also affect observers. For example, observers’ pupils dilate when viewing someone with dilated pupils, so-called pupillary contagion. Yet it is unclear how pupil size and emotional expression interact as signals. Further, examining individual differences in emotional reactivity to others can shed light on its mechanisms and potential outcomes. In the current study, adults’ (N = 453) pupil size was assessed while they viewed images of the eye region of individuals varying in emotional expression (neutral, happy, sad, fearful, angry) and pupil size (large, medium, small). Participants showed pupillary contagion regardless of the emotional expression. Individual differences in demographics (gender, age, socioeconomic status) and psychosocial factors (anxiety, depression, sleep problems) were also examined, yet the only factor related to pupillary contagion was socioeconomic status, with higher socioeconomic status predicting less pupillary contagion for emotionally-neutral stimuli. The results suggest that while pupillary contagion is a robust phenomenon, it can vary meaningfully across individuals.

Investigating effortful speech perception using fNIRS and pupillometry measures.

demonstrated that changes in pupil dilation were positively correlated with the self-reported task difficulty levels and negatively correlated with the task performance scores. A significant and negative correlation between the two behavioral measures was also found. That is, as perceived task demands increased and task performance scores decreased, pupils dilated more. fNIRS measures (cerebral oxygenation) in the left AC and LFC were both negatively correlated with the self-reported task difficulty levels and positively correlated with task performance scores. These results suggest that pupillometry measures can indicate task demands and listening effort; whereas, fNIRS measures using a similar paradigm seem to reflect speech processing, but not effort.

Enlargement of female pupils when perceiving something cute

It is reported that women’s pupils dilate when they see a baby; it is unclear if this pupillary response is caused by the perception of cuteness itself. Since many objects besides babies can be perceived as cute, we investigated whether the perception of cuteness, or the type of object observed, is related to pupil dilation. In the first experiment, female participants were requested to rate the subjective cuteness of greyscale pictures of objects such as animals and foods; their pupil sizes were measured. The results showed a significant positive correlation between perceived cuteness and participants’ pupil dilation. In the second experiment, participants rated the cuteness of images of female faces. Results revealed a significant negative correlation between perceived cuteness and pupil dilation. In our study, perceiving cuteness enlarged female observers’ pupils except when observing female faces. Positive reactions associated with cuteness may be premised on the existence of unconscious perceptual alterations and physical responses.

The cognitive workload of air pistol shooters on the aiming task

Background: Pupil dilations of the eye are known to correspond to central cognitive processes. Pupil size variations have been used for many years as a window on the mind, and various pupillary features have been associated with a number of cognitive processes Aim: The aim of this study is to examine levels of cognitive workload that occur during the aiming task of air-pistol shooting athletes according to the change that takes place in the pupils. Methods: Eight right-handed skilled male shooters (4 elite and 4 novice) were recruited in this study. A total of 320 shots - 160 dry (unscored) shots and 160 live (scored) shots - were recorded with a mobile eye-tracking device. The obtained pupil data were subjected to biometric analysis. The athletes’ pupil sizes were determined with the independent samples t-test according to whether the shot made was scored or unscored and whether the athletes were elite or novice. The athletes’ pupil size values at the beginning and end of the shot were obtained using the paired samples t-test. Results: On completion of the shot, the sport shooters’ pupils dilated on average by 2.85 mm at a rate of 32.5% compared to the situation at shot onset. Moreover, novice shooters’ pupils dilated more when firing scored shots than when firing unscored shots. According to the research results, it can be said that sports shooters engage in intensive cognitive processes while aiming and trigger pulling immediately before firing the shot. Novice shooters engage in more intensive cognitive processes when firing scored shots than when firing unscored shots. Conclusion: According to the results of the research, it can be said that in the sport of air pistol shooting, giving priority to dry (unscored) shooting training can contribute positively to the development of novice athletes’ aiming task skills and cognitive load capacity. Keywords: Cognitive workload, cognitive processes, eye tracking, air pistol, shooting

Pupil size in the evaluation of static and dynamic stimuli in peripheral vision.

It has been evidenced that in attention-window tasks, the participants fixate on the center of a screen while inspecting two stimuli that appear at the same time in parafoveal vision. Such tasks have successfully been used to estimate a person’s breadth of attention under various conditions. While behavioral investigations of visual attention have often made use of response accuracy, recent research has shown that the pupil size can also be used to track shifts of attention to the periphery. The main finding of previous studies is that the harder the evaluation of the stimuli becomes, e.g., because they appear farther away from the central fixation point, the stronger the pupils dilate. In this paper, we present experimental data suggesting that in an attention-window task, the pupil size can also be used to assess whether the participants attend to static, non-moving, or dynamic, moving stimuli. That is, regression models containing information on presentation mode (static vs. dynamic) and the visual angle between spatially separated stimuli better predict accuracy of perception and pupil dilation than model without these sources of information. This finding is useful for researchers who aim at understanding the human attentional system, including potential differences in its sensitivity to static and dynamic objects.

Replicating five pupillometry studies of Eckhard Hess

Several papers by Eckhard Hess from the 1960s and 1970s report that the pupils dilate or constrict according to the interest value, arousing content, or mental demands of visual stimuli. However, Hess mostly used small sample sizes and undocumented luminance control. In a first experiment (N = 182) and a second preregistered experiment (N = 147), we replicated five studies of Hess using modern equipment. Our experiments (1) did not support the hypothesis of gender differences in pupil diameter change with respect to baseline (PC) when viewing stimuli of different interest value, (2) showed that solving more difficult multiplications yields a larger PC in the seconds before providing an answer and a larger maximum PC, but a smaller PC at a fixed time after the onset of the multiplication, (3) did not support the hypothesis that participants’ PC mimics the pupil diameter in a pair of schematic eyes but not in single-eyed or three-eyed stimuli, (4) did not support the hypothesis of gender differences in PC when watching a video of a male trying to escape a mob, and (5) supported the hypothesis that arousing words yield a higher PC than non-arousing words. Although we did not observe consistent gender differences in PC, additional analyses showed gender differences in eye movements towards erogenous zones. Furthermore, PC strongly correlated with the luminance of the locations where participants looked. Overall, our replications confirm Hess’s findings that pupils dilate in response to mental demands and stimuli of an arousing nature. Hess’s hypotheses regarding pupil mimicry and gender differences in pupil dilation did not replicate.

Is accommodation a confounder in pupillometry research?

Much psychological research uses pupil diameter measurements to investigate the cognitive and emotional effects of visual stimuli. A potential problem is that accommodating at a nearby point causes the pupil to constrict. This study examined to what extent accommodation is a confounder in pupillometry research. Participants solved multiplication problems at different distances (Experiment 1) and looked at line drawings with different monocular depth cues (Experiment 2) while their pupil diameter, refraction, and vergence angle were recorded using a photorefractor. Experiment 1 showed that the pupils dilated while performing the multiplications, for all presentation distances. Pupillary constriction due to accommodation was not strong enough to override pupil dilation due to cognitive load. Experiment 2 showed that monocular depth cues caused a small shift in refraction in the expected direction. We conclude that, for the young student sample we used, pupil diameter measurements are not substantially affected by accommodation.

Non-spatial skills differ in the front and rear peri-personal space

In measuring behavioural and pupillary responses to auditory oddball stimuli delivered in the front and rear peri-personal space, we find that pupils dilate in response to rare stimuli, both target and distracters. Dilation in response to targets is stronger than the response to distracters, implying a task relevance effect on pupil responses. Crucially, pupil dilation in response to targets is also selectively modulated by the location of sound sources: stronger in the front than in the rear peri-personal space, in spite of matching behavioural performance. This supports the concept that even non-spatial skills, such as the ability to alert in response to behaviourally relevant events, are differentially engaged across subregions of the peri-personal space.

Pupillometric VoE paradigm reveals that 18- but not 10-month-olds spontaneously represent occluded objects (but not empty sets).

Object permanence has been investigated with a variety of paradigms and measures, yielding heterogeneous findings. The current study employed a novel Violation-of-Expectation paradigm measuring pupil dilation as indicator of cognitive effort and surprise. Across repeated trials, infants watched videos of animated toys either stopping in an open door frame or moving across the open door frame off screen. The door then closed and opened up again to reveal either the toy, or an empty space. In Experiment 1, 18-month-olds’s pupils dilated in response to the unexpected empty outcome more than to the expected empty outcome, establishing the paradigm as a suitable measure of violation of object expectation. Using the same paradigm, Experiment 2 revealed an absence of this object expectation effect for 10-month-olds. Results are discussed with regard to paradigmatic aspects and developmental differences. It is suggested that young infants do not automatically represent occluded objects upon perceiving occlusion events, and that occlusion events may initially require relevance in terms of individual activity or social interaction.

Pupil Dilation Is Sensitive to Semantic Ambiguity and Acoustic Degradation.

Speech comprehension is challenged by background noise, acoustic interference, and linguistic factors, such as the presence of words with more than one meaning (homonyms and homophones). Previous work suggests that homophony in spoken language increases cognitive demand. Here, we measured pupil dilation—a physiological index of cognitive demand—while listeners heard high-ambiguity sentences, containing words with more than one meaning, or well-matched low-ambiguity sentences without ambiguous words. This semantic-ambiguity manipulation was crossed with an acoustic manipulation in two experiments. In Experiment 1, sentences were masked with 30-talker babble at 0 and +6 dB signal-to-noise ratio (SNR), and in Experiment 2, sentences were heard with or without a pink noise masker at –2 dB SNR. Speech comprehension was measured by asking listeners to judge the semantic relatedness of a visual probe word to the previous sentence. In both experiments, comprehension was lower for high- than for low-ambiguity sentences when SNRs were low. Pupils dilated more when sentences included ambiguous words, even when no noise was added (Experiment 2). Pupil also dilated more when SNRs were low. The effect of masking was larger than the effect of ambiguity for performance and pupil responses. This work demonstrates that the presence of homophones, a condition that is ubiquitous in natural language, increases cognitive demand and reduces intelligibility of speech heard with a noisy background.

Supportive hand-holding attenuates pupillary responses to stress in adult couples.

BackgroundSocial relationships, particularly marriage, have been shown to ameliorate the potentially pathogenic impact of stressful events but prior research has been mostly aimed at downstream effects, with less research on real-time reactivity. Pupillometry is an innovative procedure that allows us to see the effects of acute stress in real time. The muscles that control pupil size are linked to the autonomic nervous system, so that when stressed, the pupils dilate; this occurs within 200ms. This quick response allows us to see the immediate effects of acute stress on the autonomic nervous system (ANS), and the real-time effects of social support in buffering stress.PurposeThe purpose of this study is to examine the dampening effects of received social support on the ANS’s pupillary response.MethodsEighty individuals (40 couples) were randomly assigned to either a spousal support (i.e., spouse hand-holding) or non-support condition (i.e., alone) and administered a Stroop task while pupil dilation was measured.ResultsThe Stroop task elicited a stress reaction in terms of pupil dilation in response to the incongruent task trials. Participants in the support condition showed accelerated habituation to the stress task (p < .001), and less pupil reactivity (p < .001) providing evidence for buffering effects of social support via spousal presence and hand-holding.ConclusionsThese results reveal the speed at which stress-buffering occurs, suggesting that pupillometry could be a good method to address the immediate dampening effects of social support.

Elaborate pupils in skates may help camouflage the eye.

The skate Leucoraja erinacea is a bottom-dweller that buries into the substrate with its eyes protruding, revealing elaborately shaped pupils. It has been suggested that such pupil shapes may camouflage the eye, yet this has never been tested. Here, we asked whether skate pupils dilate or constrict depending on background spatial frequency. In experiment 1, the skates' pupillary response to three artificial checkerboards of different spatial frequencies was recorded. Results showed that pupils did not change in response to spatial frequency. In experiment 2, in which skates buried into three natural substrates of different spatial frequencies, such that their eyes protruded, pupils showed a subtle but statistically significant response to changes in substrate spatial frequency. Although light intensity is the primary factor determining pupil dilation, our results show that pupils also change depending on the spatial frequency of natural substrates, which suggests that pupils may aid in camouflaging the eye.

Respiratory fluctuations in pupil diameter are not maintained during cognitive tasks

Pupil diameter fluctuation throughout the respiratory cycle is autonomically controlled in the resting state, as pupils dilate during inspiration and constrict during expiration. Furthermore, pupil size is differentially modulated by cognitive states between task engagement and disengagement. To determine whether respiratory-dependent fluctuations in pupil size are maintained during a cognitive task, we employed healthy human subjects performing a delayed matching-to-sample task with a short delay and measured their pupil sizes and R wave-to-R wave intervals (RRIs). We detected respiratory fluctuations in pupil size and the RRI during the delay period immediately before the discrimination stage of the task. During the discrimination stage, the cognitive state with the higher task engagement yielded more pupil dilation. However, respiratory fluctuations in pupil size were abolished, whereas those in the RRI were still discernible during the discrimination stage. Our results suggest that an alternative control mechanism involving the cognitive state associated with task engagement overrides the respiratory-related autonomic control of pupil diameter.

Pupil-linked arousal modulates behavior in rats performing a whisker deflection direction discrimination task.

Non-luminance-mediated changes in pupil size have been widely used to index arousal state. Recent animal studies have demonstrated correlations between behavioral state-related pupil dynamics and sensory processing. However, the relationship between pupil-linked arousal and behavior in animals performing perceptual tasks has not been fully elucidated. In the present study, we trained head-fixed rats to discriminate between directions of whisker movements using a Go/No-Go discrimination paradigm while imaging their pupils. Reaction times in this discrimination task were significantly slower than in previously reported detection tasks with similar setup, suggesting that discrimination required an increased cognitive load. We found the pupils dilated for all trials following stimulus presentation. Interestingly, in correct rejection trials, where pupil dilations solely resulted from cognitive processing, dilations were larger for more difficult stimuli. Baseline pupil size before stimulus presentation strongly correlated with behavior, as perceptual sensitivity peaked at intermediate pupil baselines and reaction time was fastest at large baselines. We further explored these relationships by investigating to what extent pupil baseline was predictive of upcoming behavior and found that a Bayesian decoder had significantly greater-than-chance probability in correctly predicting behavioral outcomes. Moreover, the outcome of the previous trial showed a strong correlation with behavior on present trials. Animals were more liberal and faster in responding following hit trials, whereas perceptual sensitivity was greatest following correct rejection trials. Taken together, these results suggest a tight correlation between pupil dynamics, perceptual performance, and reaction time in behaving rats, all of which are modulated by fluctuating arousal state. NEW & NOTEWORTHY In this study, we for the first time demonstrated that head-fixed rats were able to discriminate different directions of whisker movement. Interestingly, we found that the pupil dilated more when discriminating more difficult stimuli, a phenomenon reported in human subjects but not in animals. Baseline pupil size before stimulus presentation was found to strongly correlate with behavior, and a Bayesian decoder had significantly greater-than-chance probability in correctly predicting behavioral outcomes based on the baseline pupil size.

Pupillary Responses to Robotic and Human Emotions: The Uncanny Valley and Media Equation Confirmed.

Physiological responses during human–robots interaction are useful alternatives to subjective measures of uncanny feelings for nearly humanlike robots (uncanny valley) and comparable emotional responses between humans and robots (media equation). However, no studies have employed the easily accessible measure of pupillometry to confirm the uncanny valley and media equation hypotheses, evidence in favor of the existence of these hypotheses in interaction with emotional robots is scarce, and previous studies have not controlled for low level image statistics across robot appearances. We therefore recorded pupil size of 40 participants that viewed and rated pictures of robotic and human faces that expressed a variety of basic emotions. The robotic faces varied along the dimension of human likeness from cartoonish to humanlike. We strictly controlled for confounding factors by removing backgrounds, hair, and color, and by equalizing low level image statistics. After the presentation phase, participants indicated to what extent the robots appeared uncanny and humanlike, and whether they could imagine social interaction with the robots in real life situations. The results show that robots rated as nearly humanlike scored higher on uncanniness, scored lower on imagined social interaction, evoked weaker pupil dilations, and their emotional expressions were more difficult to recognize. Pupils dilated most strongly to negative expressions and the pattern of pupil responses across emotions was highly similar between robot and human stimuli. These results highlight the usefulness of pupillometry in emotion studies and robot design by confirming the uncanny valley and media equation hypotheses.

Human eyes with dilated pupils induce pupillary contagion in infants

Being sensitive and responsive to others’ internal states is critical for social life. One reliable cue to what others might be feeling is pupil dilation because it is linked to increases in arousal. When adults view an individual with dilated pupils, their pupils dilate in response, suggesting not only sensitivity to pupil size, but a corresponding response as well. However, little is known about the origins or mechanism underlying this phenomenon of pupillary contagion. Here we show that 4- to 6-month-old infants show pupillary contagion when viewing photographs of eyes with varying pupil sizes: their pupils dilate in response to others’ large, but not small or medium pupils. The results suggest that pupillary contagion is likely driven by a transfer of arousal and that it is present very early in life in human infants, supporting the view that it could be an adaptation fundamental for social and emotional development.