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2024 |
Eser Sendesen; Didem Turkyilmaz Listening handicap in tinnitus patients by controlling extended high frequencies - Effort or fatigue? Journal Article In: Auris Nasus Larynx, vol. 51, no. 1, pp. 198–205, 2024. @article{Sendesen2024, Objective: In previous studies, the results regarding the presence of listening effort or fatigue in tinnitus patients were inconsistent. The reason for this inconsistency could be that extended high frequencies, which can cause listening handicap, were not considered. Therefore, this study aimed to evaluate the listening skills in tinnitus patients by matching the hearing thresholds at all frequencies, including the extended high frequency. Methods: Eighteen chronic tinnitus patients and thirty matched healthy controls having normal pure-tone average with symmetrical hearing thresholds was included. Subjects were evaluated with 0.125-20 kHz pure-tone audiometry, Montreal cognitive assessment test (MoCA), Tinnitus Handicap Inventory (THI), Matrix Test, Pupillometry. Results: Pupil dilatation in the 'coding' phase of the sentence presented in tinnitus patients was less than in the control group (p<0.05). There was no difference between the groups for Matrix test scores (p> 0.05) Also, there was no statistically significant correlation between THI and Pupillometry components nor between MoCA (p>0.05). Conclusion: The results were interpreted for potential listening fatigue in tinnitus patients. Considering the possible listening handicap in tinnitus patients, reducing the listening difficulties especially in noisy environments, can be added to the goals of tinnitus therapy protocols. |
Damian Koevoet; Marnix Naber; Stefan Stigchel The intensity of internal and external attention assessed with pupillometry Journal Article In: Journal of Cognition, vol. 7, pp. 1–10, 2024. @article{Koevoet2024, Not only is visual attention shifted to objects in the external world, attention can also be directed to objects in memory. We have recently shown that pupil size indexes how strongly items are attended externally, which was reflected in more precise encoding into visual working memory. Using a retro-cuing paradigm, we here replicated this finding by showing that stronger pupil constrictions during encoding were reflective of the depth of encoding. Importantly, we extend this previous work by showing that pupil size also revealed the intensity of internal attention toward content stored in visual working memory. Specifically, pupil dilation during the prioritization of one among multiple internally stored representations predicted the precision of the prioritized item. Furthermore, the dynamics of the pupillary responses revealed that the intensity of internal and external attention independently determined the precision of internalized visual representations. Our results show that both internal and external attention are not all-or-none processes, but should rather be thought of as continuous resources that can be deployed at varying intensities. The employed pupillometric approach allows to unravel the intricate interplay between internal and external attention and their effects on visual working memory. |
H. Ershaid; M. Lizarazu; D. J. McLaughlin; M. Cooke; O. Simantiraki; M. Koutsogiannaki; M. Lallier Contributions of listening effort and intelligibility to cortical tracking of speech in adverse listening conditions Journal Article In: Cortex, vol. 172, pp. 54–71, 2024. @article{Ershaid2024, Cortical tracking of speech is vital for speech segmentation and is linked to speech intelligibility. However, there is no clear consensus as to whether reduced intelligibility leads to a decrease or an increase in cortical speech tracking, warranting further investigation of the factors influencing this relationship. One such factor is listening effort, defined as the cognitive resources necessary for speech comprehension, and reported to have a strong negative correlation with speech intelligibility. Yet, no studies have examined the relationship between speech intelligibility, listening effort, and cortical tracking of speech. The aim of the present study was thus to examine these factors in quiet and distinct adverse listening conditions. Forty-nine normal hearing adults listened to sentences produced casually, presented in quiet and two adverse listening conditions: cafeteria noise and re- verberant speech. Electrophysiological responses were registered with electroencephalogram, and listening effort was estimated subjectively using self-reported scores and objectively using pupillometry. Results indicated varying impacts of adverse conditions on intelligibility, listening effort, and cortical tracking of speech, depending on the preservation of the speech temporal envelope. The more distorted envelope in the reverberant condition led to higher listening effort, as reflected in higher subjective scores, increased pupil diameter, and stronger cortical tracking of speech in the delta band. These findings suggest that using measures of listening effort in addition to those of intelligibility is useful for interpreting cortical tracking of speech results. Moreover, reading and phonological skills of participants were positively correlated with listening effort in the cafeteria condition, suggesting a special role of expert language skills in processing speech in this noisy condition. Implications for future research and theories linking atypical cortical tracking of speech and reading disorders are further discussed. |
Andy Brendler; Max Schneider; Immanuel G. Elbau; Rui Sun; Taechawidd Nantawisarakul; Dorothee Pöhlchen; Tanja Brückl; A. K. Brem; E. B. Binder; A. Erhardt; J. Fietz; N. C. Grandi; Y. Kim; S. Ilić-Ćoćić; L. Leuchs; S. Lucae; T. Namendorf; J. Pape; L. Schilbach; I. Mücke-Heim; J. Ziebula; Michael Czisch; Philipp G. Sämann; Michael D. Lee; Victor I. Spoormaker In: Scientific Reports, vol. 14, no. 344, pp. 1–11, 2024. @article{Brendler2024, Major depressive disorder (MDD) is a devastating and heterogenous disorder for which there are no approved biomarkers in clinical practice. We recently identified anticipatory hypo-arousal indexed by pupil responses as a candidate mechanism subserving depression symptomatology. Here, we conducted a replication and extension study of these findings. We analyzed a replication sample of 40 unmedicated patients with a diagnosis of depression and 30 healthy control participants, who performed a reward anticipation task while pupil responses were measured. Using a Bayesian modelling approach taking measurement uncertainty into account, we could show that the negative correlation between pupil dilation and symptom load during reward anticipation is replicable within MDD patients, albeit with a lower effect size. Furthermore, with the combined sample of 136 participants (81 unmedicated depressed and 55 healthy control participants), we further showed that reduced pupil dilation in anticipation of reward is inversely associated with anhedonia items of the Beck Depression Inventory in particular. Moreover, using simultaneous fMRI, particularly the right anterior insula as part of the salience network was negatively correlated with depressive symptom load in general and anhedonia items specifically. The present study supports the utility of pupillometry in assessing noradrenergically mediated hypo-arousal during reward anticipation in MDD, a physiological process that appears to subserve anhedonia. |
2023 |
Laoura Ziaka; Athanassios Protopapas Cognitive control beyond single-item tasks: insights from pupillometry, gaze, and behavioral measures Journal Article In: Journal of Experimental Psychology: Human Perception and Performance, vol. 49, no. 7, pp. 968–988, 2023. @article{Ziaka2023, Cognitive control has been typically examined using single-item tasks. This has implications for the generalizability of theories of control implementation. Previous studies have revealed that different control demands are posed by tasks depending on whether they present stimuli individually (i.e., single-item) or simultaneously in array format (i.e., multi-item). In the present study, we tracked within-task performance in single-item and multi-item Stroop tasks using simultaneous pupillometry, gaze, and behavioral response measures, aiming to explore the implications of format differences for cognitive control. The results indicated within-task performance decline in the multi-item version of the Stroop task, accompanied by pupil constriction and dwell time increase, in both the incongruent and the neutral condition. In contrast, no performance decline or dwell time increase was observed in the course of the single-item version of the task.We interpret these findings in terms of capacity constraints on cognitive control, with implications for cognitive control research, and highlight the need for better understanding of the cognitive demands of multi-item tasks. |
Junyi Zhou; Wenjie Zhuang Physically active undergraduates perform better on executive-related oculomotor control: Evidence from the antisaccade task and pupillometry Journal Article In: PsyCh Journal, vol. 12, no. 1, pp. 17–24, 2023. @article{Zhou2023e, Previous studies have shown that exercise can improve executive function in young and older adults. However, it remains controversial whether a sufficient amount of physical activity leads to higher-level executive function. To examine the effect of physical activity on executive function, we used eye-tracking technology and the antisaccade task in 41 young undergraduates with various levels of physical activity. Moreover, we also investigated their differences in cognitive ability by examining their pupil size during the antisaccade task. Eye-tracking results showed that physically active individuals showed shorter saccade latency and higher accuracy in the antisaccade task than their physically inactive counterparts. Furthermore, the former showed larger pupil size during the preparatory period of antisaccade. These findings suggest that individuals with higher-level physical activity have higher-level executive function. The larger pupil sizes of physically active individuals may imply that their locus coeruleus-norepinephrine system and executive-related prefrontal cortex are more active, which contributes to their higher-level cognitive ability. |
Yao Yao; Katrina Connell; Stephen Politzer-Ahles Hearing emotion in two languages: A pupillometry study of Cantonese–Mandarin bilinguals' perception of affective cognates in L1 and L2 Journal Article In: Bilingualism: Language and Cognition, vol. 26, no. 4, pp. 795–808, 2023. @article{Yao2023d, Differential affective processing has been widely documented for bilinguals: L1 affective words elicit higher levels of arousal and stronger emotionality ratings than L2 affective words (Pavlenko, 2012). In this study, we focus on two closely related Chinese languages, Mandarin and Cantonese, whose affective lexicons are highly overlapping, with shared lexical items that only differ in pronunciation across languages. We recorded L1 Cantonese – L2 Mandarin bilinguals' pupil responses to auditory tokens of Cantonese and Mandarin affective words. Our results showed that Cantonese–Mandarin bilinguals had stronger pupil responses when the affective words were pronounced in Cantonese (L1) than when the same words were pronounced in Mandarin (L2). The effect was most evident in taboo words and among bilinguals with lower L2 proficiency. We discuss the theoretical implications of the findings in the frameworks of exemplar theory and models of the bilingual lexicon. |
Yao Yao; Katrina Connell; Stephen Politzer-Ahles Hearing emotion in two languages : A pupillometry study of Cantonese – Mandarin bilinguals ' perception of affective cognates in L1 and L2 Journal Article In: Bilingualism: Language and Cognition, pp. 1–14, 2023. @article{Yao2023, Differential affective processing has been widely documented for bilinguals: L1 affective words elicit higher levels of arousal and stronger emotionality ratings than L2 affective words (Pavlenko, 2012). In this study, we focus on two closely related Chinese languages, Mandarin and Cantonese, whose affective lexicons are highly overlapping, with shared lexical items that only differ in pronunciation across languages. We recorded L1 Cantonese – L2 Mandarin bilinguals' pupil responses to auditory tokens of Cantonese and Mandarin affective words. Our results showed that Cantonese–Mandarin bilinguals had stronger pupil responses when the affective words were pronounced in Cantonese (L1) than when the same words were pronounced in Mandarin (L2). The effect was most evident in taboo words and among bilinguals with lower L2 proficiency. We discuss the theoretical implications of the findings in the frameworks of exemplar theory and models of the bilingual lexicon. © |
Weizhen Xie; Weiwei Zhang Pupillary evidence reveals the influence of conceptual association on brightness perception Journal Article In: Psychonomic Bulletin & Review, vol. 30, no. 4, pp. 1388–1395, 2023. @article{Xie2023a, Our visual experience often varies based on momentary thoughts and feelings. For example, when positive concepts are invoked, visual objects may appear brighter (e.g., a “brighter” smile). However, it remains unclear whether this phenomenological experience is driven by a genuine top-down modulation of brightness perception or by a mere response bias. To investigate this issue, we use pupillometry as a more objective measure of perceived brightness. We asked participants to judge the brightness level of an isoluminant gray color patch after evaluating the valence of a positive or negative word. We found that the gray color patch elicited greater pupillary light reflex and more frequent “brighter” responses after observers had evaluated the valence of a positive word. As pupillary light reflex is unlikely driven by voluntary control, these results suggest that the conceptual association between affect and luminance can modulate brightness perception. |
Christian Wienke; Marcus Grueschow; Aiden Haghikia; Tino Zaehle In: Journal of Neuroscience, vol. 43, no. 36, pp. 6306–6319, 2023. @article{Wienke2023, Transcutaneous auricular vagus nerve stimulation (taVNS) has been proposed to activate the locus ceruleus-noradrenaline (LC-NA) system. However, previous studies failed to find consistent modulatory effects of taVNS on LC-NA biomarkers. Previous studies suggest that phasic taVNS may be capable of modulating LC-NA biomarkers such as pupil dilation and alpha oscillations. However, it is unclear whether these effects extend beyond pure sensory vagal nerve responses. Critically, the potential of the pupillary light reflex as an additional taVNS biomarker has not been explored so far. Here, we applied phasic active and sham taVNS in 29 subjects (16 female, 13 male) while they performed an emotional Stroop task (EST) and a passive pupil light reflex task (PLRT). We recorded pupil size and brain activity dynamics using a combined Magnetoencephalography (MEG) and pupillometry design. Our results show that phasic taVNS significantly increased pupil dilation and performance during the EST. During the PLRT, active taVNS reduced and delayed pupil constriction. In the MEG, taVNS increased frontal-midline theta and alpha power during the EST, whereas occipital alpha power was reduced during both the EST and PLRT. Our findings provide evidence that phasic taVNS systematically modulates behavioral, pupillary, and electrophysiological parameters of LC-NA activity during cognitive processing. Moreover, we demonstrate for the first time that the pupillary light reflex can be used as a simple and effective proxy of taVNS efficacy. These findings have important implications for the development of noninvasive neuromodulation interventions for various cognitive and clinical applications. |
Sarah Villard; Tyler K. Perrachione; Sung-Joo Lim; Ayesha Alam; Gerald Kidd In: The Journal of the Acoustical Society of America, vol. 154, no. 2, pp. 1152–1167, 2023. @article{Villard2023, The task of processing speech masked by concurrent speech/noise can pose a substantial challenge to listeners. However, performance on such tasks may not directly reflect the amount of listening effort they elicit. Changes in pupil size and neural oscillatory power in the alpha range (8–12 Hz) are prominent neurophysiological signals known to reflect listening effort; however, measurements obtained through these two approaches are rarely correlated, suggesting that they may respond differently depending on the specific cognitive demands (and, by extension, the specific type of effort) elicited by specific tasks. This study aimed to compare changes in pupil size and alpha power elicited by different types of auditory maskers (highly confusable intelligible speech maskers, speech-envelope-modulated speech-shaped noise, and unmodulated speech-shaped noise maskers) in young, normal-hearing listeners. Within each condition, the target-to-masker ratio was set at the participant's individually estimated 75% correct point on the psychometric function. The speech masking condition elicited a significantly greater increase in pupil size than either of the noise masking conditions, whereas the unmodulated noise masking condition elicited a significantly greater increase in alpha oscillatory power than the speech masking condition, suggesting that the effort needed to solve these respective tasks may have different neural origins. |
Anna Cornelia Stausberg Who are you to judge? Investigating narcissism through pupil dilation at witness testimonials Journal Article In: Psychology, vol. 14, no. 02, pp. 144–157, 2023. @article{Stausberg2023, Various literature has explored narcissistic behaviour and its distinct emotional and aggressive nature. However, there is a significant gap in research when exploring the physiological differences of individual classified as having high narcissistic traits and tendencies. Pupillometry is a measure often utilised to measure distinct differences in emotional arousal. This measure allows for excellent insight into physiological responses to various stimuli presented. This exploratory study was set up to investigate pupillometry responses to auditory stimulation in narcissistic versus control participants. Findings were consistent with previous research; however, various limitations hindered significant findings. This pilot study is a guideline for future research as it is a first attempt at exploring a physiological relationship between emotionality and narcissism in the context of a criminal hearing. |
Olympia Simantiraki; Anita E. Wagner; Martin Cooke The impact of speech type on listening effort and intelligibility for native and non-native listeners Journal Article In: Frontiers in Neuroscience, vol. 17, pp. 1–16, 2023. @article{Simantiraki2023, Listeners are routinely exposed to many different types of speech, including artificially-enhanced and synthetic speech, styles which deviate to a greater or lesser extent from naturally-spoken exemplars. While the impact of differing speech types on intelligibility is well-studied, it is less clear how such types affect cognitive processing demands, and in particular whether those speech forms with the greatest intelligibility in noise have a commensurately lower listening effort. The current study measured intelligibility, self-reported listening effort, and a pupillometry-based measure of cognitive load for four distinct types of speech: (i) plain i.e. natural unmodified speech; (ii) Lombard speech, a naturally-enhanced form which occurs when speaking in the presence of noise; (iii) artificially-enhanced speech which involves spectral shaping and dynamic range compression; and (iv) speech synthesized from text. In the first experiment a cohort of 26 native listeners responded to the four speech types in three levels of speech-shaped noise. In a second experiment, 31 non-native listeners underwent the same procedure at more favorable signal-to-noise ratios, chosen since second language listening in noise has a more detrimental effect on intelligibility than listening in a first language. For both native and non-native listeners, artificially-enhanced speech was the most intelligible and led to the lowest subjective effort ratings, while the reverse was true for synthetic speech. However, pupil data suggested that Lombard speech elicited the lowest processing demands overall. These outcomes indicate that the relationship between intelligibility and cognitive processing demands is not a simple inverse, but is mediated by speech type. The findings of the current study motivate the search for speech modification algorithms that are optimized for both intelligibility and listening effort. |
Eser Sendesen; Samet Kılıç; Nurhan Erbil; Özgür Aydın; Didem Turkyilmaz An exploratory study of the effect of tinnitus on listening effort using EEG and pupillometry Journal Article In: Otolaryngology-Head and Neck Surgery, vol. 169, no. 5, pp. 1259–1267, 2023. @article{Sendesen2023, Objective: Previous behavioral studies on listening effort in tinnitus patients did not consider extended high-frequency hearing thresholds and had conflicting results. This inconsistency may be related that listening effort is not evaluated by the central nervous system (CNS) and autonomic nervous system (ANS), which are directly related to tinnitus pathophysiology. This study matches hearing thresholds at all frequencies, including the extended high-frequency and reduces hearing loss to objectively evaluate listening effort over the CNS and ANS simultaneously in tinnitus patients. Study Design: Case-control study. Setting: University hospital. Methods: Sixteen chronic tinnitus patients and 23 matched healthy controls having normal pure-tone averages with symmetrical hearing thresholds were included. Subjects were evaluated with 0.125 to 20 kHz pure-tone audiometry, Montreal Cognitive Assessment Test (MoCA), Tinnitus Handicap Inventory (THI), Visual Analog Scale (VAS), electroencephalography (EEG), and pupillometry. Results: Pupil dilation and EEG alpha band in the “coding” phase of the sentence presented in tinnitus patients was less than in the control group (p <.05). VAS score was higher in the tinnitus group (p <.01). Also, there was no statistically significant relationship between EEG and pupillometry components and THI or MoCA (p >.05). Conclusion: This study suggests that tinnitus patients may need to make an extra effort to listen. Also, pupillometry may not be sufficiently reliable to assess listening effort in ANS-related pathologies. Considering the possible listening difficulties in tinnitus patients, reducing the listening difficulties, especially in noisy environments, can be added to the goals of tinnitus therapy protocols. |
Hannah Mechtenberg; Cristal Giorio; Emily B. Myers Pupil dilation reflects perceptual priorities during a receptive speech task Journal Article In: Ear & Hearing, pp. 1–16, 2023. @article{Mechtenberg2023, Objectives: The listening demand incurred by speech perception fluctu- ates in normal conversation. At the acoustic-phonetic level, natural varia- tion in pronunciation acts as speedbumps to accurate lexical selection. Any given utterance may be more or less phonetically ambiguous—a problem that must be resolved by the listener to choose the correct word. This becomes especially apparent when considering two common speech registers—clear and casual—that have characteristically differ- ent levels of phonetic ambiguity. Clear speech prioritizes intelligibility through hyperarticulation which results in less ambiguity at the pho- netic level, while casual speech tends to have a more collapsed acoustic space. We hypothesized that listeners would invest greater cognitive resources while listening to casual speech to resolve the increased amount of phonetic ambiguity, as compared with clear speech. To this end, we used pupillometry as an online measure of listening effort dur- ing perception of clear and casual continuous speech in two background conditions: quiet and noise. Design: Forty-eight participants performed a probe detection task while listening to spoken, nonsensical sentences (masked and unmasked) while recording pupil size. Pupil size was modeled using growth curve analysis to capture the dynamics of the pupil response as the sentence unfolded. Results: Pupil size during listening was sensitive to the presence of noise and speech register (clear/casual). Unsurprisingly, listeners had over- all larger pupil dilations during speech perception in noise, replicating earlier work. The pupil dilation pattern for clear and casual sentences was considerably more complex. Pupil dilation during clear speech tri- als was slightly larger than for casual speech, across quiet and noisy backgrounds. Conclusions: We suggest that listener motivation could explain the larger pupil dilations to clearly spoken speech. We propose that, bounded by the context of this task, listeners devoted more resources to perceiving the speech signal with the greatest acoustic/phonetic fidelity. Further, we unexpectedly found systematic differences in pupil dilation preceding the onset of the spoken sentences. Together, these data demonstrate that the pupillary system is not merely reactive but also adaptive—sensitive to both task structure and listener motivation to maximize accurate per- ception in a limited resource system. |
Drew J. McLaughlin; Maggie E. Zink; Lauren Gaunt; Jamie Reilly; Mitchell S. Sommers; Kristin J. Van Engen; Jonathan E. Peelle Give me a break! Unavoidable fatigue effects in cognitive pupillometry Journal Article In: Psychophysiology, vol. 60, no. 7, pp. 1–20, 2023. @article{McLaughlin2023a, Pupillometry has a rich history in the study of perception and cognition. One perennial challenge is that the magnitude of the task-evoked pupil response diminishes over the course of an experiment, a phenomenon we refer to as a fatigue effect. Reducing fatigue effects may improve sensitivity to task effects and reduce the likelihood of confounds due to systematic physiological changes over time. In this paper, we investigated the degree to which fatigue effects could be ameliorated by experimenter intervention. In Experiment 1, we assigned participants to one of three groups—no breaks, kinetic breaks (playing with toys, but no social interaction), or chatting with a research assistant—and compared the pupil response across conditions. In Experiment 2, we additionally tested the effect of researcher observation. Only breaks including social interaction significantly reduced the fatigue of the pupil response across trials. However, in all conditions we found robust evidence for fatigue effects: that is, regardless of protocol, the task-evoked pupil response was substantially diminished (at least 60%) over the duration of the experiment. We account for the variance of fatigue effects in our pupillometry data using multiple common statistical modeling approaches (e.g., linear mixed-effects models of peak, mean, and baseline pupil diameters, as well as growth curve models of time-course data). We conclude that pupil attenuation is a predictable phenomenon that should be accommodated in our experimental designs and statistical models. |
Drew J. McLaughlin; Jackson S. Colvett; Julie M. Bugg; Kristin J. Van Engen Sequence effects and speech processing: Cognitive load for speaker-switching within and across accents Journal Article In: Psychonomic Bulletin & Review, pp. 1–11, 2023. @article{McLaughlin2023, Prior work in speech processing indicates that listening tasks with multiple speakers (as opposed to a single speaker) result in slower and less accurate processing. Notably, the trial-to-trial cognitive demands of switching between speakers or switching between accents have yet to be examined. We used pupillometry, a physiological index of cognitive load, to examine the demands of processing first (L1) and second (L2) language-accented speech when listening to sentences produced by the same speaker consecutively (no switch), a novel speaker of the same accent (within-accent switch), and a novel speaker with a different accent (across-accent switch). Inspired by research on sequential adjustments in cognitive control, we aimed to identify the cognitive demands of accommodating a novel speaker and accent by examining the trial-to-trial changes in pupil dilation during speech processing. Our results indicate that switching between speakers was more cognitively demanding than listening to the same speaker consecutively. Additionally, switching to a novel speaker with a different accent was more cognitively demanding than switching between speakers of the same accent. However, there was an asymmetry for across-accent switches, such that switching from an L1 to an L2 accent was more demanding than vice versa. Findings from the present study align with work examining multi-talker processing costs, and provide novel evidence that listeners dynamically adjust cognitive processing to accommodate speaker and accent variability. We discuss these novel findings in the context of an active control model and auditory streaming framework of speech processing. |
Jacie R. McHaney; William L. Schuerman; Matthew K. Leonard; Bharath Chandrasekaran Transcutaneous auricular vagus nerve stimulation modulates performance but not pupil size during nonnative speech category learning Journal Article In: Journal of Speech, Language, and Hearing Research, vol. 66, no. 10, pp. 3825–3843, 2023. @article{McHaney2023, Purpose: Subthreshold transcutaneous auricular vagus nerve stimulation (taVNS) synchronized with behavioral training can selectively enhance nonnative speech category learning in adults. Prior work has demonstrated that behavioral performance increases when taVNS is paired with easier-to-learn Mandarin tone categories in native English listeners, relative to when taVNS is paired with harder-to-learn Mandarin tone categories or without taVNS. Mechanistically, this temporally precise plasticity has been attributed to noradrenergic modulation. However, prior work did not specifically utilize methodologies that indexed nor-adrenergic modulation and, therefore, was unable to explicitly test this hypothesis. Our goal for this study was to use pupillometry to gain mechanistic insights into taVNS behavioral effects. Method: Thirty-eight participants learned to categorize Mandarin tones while pupil-lometry was recorded. In a double-blinded design, participants were divided into two taVNS groups that, as in the prior study, differed according to whether taVNS was paired with easier-to-learn tones or harder-to-learn tones. Learning performance and pupillary responses were measured using linear mixed-effects models. Results: We found that taVNS did not have any tone-specific or group behav-ioral or pupillary effects. However, in an exploratory analysis, we observed that taVNS did lead to faster rates of learning on trials paired with stimulation, particularly for those who were stimulated at lower amplitudes. Conclusions: Our results suggest that pupillary responses may not be a reliable marker of locus coeruleus–norepinephrine system activity in humans. However, future research should systematically examine the effects of stimulation amplitude on both behavior and pupillary responses. |
Audrey Mazancieux; Franck Mauconduit; Alexis Amadon; Jan Willem de Gee; Tobias H. Donner; Florent Meyniel Brainstem fMRI signaling of surprise across different types of deviant stimuli Journal Article In: Cell Reports, vol. 42, no. 11, pp. 1–15, 2023. @article{Mazancieux2023, Detection of deviant stimuli is crucial to orient and adapt our behavior. Previous work shows that deviant stimuli elicit phasic activation of the locus coeruleus (LC), which releases noradrenaline and controls central arousal. However, it is unclear whether the detection of behaviorally relevant deviant stimuli selectively triggers LC responses or other neuromodulatory systems (dopamine, serotonin, and acetylcholine). We combine human functional MRI (fMRI) recordings optimized for brainstem imaging with pupillometry to perform a mapping of deviant-related responses in subcortical structures. Participants have to detect deviant items in a “local-global” paradigm that distinguishes between deviance based on the stimulus probability and the sequence structure. fMRI responses to deviant stimuli are distributed in many cortical areas. Both types of deviance elicit responses in the pupil, LC, and other neuromodulatory systems. Our results reveal that the detection of task-relevant deviant items recruits the same multiple subcortical systems across computationally different types of deviance. |
Sebastiaan Mathôt; Ana Vilotijević Methods in cognitive pupillometry: Design, preprocessing, and statistical analysis Journal Article In: Behavior Research Methods, vol. 55, no. 6, pp. 3055–3077, 2023. @article{Mathot2023a, Cognitive pupillometry is the measurement of pupil size to investigate cognitive processes such as attention, mental effort, working memory, and many others. Currently, there is no commonly agreed-upon methodology for conducting cognitive-pupillometry experiments, and approaches vary widely between research groups and even between different experiments from the same group. This lack of consensus makes it difficult to know which factors to consider when conducting a cognitive-pupillometry experiment. Here we provide a comprehensive, hands-on guide to methods in cognitive pupillometry, with a focus on trial-based experiments in which the measure of interest is the task-evoked pupil response to a stimulus. We cover all methodological aspects of cognitive pupillometry: experimental design, preprocessing of pupil-size data, and statistical techniques to deal with multiple comparisons when testing pupil-size data. In addition, we provide code and toolboxes (in Python) for preprocessing and statistical analysis, and we illustrate all aspects of the proposed workflow through an example experiment and example scripts. |
Priscila López-Beltrán Heritage speakers' processing of the Spanish subjunctive: A pupillometric study Journal Article In: Linguistic Approaches to Bilingualism, pp. 1–47, 2023. @article{LopezBeltran2023, We investigated linguistic knowledge of subjunctive mood in heritage speakers of Spanish who live in a long-standing English-Spanish bilingual community in Albuquerque, New Mexico. Three experiments examine the constraints on subjunctive selection. Experiment 1 and Experiment 2 employed pupillometry to investigate heritage speakers' online sensitivity to the presence of the subjunctive with non-variable governors (Lexical conditioning) and with negated governors (Structural conditioning). Experiment 3 employed an elicited production task to examine production of subjunctive in the same contexts. The findings of the heritage group were compared to those of a group ofSpanish-dominant Mexican bilinguals. Results showed that in comprehension and production, heritage speakers were as sensitive as the Spanish-dominant bilinguals to the lexical and structural factors that condition mood selection. In comprehension, the two groups experienced an increased pupillary dilation in conditions where the indicative was used but the subjunctive was expected. In addition, high- frequency governors and irregular subordinate verbs boosted participants' sensitivity to the presence of the subjunctive. In production, there were no significant differences between heritage speakers and Spanish-dominant bilinguals when producing the subjunctive with non-variable and negated governors. |
Elke B. Lange; Lauren K. Fink Eye blinking, musical processing, and subjective states—A methods account Journal Article In: Psychophysiology, vol. 60, no. 10, pp. 1–20, 2023. @article{Lange2023, Affective sciences often make use of self-reports to assess subjective states. Seeking a more implicit measure for states and emotions, our study explored spontaneous eye blinking during music listening. However, blinking is understudied in the context of research on subjective states. Therefore, a second goal was to explore different ways of analyzing blink activity recorded from infra-red eye trackers, using two additional data sets from earlier studies differing in blinking and viewing instructions. We first replicate the effect of increased blink rates during music listening in comparison with silence and show that the effect is not related to changes in self-reported valence, arousal, or to specific musical features. Interestingly, but in contrast, felt absorption reduced participants' blinking. The instruction to inhibit blinking did not change results. From a methodological perspective, we make suggestions about how to define blinks from data loss periods recorded by eye trackers and report a data-driven outlier rejection procedure and its efficiency for subject-mean analyses, as well as trial-based analyses. We ran a variety of mixed effects models that differed in how trials without blinking were treated. The main results largely converged across accounts. The broad consistency of results across different experiments, outlier treatments, and statistical models demonstrates the reliability of the reported effects. As recordings of data loss periods come for free when interested in eye movements or pupillometry, we encourage researchers to pay attention to blink activity and contribute to the further understanding of the relation between blinking, subjective states, and cognitive processing. |
Charlene L. M. Lam; Tom J. Barry; Jenny Yiend; Tatia M. C. Lee The role of consciousness in threat extinction learning Journal Article In: Consciousness and Cognition, vol. 116, pp. 1–12, 2023. @article{Lam2023, Extinction learning is regarded as a core mechanism underlying exposure therapy. The extent to which learned threats can be extinguished without conscious awareness is a controversial and on-going debate. We investigated whether implicit vs. explicit exposure to a threatened stimulus can modulate defence responses measured using pupillometry. Healthy participants underwent a threat conditioning paradigm in which one of the conditioned stimuli (CS) was perceptually suppressed using continuous flash suppression (CFS). Participants' pupillary responses, CS pleasantness ratings, and trial-by-trial awareness of the CS were recorded. During Extinction, participants' pupils dilated more in the trials in which they were unaware of the CS than in those in which they were aware of it (Cohen's d = 0.57). After reinstatement, the percentage of fear recovery was greater for the CFS-suppressed CS than the CS with full awareness. The current study suggests that the modulation of fear responses by extinction with reduced visual awareness is weaker compared to extinction with full perceptual awareness. |
Damian Koevoet; Christoph Strauch; Marnix Naber; Stefan Van Stigchel The costs of paying overt and covert attention assessed with pupillometry Journal Article In: Psychological Science, vol. 34, no. 8, pp. 887–898, 2023. @article{Koevoet2023a, Attention can be shifted with or without an accompanying saccade (i.e., overtly or covertly, respectively). Thus far, it is unknown how cognitively costly these shifts are, yet such quantification is necessary to understand how and when attention is deployed overtly or covertly. In our first experiment (N = 24 adults), we used pupillometry to show that shifting attention overtly is more costly than shifting attention covertly, likely because planning saccades is more complex. We pose that these differential costs will, in part, determine whether attention is shifted overtly or covertly in a given context. A subsequent experiment (N = 24 adults) showed that relatively complex oblique saccades are more costly than relatively simple saccades in horizontal or vertical directions. This provides a possible explanation for the cardinal-direction bias of saccades. The utility of a cost perspective as presented here is vital to furthering our understanding of the multitude of decisions involved in processing and interacting with the external world efficiently. |
Damian Koevoet; Marnix Naber; Christoph Strauch; Rosyl S. Somai; Stefan Van der Stigchel Differential aspects of attention predict the depth of visual working memory encoding: Evidence from pupillometry Journal Article In: Journal of Vision, vol. 23, no. 6, pp. 1–11, 2023. @article{Koevoet2023, What determines how much one encodes into visual working memory? Traditionally, encoding depth is considered to be indexed by spatiotemporal properties of gaze, such as gaze position and dwell time. Although these properties inform about where and how long one looks, they do not necessarily inform about the current arousal state or how strongly attention is deployed to facilitate encoding. Here, we found that two types of pupillary dynamics predict how much information is encoded during a copy task. The task involved encoding a spatial pattern of multiple items for later reproduction. Results showed that smaller baseline pupil sizes preceding and stronger pupil orienting responses during encoding predicted that more information was encoded into visual working memory. Additionally, we show that pupil size reflects not only how much but also how precisely material is encoded. We argue that a smaller pupil size preceding encoding is related to increased exploitation, whereas larger pupil constrictions signal stronger attentional (re)orienting to the to-be-encoded pattern. Our findings support the notion that the depth of visual working memory encoding is the integrative outcome of differential aspects of attention: how alert one is, how much attention one deploys, and how long it is deployed. Together, these factors determine how much information is encoded into visual working memory. |
Jiae Kim; Jiyeon Lee; Sang Beom Jun; Jee Eun Sung Pupillometry as a window to detect cognitive aging in the brain Journal Article In: Biomedical Engineering Letters, pp. 91–101, 2023. @article{Kim2023b, This study investigated whether there are aging-related differences in pupil dilation (pupillometry) while the cognitive load is manipulated using digit- and word-span tasks. A group of 17 younger and 15 cognitively healthy older adults performed digit- and word-span tasks. Each task comprised three levels of cognitive loads with 10 trials for each level. For each task, the recall accuracy and the slope of pupil dilation were calculated and analyzed. The raw signal of measured pupil size was low-pass filtered and interpolated to eliminate blinking artifacts and spike noises. Two-way ANOVA was used for statistical analyses. For the recall accuracy, the significant group differences emerged as the span increases in digit-span (5- vs. 7-digit) and word-span (4- vs. 5-word) tasks, while the group differences were not significant on 3-digit- and 3-word-span tasks with lower cognitive load. In digit-span tasks, there was no aging-related difference in the slope of pupil dilation. However, in word-span tasks, the slope of pupil dilation differed significantly between two groups as cognitive load increased, indicating that older adults presented a higher pupil dilation slope than younger adults especially under the conditions with higher cognitive load. The current study found significant aging effects in the pupil dilations under the more cognitive demanding span tasks when the types of span tasks varied (e.g., digit vs. word). The manipulations successfully elicited differential aging effects, given that the aging effects became most salient under word-span tasks with greater cognitive load especially under the maximum length. |
Jeff Huang; Donald Brien; Brian C. Coe; Giulia Longoni; Donald J. Mabbott; Douglas P. Munoz; E. Ann Yeh Delayed oculomotor response associates with optic neuritis in youth with demyelinating disorders Journal Article In: Multiple Sclerosis and Related Disorders, vol. 79, pp. 1–9, 2023. @article{Huang2023a, Introduction: Impairment in visual and cognitive functions occur in youth with demyelinating disorders such as multiple sclerosis, neuromyelitis optica spectrum disorder, and myelin oligodendrocyte glycoprotein antibody-associated disease. Quantitative behavioral assessment using eye-tracking and pupillometry can provide functional metrics for important prognostic and clinically relevant information at the bedside. Methods: Children and adolescents diagnosed with demyelinating disorders and healthy, age-matched controls completed an interleaved pro- and anti-saccade task using video-based eye-tracking and underwent spectral-domain optical coherence tomography examination for evaluation of retinal nerve fiber layer and ganglion cell inner plexiform layer thickness. Low-contrast visual acuity and Symbol Digit Modalities Test were performed for visual and cognitive functional assessments. We assessed saccade and pupil parameters including saccade reaction time, direction error rate, pupil response latency, peak constriction time, and peak constriction and dilation velocities. Generalized Estimating Equations were used to examine the association of eye-tracking parameters with optic neuritis history, structural metrics, and visual and cognitive scores. Results: The study included 36 demyelinating disorders patients, aged 8–18 yrs. (75% F; median = 15.22 yrs. |
Hengda He; Linbi Hong; Paul Sajda Pupillary response is associated with the reset and switching of functional brain networks during salience processing Journal Article In: PLoS Computational Biology, vol. 19, no. 5, pp. 1–29, 2023. @article{He2023a, The interface between processing internal goals and salient events in the environment involves various top-down processes. Previous studies have identified multiple brain areas for salience processing, including the salience network (SN), dorsal attention network, and the locus coeruleus-norepinephrine (LC-NE) system. However, interactions among these systems in salience processing remain unclear. Here, we simultaneously recorded pupillometry, EEG, and fMRI during an auditory oddball paradigm. The analyses of EEG and fMRI data uncovered spatiotemporally organized target-associated neural correlates. By modeling the target-modulated effective connectivity, we found that the target-evoked pupillary response is associated with the network directional couplings from late to early subsystems in the trial, as well as the network switching initiated by the SN. These findings indicate that the SN might cooperate with the pupil-indexed LC-NE system in the reset and switching of cortical networks, and shed light on their implications in various cognitive processes and neurological diseases. |
Althea Frisanco; Marco Biella; Marco Brambilla; Mariska E. Kret All that meets the eye: The contribution of reward processing and pupil mimicry on pupillary reactions to facial trustworthiness Journal Article In: Current Psychology, vol. 42, no. 14, pp. 11685–11692, 2023. @article{Frisanco2023, The present work investigates pupillary reactions induced by exposure to faces with different levels of trustworthiness. Participants' (N = 69) pupillary changes were recorded while they viewed white male faces with a neutral expression varying on facial trustworthiness. Results suggest that reward processing and pupil mimicry are relevant mechanisms driving participants' pupil reactions. However, when including both factors in one statistical model, pupil mimicry seems to be a stronger predictor than reward processing of participants' pupil dilation. Results are discussed in light of pupillometry evidence. |
Julia Fietz; Dorothee Pöhlchen; Elisabeth B. Binder; Angelika Erhardt; Susanne Lucae; Norma C. Grandi; Tamara Namendorf; Immanuel Elbau; Laura Leuchs; Leonhard Schilbach; Sanja Ilić-Ćoćić; Julius Ziebula; Iven-Alex Mücke-Heim; Yeho Kim; Julius Pape; Tanja M. Brückl; Anna-Katharine Brem; Frank Padberg; Michael Czisch; Philipp G. Sämann; Victor I. Spoormaker In: Biological Psychiatry: Cognitive Neuroscience and Neuroimaging, no. 24, pp. 1–8, 2023. @article{Fietz2023, Background Neurocognitive 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. Methods Our 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. Results The 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. Conclusions Pupil 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. |
Jamal Esmaily; Sajjad Zabbah; Reza Ebrahimpour; Bahador Bahrami Interpersonal alignment of neural evidence accumulation to social exchange of confidence Journal Article In: eLife, vol. 12, pp. 1–27, 2023. @article{Esmaily2023, Private, subjective beliefs about uncertainty have been found to have idiosyncratic computational and neural substrates yet, humans share such beliefs seamlessly and cooperate successfully. Bringing together decision making under uncertainty and interpersonal alignment in communication, in a discovery plus pre-registered replication design, we examined the neuro-computational basis of the relationship between privately held and socially shared uncertainty. Examining confidence-speed-accuracy trade-off in uncertainty-ridden perceptual decisions under social vs isolated context, we found that shared (i.e. reported confidence) and subjective (inferred from pupillometry) uncertainty dynamically followed social information. An attractor neural network model incorporating social information as top-down additive input captured the observed behavior and demonstrated the emergence of social alignment in virtual dyadic simulations. Electroencephalography showed that social exchange of confidence modulated the neural signature of perceptual evidence accumulation in the central parietal cortex. Our findings offer a neural population model for interpersonal alignment of shared beliefs. |
Ciara Egan; Joshua S. Payne; Manon W. Jones In: Neuropsychologia, vol. 184, pp. 1–8, 2023. @article{Egan2023, Readers with developmental dyslexia are known to be impaired in representing and accessing phonology, but their ability to process meaning is generally considered to be intact. However, neurocognitive studies show evidence of a subtle semantic processing deficit in dyslexic readers, relative to their typically-developing peers. Here, we compared dyslexic and typical adult readers on their ability to judge semantic congruency (congruent vs. inconcongruent) in short, two-word phrases, which were further manipulated for phonological relatedness (alliterating vs. non-alliterating); “dazzling-diamond”; “sparkling-diamond”; “dangerous-diamond”; and “creepy-diamond”. At the level of behavioural judgement, all readers were less accurate when evaluating incongruent alliterating items compared with incongruent non-aliterating, suggesting that phonological patterning creates the illusion of semantic congruency (as per Egan et al., 2020). Dyslexic readers showed a similar propensity for this form-meaning relationship despite a phonological processing impairment as evidenced in the cognitive and literacy indicative assessments. Dyslexic readers also showed an overall reduction in the ability to accurately judge semantic congruency, suggestive of a subtle semantic impairment. Whilst no group differences emerged in the electrophysiological measures, our pupil dilation measurements revealed a global tendency for dyslexic readers to manifest a reduced attentional response to these word stimuli, compared with typical readers. Our results show a broad manifestation of neurocognitive differences in adult dyslexic and typical readers' processing of print, at the level of autonomic arousal as well as in higher level semantic judgements. |
Tjerk T. Dercksen; Andreas Widmann; Nicole Wetzel Salient omissions—pupil dilation in response to unexpected omissions of sound and touch Journal Article In: Frontiers in Psychiatry, vol. 14, pp. 1–11, 2023. @article{Dercksen2023, Introduction: Recent theories describe perception as an inferential process based on internal predictive models adjusted by means of prediction violations (prediction error). To study and demonstrate predictive processing in the brain the use of unexpected stimulus omissions has been suggested as a promising approach as the evoked brain responses are uncontaminated by responses to stimuli. Here, we aimed to investigate the pupil's response to unexpected stimulus omissions in order to better understand surprise and orienting of attention resulting from prediction violation. So far only few studies have used omission in pupillometry research and results have been inconsistent. Methods: This study adapted an EEG paradigm that has been shown to elicit omission responses in auditory and somatosensory modalities. Healthy adults pressed a button at their own pace, which resulted in the presentation of sounds or tactile stimuli in either 88%, 50% or 0% (motor-control) of cases. Pupil size was recorded continuously and averaged to analyze the pupil dilation response associated with each condition. Results: Results revealed that omission responses were observed in both modalities in the 88%-condition compared to motor-control. Similar pupil omission responses were observed between modalities, suggesting modality-unspecific activation of the underlying brain circuits. Discussion: In combination with previous omission studies using EEG, the findings demonstrate predictive models in brain processing and point to the involvement of subcortical structures in the omission response. Our pupillometry approach is especially suitable to study sensory prediction in vulnerable populations within the psychiatric field. |
Wolf Culemann; Leana Neuber; Angela Heine Pupil vs. eyelid: Evaluating the accuracy of blink detection in pupil-based eye tracking devices Journal Article In: Procedia Computer Science, vol. 225, pp. 2008–2017, 2023. @article{Culemann2023, Spontaneous eye blinks as a means to tap into human cognitive and affective processes have been demonstrated to offer valuable Spontaneous eye blinks as a means to tap into human cognitive and affective processes have been demonstrated to offer valuable insight in various fields of application, such as human-computer-interaction and driver drowsiness detection. Furthermore, studies insight in various fields of application, such as human-computer-interaction and driver drowsiness detection. Furthermore, studies focusing on gaze tracking and pupillometry have to deal with eye blinks as artifacts, leading to distortions in gaze and pupil data. insight in various fields of application, such as human-computer-interaction and driver drowsiness detection. Furthermore, studies focusing on gaze tracking and pupillometry have to deal with eye blinks as artifacts, leading to distortions in gaze and pupil data. focusing on gaze tracking and pupillometry have to deal with eye blinks as artifacts, leading to distortions in gaze and pupil data. Accurate detection of blinks and calculation of their duration is, consequently, essential for both areas of research and development. In the present study, the accuracy of standard blink detection approaches based on pupil-based eye trackers was investigated with Accurate detection of blinks and calculation of their duration is, consequently, essential for both areas of research and development. Accurate detection of blinks and calculation of their duration is, consequently, essential for both areas of research and development. In the present study, the accuracy of standard blink detection approaches based on pupil-based eye trackers was investigated with In the present study, the accuracy of standard blink detection approaches based on pupil-based eye trackers was investigated with respect to blink durations as well as blink occurrence detection for different eye closure thresholds. We compared the results respect to blink durations as well as blink occurrence detection for different eye closure thresholds. We compared the results respect to blink durations as well as blink occurrence detection for different eye closure thresholds. We compared the results from two different detection algorithms and two eye tracking devices against ground truth data, i.e. blink events that were labeled manually on a frame-by-frame basis from video recordings. The Eye Aspect Ratio was used for the estimation of blink amplitudes from two different detection algorithms and two eye tracking devices against ground truth data, i.e. blink events that were labeled manually on a frame-by-frame basis from video recordings. The Eye Aspect Ratio was used for the estimation of blink amplitudes and eye closure. The results reveal significant differences in blink detection rate and blink duration depending on the method from two different detection algorithms and two eye tracking devices against ground truth data, i.e. blink events that were labeled manually on a frame-by-frame basis from video recordings. The Eye Aspect Ratio was used for the estimation of blink amplitudes and eye closure. The results reveal significant differences in blink detection rate and blink duration depending on the method used. Blink detection by pupil-based eye tracking devices is demonstrated to be systematically influenced by the choice of device, and eye closure. The results reveal significant differences in blink detection rate and blink duration depending on the method used. Blink detection by pupil-based eye tracking devices is demonstrated to be systematically influenced by the choice of device, detection algorithm, and blink definition. Our results highlight the importance of discussing different approaches to blink detection used. Blink detection by pupil-based eye tracking devices is demonstrated to be systematically influenced by the choice of device, when comparing blink metrics across different devices and applications. In addition, we encourage dialogue within the community to transparently communicate blink detection methods and discuss their impact on study outcomes. detection algorithm, and blink definition. Our results highlight the importance of discussing different approaches to blink detection detection algorithm, and blink definition. Our results highlight the importance of discussing different approaches to blink detection when comparing blink metrics across different devices and applications. In addition, we encourage dialogue within the community when comparing blink metrics across different devices and applications. In addition, we encourage dialogue within the community to transparently communicate blink detection methods and discuss their impact on study outcomes. |
M. Eric Cui; Björn Herrmann Eye movements decrease during effortful speech listening Journal Article In: Journal of Neuroscience, vol. 43, no. 32, pp. 5856–5869, 2023. @article{Cui2023, Hearing impairment affects many older adults but is often diagnosed decades after speech comprehension in noisy situations has become effortful. Accurate assessment of listening effort may thus help diagnose hearing impairment earlier. However, pupillometry—the most used approach to assess listening effort—has limitations that hinder its use in practice. The current study explores a novel way to assess listening effort through eye movements. Building on cognitive and neurophysiological work, we examine the hypothesis that eye movements decrease when speech listening becomes challenging. In three experiments with human participants from both sexes, we demonstrate, consistent with this hypothesis, that fixation duration increases and spatial gaze dispersion decreases with increasing speech masking. Eye movements decreased during effortful speech listening for different visual scenes (free viewing, object tracking) and speech materials (simple sentences, naturalistic stories). In contrast, pupillometry was less sensitive to speech masking during story listening, suggesting pupillometric measures may not be as effective for the assessments of listening effort in naturalistic speech-listening paradigms. Our results reveal a critical link between eye movements and cognitive load, suggesting that neural activity in the brain regions that support the regulation of eye movements, such as frontal eye field and superior colliculus, are modulated when listening is effortful. |
Sophie L. Cronin; Ottmar V. Lipp; Welber Marinovic Pupil dilation during encoding, but not type of auditory stimulation, predicts recognition success in face memory Journal Article In: Biological Psychology, vol. 178, pp. 1–10, 2023. @article{Cronin2023, We encounter and process information from multiple sensory modalities in our daily lives, and research suggests that learning can be more efficient when contexts are multisensory. In this study, we were interested in whether face identity recognition memory might be improved in multisensory learning conditions, and to explore associated changes in pupil dilation during encoding and recognition. In two studies participants completed old/new face recognition tasks wherein visual face stimuli were presented in the context of sounds. Faces were learnt alongside no sound, low arousal sounds (Experiment 1), high arousal non-face relevant, or high arousal face relevant (Experiment 2) sounds. We predicted that the presence of sounds during encoding would improve later recognition accuracy, however, the results did not support this with no effect of sound condition on memory. Pupil dilation, however, was found to predict later successful recognition both at encoding and during recognition. While these results do not provide support to the notion that face learning is improved under multisensory conditions relative to unisensory conditions, they do suggest that pupillometry may be a useful tool to further explore face identity learning and recognition. |
Gonçalo Cosme; Patrícia Arriaga; Pedro J. Rosa; Mitul A. Mehta; Diana Prata Temporal profile of intranasal oxytocin in the human autonomic nervous system at rest: An electrocardiography and pupillometry study Journal Article In: Journal of Psychopharmacology, vol. 37, no. 6, pp. 566–576, 2023. @article{Cosme2023, Background: Human social behavior is modulated by oxytocin (OT). Intranasal administration of OT (IN-OT) is a noninvasive route shown to elicit changes in the autonomic nervous system (ANS) activity; however, IN-OT's effect on the temporal profile of ANS activity at rest is yet to be described. Aims: We aimed to describe the temporal profile of IN-OT at six 10-min time windows from 15- to 100-min post-administration in 20 male participants at rest while continuously recording their pupillary in an eyes-open condition and cardiac activity in eyes-open and eyes-closed conditions. Methods: We used a double-blind, placebo-controlled, within-subjects design study where we extracted two proxies of parasympathetic nervous system (PNS) activity: high-frequency heart rate variability (HF-HRV) and pupillary unrest index (PUI); and a proxy of sympathetic nervous system activity: sample entropy of the pupillary unrest. Results: In the eyes-open condition, we found an effect of IN-OT on the proxies of PNS activity: decreased PUI in the three-time windows post-administration spanning 65–100 min, and as an exploratory finding, an increased HF-HRV in the 80–85 min time window. Conclusions: We suggest there is a role of OT in PNS regulation that may be consistent with OT's currently theorized role in the facilitation of alertness and approach behavior. |
Joseph Colantonio; Igor Bascandziev; Maria Theobald; Garvin Brod; Elizabeth Bonawitz In: Entropy, vol. 25, no. 2, pp. 1–24, 2023. @article{Colantonio2023, Bayesian models allow us to investigate children's belief revision alongside physiological states, such as “surprise”. Recent work finds that pupil dilation (or the “pupillary surprise response”) following expectancy violations is predictive of belief revision. How can probabilistic models inform the interpretations of “surprise”? Shannon Information considers the likelihood of an observed event, given prior beliefs, and suggests stronger surprise occurs following unlikely events. In contrast, Kullback–Leibler divergence considers the dissimilarity between prior beliefs and updated beliefs following observations—with greater surprise indicating more change between belief states to accommodate information. To assess these accounts under different learning contexts, we use Bayesian models that compare these computational measures of “surprise” to contexts where children are asked to either predict or evaluate the same evidence during a water displacement task. We find correlations between the computed Kullback–Leibler divergence and the children's pupillometric responses only when the children actively make predictions, and no correlation between Shannon Information and pupillometry. This suggests that when children attend to their beliefs and make predictions, pupillary responses may signal the degree of divergence between a child's current beliefs and the updated, more accommodating beliefs. |
Ivory Y. Chen; Aytaç Karabay; Sebastiaan Mathȏt; Howard Bowman; Elkan G. Akyürek Concealed identity information detection with pupillometry in rapid serial visual presentation Journal Article In: Psychophysiology, vol. 60, no. 1, pp. 1–14, 2023. @article{Chen2023n, The concealed information test (CIT) relies on bodily reactions to stimuli that are hidden in mind. However, people can use countermeasures, such as purposely focusing on irrelevant things, to confound the CIT. A new method designed to prevent countermeasures uses rapid serial visual presentation (RSVP) to present stimuli on the fringe of awareness. Previous studies that used RSVP in combination with electroencephalography (EEG) showed that participants exhibit a clear reaction to their real first name, even when they try to prevent such a reaction (i.e., when their name is concealed information). Because EEG is not easily applicable outside the laboratory, we investigated here whether pupil size, which is easier to measure, can also be used to detect concealed identity information. In our first study, participants adopted a fake name, and searched for this name in an RSVP task, while their pupil sizes were recorded. Apart from this fake name, their real name and a control name also appeared in the task. We found pupil dilation in response to the task-irrelevant real name, as compared to control names. However, while most participants showed this effect qualitatively, it was not statistically significant for most participants individually. In a second study, we preregistered the proof-of-concept methodology and replicated the original findings. Taken together, our results show that the current RSVP task with pupillometry can detect concealed identity information at a group level. Further development of the method is needed to create a valid and reliable concealed identity information detector at the individual level. |
Yuqing Cai; Christoph Strauch; Stefan Van der Stigchel; Marnix Naber Open-DPSM: An open-source toolkit for modeling pupil size changes to dynamic visual inputs Journal Article In: Behavior Research Methods, pp. 1–17, 2023. @article{Cai2023, Pupil size change is a widely adopted, sensitive indicator for sensory and cognitive processes. However, the interpretation of these changes is complicated by the influence of multiple low-level effects, such as brightness or contrast changes, posing challenges to applying pupillometry outside of extremely controlled settings. Building on and extending previous models, we here introduce Open Dynamic Pupil Size Modeling (Open-DPSM), an open-source toolkit to model pupil size changes to dynamically changing visual inputs using a convolution approach. Open-DPSM incorporates three key steps: (1) Modeling pupillary responses to both luminance and contrast changes; (2) Weighing of the distinct contributions of visual events across the visual field on pupil size change; and (3) Incorporating gaze-contingent visual event extraction and modeling. These steps improve the prediction of pupil size changes beyond the here-evaluated benchmarks. Open-DPSM provides Python functions, as well as a graphical user interface (GUI), enabling the extension of its applications to versatile scenarios and adaptations to individualized needs. By obtaining a predicted pupil trace using video and eye-tracking data, users can mitigate the effects of low-level features by subtracting the predicted trace or assess the efficacy of the low-level feature manipulations a priori by comparing estimated traces across conditions. |
Tom Bullock; Mary H. Maclean; Tyler Santander; Alexander P. Boone; Viktoriya Babenko; Neil M. Dundon; Alexander Stuber; Liann Jimmons; Jamie Raymer; Gold N. Okafor; Michael B. Miller; Barry Giesbrecht; Scott T. Grafton; Janice Marshall Habituation of the stress response multiplex to repeated cold pressor exposure Journal Article In: Frontiers in Psychology, vol. 13, pp. 1–23, 2023. @article{Bullock2023, Humans show remarkable habituation to aversive events as reflected by changes of both subjective report and objective measures of stress. Although much experimental human research focuses on the effects of stress, relatively little is known about the cascade of physiological and neural responses that contribute to stress habituation. The cold pressor test (CPT) is a common method for inducing acute stress in human participants in the laboratory; however, there are gaps in our understanding of the global state changes resulting from this stress-induction technique and how these responses change over multiple exposures. Here, we measure the stress response to repeated CPT exposures using an extensive suite of physiologic measures and state-of-the-art analysis techniques. In two separate sessions on different days, participants underwent five 90 s CPT exposures of both feet and five warm water control exposures, while electrocardiography (ECG), impedance cardiography, continuous blood pressure, pupillometry, scalp electroencephalography (EEG), salivary cortisol and self-reported pain assessments were recorded. A diverse array of adaptive responses are reported that vary in their temporal dynamics within each exposure as well as habituation across repeated exposures. During cold-water exposure there was a cascade of changes across several cardiovascular measures (elevated heart rate (HR), cardiac output (CO) and Mean Arterial Pressure (MAP) and reduced left ventricular ejection time (LVET), stroke volume (SV) and high- frequency heart rate variability (HF)). Increased pupil dilation was observed, as was increased power in low-frequency bands (delta and theta) across frontal EEG electrode sites. Several cardiovascular measures also habituated over repeated cold-water exposures (HR, MAP, CO, SV, LVET) as did pupil dilation and alpha frequency activity across the scalp. Anticipation of cold water induced stress effects in the time-period immediately prior to exposure, indexed by increased pupil size and cortical disinhibition in the alpha and beta frequency bands across central scalp sites. These results provide comprehensive insight into the evolution of a diverse array of stress responses to an acute noxious stressor, and how these responses adaptively contribute to stress habituation. |
Andreas Bär; Hannah E. Bär; Max Schneider; Fritz Renner The pupil as a window to the mind's eye: Greater emotionality of episodic imagery than verbal visualisation of rewarding activities Journal Article In: Journal of Experimental Psychopathology, vol. 14, no. 2, pp. 1–11, 2023. @article{Baer2023, Episodic imagery has been shown to amplify emotion more than abstract verbal representations. This may prove useful for clinical interventions aiming to motivate adaptive behaviours. However, most findings rely on self-report measures and verbal control conditions not designed to actively prevent automatic engagement in episodic imagery. We thus investigated the difference in emotionality between Episodic Imagery (EI) and Verbal Visualisation (VV) using pupil dilation as a physiological measure of emotional arousal. A sample of 75 participants listened to audio recordings describing activities in a positive manner. Subjects were randomly assigned to the EI or VV condition. Participants in the EI condition imagined performing the described activity, while participants in the VV condition visualised the words constituting the descriptions. As predicted, EI led to greater pupil dilation than VV, independent of mental effort. Self-reported anticipatory reward assessed throughout the task was also greater for EI than VV, yet no difference was found for arousal, anticipated reward or motivation. Our findings extend previous work demonstrating the property of episodic imagery to amplify emotion to a physiological level using pupillometry. However, we did not find a transfer to motivation, which is in line with previous studies using verbal control conditions for episodic imagery. |
Jordan C. Abramowitz; Matthew J. Goupell; Kristina DeRoy Milvae Cochlear–implant simulated signal degradation exacerbates listening effort in older listeners Journal Article In: Ear & Hearing, pp. 1–10, 2023. @article{Abramowitz2023, Objectives: Individuals with cochlear implants (CIs) often report that listening requires high levels of effort. Listening effort can increase with decreasing spectral resolution, which occurs when listening with a CI, and can also increase with age. What is not clear is whether these factors interact; older CI listeners potentially experience even higher listening effort with greater signal degradation than younger CI listeners. This study used pupillometry as a physiological index of listening effort to examine whether age, spectral resolution, and their interaction affect listening effort in a simulation of CI listening. Design: Fifteen younger normal-hearing listeners (ages 18 to 31 years) and 15 older normal-hearing listeners (ages 65 to 75 years) participated in this experiment; they had normal hearing thresholds from 0.25 to 4 kHz. Participants repeated sentences presented in quiet that were either unprocessed or vocoded, simulating CI listening. Stimuli frequency spectra were limited to below 4 kHz (to control for effects of age-related high-frequency hearing loss), and spectral resolution was decreased by decreasing the number of vocoder channels, with 32-, 16-, and 8-channel conditions. Behavioral speech recognition scores and pupil dilation were recorded during this task. In addition, cognitive measures of working memory and processing speed were obtained to examine if individual differences in these measures predicted changes in pupil dilation. Results: For trials where the sentence was recalled correctly, there was a significant interaction between age and spectral resolution, with significantly greater pupil dilation in the older normal-hearing listeners for the 8- and 32-channel vocoded conditions. Cognitive measures did not predict pupil dilation. Conclusions: There was a significant interaction between age and spectral resolution, such that older listeners appear to exert relatively higher listening effort than younger listeners when the signal is highly degraded, with the largest effects observed in the eight-channel condition. The clinical implication is that older listeners may be at higher risk for increased listening effort with a CI. |
Leonie Nowack; Hermann J. Müller; Markus Conci Changes in attentional breadth scale with the demands of Kanizsa-figure object completion–evidence from pupillometry Journal Article In: Attention, Perception, and Psychophysics, pp. 1–18, 2023. @article{Nowack2023, The present study investigated whether the integration of separate parts into a whole-object representation varies with the amount of available attentional resources. To this end, two experiments were performed, which required observers to maintain central fixation while searching in peripheral vision for a target among various distractor configurations. The target could either be a “grouped” whole-object Kanizsa figure, or an “ungrouped” configuration of identical figural parts, but which do not support object completion processes to the same extent. In the experiments, accuracies and changes in pupil size were assessed, with the latter reflecting a marker of the covert allocation of attention in the periphery. Experiment 1 revealed a performance benefit for grouped (relative to ungrouped) targets, which increased with decreasing distance from fixation. By contrast, search for ungrouped targets was comparably poor in accuracy without revealing any eccentricity-dependent variation. Moreover, measures of pupillary dilation mirrored this eccentricity-dependent advantage in localizing grouped targets. Next, in Experiment 2, an additional attention-demanding foveal task was introduced in order to further reduce the availability of attentional resources for the peripheral detection task. This additional task hampered performance overall, alongside with corresponding pupil size changes. However, there was still a substantial benefit for grouped over ungrouped targets in both the behavioral and the pupillometric data. This shows that perceptual grouping scales with the allocation of attention even when only residual attentional resources are available to trigger the representation of a complete (target) object, thus illustrating that object completion operates in the “near absence” of attention. |
Mihaela Beatrice Neagu; Abigail A. Kressner; Helia Relaño-Iborra; Per Bækgaard; Torsten Dau; Dorothea Wendt Investigating the reliability of pupillometry as a measure of individualized listening effort Journal Article In: Trends in Hearing, vol. 27, pp. 1–20, 2023. @article{Neagu2023, Recordings of the pupillary response have been used in numerous studies to assess listening effort during a speech-in-noise task. Most studies focused on averaged responses across listeners, whereas less is known about pupil dilation as an indicator of the individuals' listening effort. The present study investigated the reliability of several pupil features as potential indicators of individual listening effort and the impact of different normalization procedures on the reliability. The pupil diameters of 31 normal-hearing listeners were recorded during multiple visits while performing a speech-in-noise task. The signal-to-noise ratios (SNRs) of the stimuli ranged from (Formula presented.) 12 dB to (Formula presented.) 4 dB. All listeners were measured twice at separate visits, and 11 were re-tested at a third visit. To examine the reliability of the pupil responses across visits, the intraclass correlation coefficient was applied to the peak and mean pupil dilation and to the temporal features of the pupil response, extracted using growth curve analysis. The reliability of the pupillary response was assessed in relation to SNR and different normalization procedures over multiple visits. The most reliable pupil features were the traditional mean and peak pupil dilation. The highest reliability results were obtained when the data were baseline-corrected and normalized to the individual pupil response range across all visits. Moreover, the present study results showed only a minor impact of the SNR and the number of visits on the reliability of the pupil response. Overall, the results may provide an important basis for developing a standardized test for pupillometry in the clinic. |
Yoichi Mukai; Juhani Järvikivi; Benjamin V. Tucker The role of phonology-to-orthography consistency in predicting the degree of pupil dilation induced in processing reduced and unreduced speech Journal Article In: Applied Psycholinguistics, vol. 44, no. 5, pp. 784–815, 2023. @article{Mukai2023, The relationship between the ways in which words are pronounced and spelled has been shown to affect spoken word processing, and a consistent relationship between pronunciation and spelling has been reported as a possible cause of unreduced pronunciations being easier to process than reduced counterparts although reduced pronunciations occur more frequently. In the present study, we investigate the effect of pronunciation-to-spelling consistency for reduced and unreduced pronunciations in L1 and L2 listeners of a logographic language. More precisely, we compare L1 and L2 Japanese listeners to probe whether they use orthographic information differently when processing reduced and unreduced speech. Using pupillometry, the current study provides evidence that extends the hypothesis about the role of orthography in the processing of reduced speech. Orthographic realization matters in processing for L1 and L2 advanced listeners. More specifically, how consistent the orthographic realization is with its phonological form (phonology-to-orthography consistency) modulates the extent to which reduced pronunciation induces additional processing costs. The results are further discussed in terms of their implications for how listeners process reduced speech and the role of the orthographic form in speech processing. |
Sreenivasan Meyyappan; Abhijit Rajan; George R. Mangun; Mingzhou Ding Top-down control of the left visual field bias in cued visual spatial attention Journal Article In: Cerebral Cortex, vol. 33, no. 9, pp. 5097–5107, 2023. @article{Meyyappan2023, A left visual field (LVF) bias in perceptual judgments, response speed, and discrimination accuracy has been reported in humans. Cognitive factors, such as visual spatial attention, are known to modulate or even eliminate this bias. We investigated this problem by recording pupillometry together with functional magnetic resonance imaging (fMRI) in a cued visual spatial attention task. We observed that (i) the pupil was significantly more dilated following attend-right than attend-left cues, (ii) the task performance (e.g. reaction time [RT]) did not differ between attend-left and attend-right trials, and (iii) the difference in cue-related pupil dilation between attend-left and attend-right trials was inversely related to the corresponding difference in RT. Neuroscientically, correlating the difference in cue-related pupil dilation with the corresponding cue-related fMRI difference yielded activations primarily in the right hemisphere, including the right intraparietal sulcus and the right ventrolateral prefrontal cortex. These results suggest that (i) there is an asymmetry in visual spatial attention control, with the rightward attention control being more effortful than the leftward attention control, (ii) this asymmetry underlies the reduction or the elimination of the LVF bias, and (iii) the components of the attentional control networks in the right hemisphere are likely part of the neural substrate of the observed asymmetry in attentional control. |
2022 |
Cherie Zhou; Monicque M. Lorist; Sebastiaan Mathôt Is categorization in visual working memory a way to reduce mental effort? A pupillometry study Journal Article In: Cognitive Science, vol. 46, no. 9, pp. 1–15, 2022. @article{Zhou2022a, Recent studies on visual working memory (VWM) have shown that visual information can be stored in VWM as continuous (e.g., a specific shade of red) as well as categorical representations (e.g., the general category red). It has been widely assumed, yet never directly tested, that continuous representations require more VWM mental effort than categorical representations; given limited VWM capacity, this would mean that fewer continuous, as compared to categorical, representations can be maintained simultaneously. We tested this assumption by measuring pupil size, as a proxy for mental effort, in a delayed estimation task. Participants memorized one to four ambiguous (boundaries between adjacent color categories) or prototypical colors to encourage continuous or categorical representations, respectively; after a delay, a probe indicated the location of the to-be-reported color. We found that, for memory load 1, pupil size was larger while maintaining ambiguous as compared to prototypical colors, but without any difference in memory precision; this suggests that participants relied on an effortful continuous representation to maintain a single ambiguous color, thus resulting in pupil dilation while preserving precision. Strikingly, this effect gradually inverted, such that for memory load 4, pupil size was smaller while maintaining ambiguous and prototypical colors, but memory precision was now substantially reduced for ambiguous colors; this suggests that with increased memory load participants increasingly relied on categorical representations for ambiguous colors (which are by definition a poor fit to any category). Taken together, our results suggest that continuous representations are more effortful than categorical representations and that very few continuous representations (perhaps only one) can be maintained simultaneously. |
Atsushi Yokoi; Jeffrey Weiler Pupil diameter tracked during motor adaptation in humans Journal Article In: Journal of Neurophysiology, vol. 128, no. 5, pp. 1224–1243, 2022. @article{Yokoi2022, Pupil diameter is known as a noninvasive window into individuals' internal states. Despite the growing use of pupillometry in cognitive learning studies, it receives little attention in motor learning studies. Here, we characterized the pupil responses in a short-term reach adaptation paradigm by measuring pupil diameter of human participants while they adapted to abrupt, gradual, or switching force field conditions. Our results demonstrate how surprise and uncertainty reflected in pupil diameter develop during motor adaptation.Pupil diameter, under constant illumination, is known to reflect individuals' internal states, such as surprise about observation and environmental uncertainty. Despite the growing use of pupillometry in cognitive learning studies as an additional measure for examining internal states, few studies have used pupillometry in human motor learning studies. Here, we provide the first detailed characterization of pupil diameter changes in a short-term reach adaptation paradigm. We measured pupil changes in 121 human participants while they adapted to abrupt, gradual, or switching force field conditions. Sudden increases in movement error caused by the introduction/reversal of the force field resulted in strong phasic pupil dilation during movement accompanied by a transient increase in tonic premovement baseline pupil diameter in subsequent trials. In contrast, pupil responses were reduced when the force field was gradually introduced, indicating that large, unexpected errors drove the changes in pupil responses. Interestingly, however, error-induced pupil responses gradually became insensitive after experiencing multiple force field reversals. We also found an association between baseline pupil diameter and incidental knowledge of the gradually introduced perturbation. Finally, in all experiments, we found a strong co-occurrence of larger baseline pupil diameter with slower reaction and movement times after each rest break. Collectively, these results suggest that tonic baseline pupil diameter reflects one's belief about environmental uncertainty, whereas phasic pupil dilation during movement reflects surprise about a sensory outcome (i.e., movement error), and both effects are modulated by novelty. Our results provide a new approach for nonverbally assessing participants' internal states during motor learning. |
Weizhen Xie; JC Lynne Lu Sing; Ana Martinez-Flores; Weiwei Zhang Induced negative arousal modulates the speed of visual working memory consolidation Journal Article In: Emotion, vol. 22, no. 1, pp. 179–197, 2022. @article{Xie2022a, This study examines how induced negative arousal influences the consolidation of fragile sensory inputs into durable working memory (WM) representations. Participants performed a visual WM change detection task with different amounts of encoding time manipulated by random pattern masks inserted at different levels of memory-and-mask Stimulus Onset Asynchrony (SOA). Prior to the WM task, negative or neutral emotion was induced using audio clips from the International Affective Digital Sounds (IADS). Pupillometry was simultaneously recorded to provide an objective measure of induced arousal. Self-report measures of early-life stress (i.e., adverse childhood experiences) and current mood states (i.e., depressed mood and anxious feeling) were also collected as covariates. We find that participants initially remember a comparable number of WM items under a short memory-and-mask SOA of 100 ms across emotion conditions, but then encode more items into WM at a longer memory-and-mask SOA of 333 ms under induced negative arousal. These findings suggest that induced negative arousal speeds up WM consolidation. Yet, induced negative arousal does not seem to significantly affect participants' WM storage capacity estimated from a separate no mask condition. Furthermore, this emotional effect on WM consolidation speed is moderated by key affect-related individual differences. Participants who have greater pupil responses to negative IADS sounds or have more early-life stress show faster WM consolidation under induced negative arousal. Collectively, our findings reveal a critical role of phasic adrenergic responses in the rapid consolidation of visual WM content and identify potential moderators of this association. |
Thomas Wilschut; Sebastiaan Mathôt Interactions between visual working memory, attention, and color categories: A pupillometry study Journal Article In: Journal of Cognition, vol. 5, no. 1, pp. 1–12, 2022. @article{Wilschut2022, Recent studies have found that visual working memory (VWM) for color shows a categorical bias: Observers typically remember colors as more prototypical to the category they belong to than they actually are. Here, we further examine color-category effects on VWM using pupillometry. Participants remembered a color for later reproduction on a color wheel. During the retention interval, a colored probe was presented, and we measured the pupil constriction in response to this probe, assuming that the strength of constriction reflects the visual saliency of the probe. We found that the pupil initially constricted most strongly for non-matching colors that were maximally different from the memorized color; this likely reflects a lack of visual adaptation for these colors, which renders them more salient than memory-matching colors (which were shown before). Strikingly, this effect reversed later in time, such that pupil constriction was more prolonged for memory-matching colors as compared to non-matching colors; this likely reflects that memory-matching colors capture attention more strongly, and perhaps for a longer time, than non-matching colors do. We found no effects of color categories on pupil constriction: After controlling for color distance, (non-matching) colors from the same category as the memory color did not result in a different pupil response as compared to colors from a different category; however, we did find that behavioral responses were biased by color categories. In summary, we found that pupil constriction to colored probes reflects both visual adaptation and VWM content, but, unlike behavioral measures, is not notably affected by color categories. |
I. K. Wardhani; B. H. Janssen; C. N. Boehler Investigating the relationship between background luminance and self-reported valence of auditory stimuli Journal Article In: Acta Psychologica, vol. 224, pp. 1–10, 2022. @article{Wardhani2022a, The present study investigated the effect of background luminance on the self-reported valence ratings of auditory stimuli, as suggested by some earlier work. A secondary aim was to better characterise the effect of auditory valence on pupillary responses, on which the literature is inconsistent. Participants were randomly presented with sounds of different valence categories (negative, neutral, and positive) obtained from the IADS-E database. At the same time, the background luminance of the computer screen (in blue hue) was manipulated across three levels (i.e., low, medium, and high), with pupillometry confirming the expected strong effect of luminance on pupil size. Participants were asked to rate the valence of the presented sound under these different luminance levels. On a behavioural level, we found evidence for an effect of background luminance on the self-reported valence rating, with generally more positive ratings as background luminance increased. Turning to valence effects on pupil size, irrespective of background luminance, interestingly, we observed that pupils were smallest in the positive valence and the largest in negative valence condition, with neutral valence in between. In sum, the present findings provide evidence concerning a relationship between luminance perception (and hence pupil size) and self-reported valence of auditory stimuli, indicating a possible cross-modal interaction of auditory valence processing with completely task-irrelevant visual background luminance. We furthermore discuss the potential for future applications of the current findings in the clinical field. |
Shamini Warda; Jaana Simola; Devin B. Terhune Pupillometry tracks errors in interval timing Journal Article In: Behavioral Neuroscience, vol. 13, no. 2, pp. 495–502, 2022. @article{Warda2022, Recent primate studies suggest a potential link between pupil size and subjectively elapsed duration. Here, we sought to investigate the relationship between pupil size and perceived duration in human participants performing two temporal bisection tasks in the subsecond and suprasecond interval ranges. In the subsecond task, pupil diameter was greater during stimulus processing when shorter intervals were overestimated but also during and after stimulus offset when longer intervals were underestimated. By contrast, in the suprasecond task, larger pupil diameter was observed only in the late stimulus offset phase prior to response prompts when longer intervals were underestimated. This pattern of results suggests that pupil diameter relates to an error monitoring mechanism in interval timing. These results are at odds with a direct relationship between pupil size and the perception of duration but suggest that pupillometric variation might play a key role in signifying errors related to temporal judgments. |
Stella D. Voulgaropoulou; Fasya Fauzani; Janine Pfirrmann; Claudia Vingerhoets; Thérèse Amelsvoort; Dennis Hernaus Asymmetric effects of acute stress on cost and benefit learning Journal Article In: Psychoneuroendocrinology, vol. 138, pp. 1–10, 2022. @article{Voulgaropoulou2022, Background: Humans are continuously exposed to stressful challenges in everyday life. Such stressful events trigger a complex physiological reaction – the fight-or-flight response – that can hamper flexible decision-making and learning. Inspired by key neural and peripheral characteristics of the fight-or-flight response, here, we ask whether acute stress changes how humans learn about costs and benefits. Methods: Healthy adults were randomly exposed to an acute stress (age mean=23.48, 21/40 female) or no-stress control (age mean=23.80, 22/40 female) condition, after which they completed a reinforcement learning task in which they minimize cost (physical effort) and maximize benefits (monetary rewards). During the task pupillometry data were collected. A computational model of cost-benefit reinforcement learning was employed to investigate the effect of acute stress on cost and benefit learning and decision-making. Results: Acute stress improved learning to maximize rewards relative to minimizing physical effort (Condition-by-Trial Type interaction: F(1,78)= 6.53 |
Jeshua Tromp; Sander Nieuwenhuis; Peter Murphy The effects of neural gain on reactive cognitive control Journal Article In: Computational Brain and Behavior, vol. 5, no. 3, pp. 422–433, 2022. @article{Tromp2022, Fluctuations in global neural gain, arising from brainstem arousal systems, have been found to shape attention, learning, and decision-making as well as cortical state. Comparatively, little is known about how fluctuations in neural gain affect cognitive control. In the present study, we examined this question using a combination of behavioral methods, pupillometry, and computational modeling. Simulations of a comprehensive model of the Stroop task incorporating task conflict and both proactive and reactive forms of control indicated that increasing global gain led to an overall speeding of reaction times, increased Stroop interference, and decreased Stroop facilitation. Pupil analyses revealed that the pre-trial pupil derivative (i.e., rate of change), a putative non-invasive index of global gain, showed the same diagnostic relationships with the Stroop-task performance of human participants. An analysis of the internal model dynamics suggested that a gain-related increase in task conflict and corresponding (within-trial) increase in reactive control are vital for understanding this pattern of behavioral results. Indeed, a similar connectionist model without this task-conflict-control loop could not account for the results. Our study suggests that spontaneous fluctuations in neural gain can have a significant impact on reactive cognitive control. |
Chiara Tortelli; Antonella Pomè; Marco Turi; Roberta Igliozzi; David C. Burr; Paola Binda Contextual information modulates pupil size in autistic children Journal Article In: Frontiers in Neuroscience, vol. 16, pp. 1–10, 2022. @article{Tortelli2022, Recent Bayesian models suggest that perception is more “data-driven” and less dependent on contextual information in autistic individuals than others. However, experimental tests of this hypothesis have given mixed results, possibly due to the lack of objectivity of the self-report methods typically employed. Here we introduce an objective no-report paradigm based on pupillometry to assess the processing of contextual information in autistic children, together with a comparison clinical group. After validating in neurotypical adults a child-friendly pupillometric paradigm, in which we embedded test images within an animation movie that participants watched passively, we compared pupillary response to images of the sun and meaningless control images in children with autism vs. age- and IQ-matched children presenting developmental disorders unrelated to the autistic spectrum. Both clinical groups showed stronger pupillary constriction for the sun images compared with control images, like the neurotypical adults. However, there was no detectable difference between autistic children and the comparison group, despite a significant difference in pupillary light responses, which were enhanced in the autistic group. Our report introduces an objective technique for studying perception in clinical samples and children. The lack of statistically significant group differences in our tests suggests that autistic children and the comparison group do not show large differences in perception of these stimuli. This opens the way to further studies testing contextual processing at other levels of perception. |
Christoph Strauch; Christophe Romein; Marnix Naber; Stefan Van der Stigchel; Antonia F. Ten Brink The orienting response drives pseudoneglect—Evidence from an objective pupillometric method Journal Article In: Cortex, vol. 151, pp. 259–271, 2022. @article{Strauch2022, Spatial attention is generally slightly biased leftward (“pseudoneglect”), a phenomenon typically assessed with paper-and-pencil tasks, limited by the requirement of explicit responses and the inability to assess on a subsecond timescale. Pseudoneglect is often stable within experiments, but differs vastly between investigations and is sometimes directed to the left, sometimes to the right. To date, no exhaustive explanation to this phenomenon has been provided. Here, we objectively assessed lateralized attention over time, exploiting the phenomenon that changes in the pupil reflect the allocation of attention in space. Pupil sizes of 41 healthy participants fixating the center were influenced stronger by the differential background luminance of the left side compared to the right side of the visual display. These differences were mainly driven by visual information in the periphery. Differences in pupil sizes positively related with greyscales scores. Time-based analyses within trials show strongest effects early on. With increasing trial number (not time), the initial leftward bias shifted central in pupillometry-based and greyscales measures. This suggests that the orienting response determines the degree of attention bias. In our amplification hypothesis we pose that the quality of pseudoneglect (i.e., the direction) is determined by higher order factors such as hemispheric imbalances, whereas the quantity (i.e., the degree) is determined by the orienting network. This account might explain numerous—previously thought opposing—findings. We here show how pupil light responses reveal pseudoneglect, in a next step, this might allow clinical diagnosis of hemispatial neglect. |
Adi Shechter; Ronen Hershman; David L. Share A pupillometric study of developmental and individual differences in cognitive effort in visual word recognition Journal Article In: Scientific Reports, vol. 12, no. 1, pp. 1–7, 2022. @article{Shechter2022, Throughout the history of modern psychology, the neural basis of cognitive performance, and particularly its efficiency, has been assumed to be an essential determinant of developmental and individual differences in a wide range of human behaviors. Here, we examine one aspect of cognitive efficiency—cognitive effort, using pupillometry to examine differences in word reading among adults (N = 34) and children (N = 34). The developmental analyses confirmed that children invested more effort in reading than adults, as indicated by larger and sustained pupillary responses. The within-age (individual difference) analyses comparing faster (N = 10) and slower (N = 10) performers revealed that in both age groups, the faster readers demonstrated accelerated pupillary responses compared to slower readers, although both groups invested a similar overall degree of cognitive effort. These findings have the potential to open up new avenues of research in the study of skill growth in word recognition and many other domains of skill learning. |
Nir Shalev; Anna C. Nobre Eyes wide open: Regulation of arousal by temporal expectations Journal Article In: Cognition, vol. 224, pp. 1–12, 2022. @article{Shalev2022, Maintaining adequate levels of arousal is essential for sustaining performance on extended tasks. To investigate arousal in prolonged tasks such as driving studies have traditionally used monotonous task designs. Both ecological and experimental settings often contain embedded temporal regularities, but it is unknown whether these enable adaptive modulation of arousal. We explored whether temporal predictability can modulate arousal according to the timing of anticipated relevant events. In two experiments, we manipulated the temporal predictability of events to test for behavioural benefits and arousal modulation, using pupillometry as a proxy measure. High temporal predictability significantly lowered the tonic level of arousal briefly increased arousal in anticipation of upcoming stimuli, whereas low temporal predictability resulted in tonically elevated arousal. These novel findings suggest that arousal levels flexibly adapt to the temporal structures of events and bring about energy efficiencies in the context of high levels of behavioural performance. |
Elena Selezneva; Nicole Wetzel The impact of probabilistic cues on sound-related pupil dilation and ERP responses in 7–9-year-old children Journal Article In: Auditory Perception & Cognition, vol. 5, no. 1-2, pp. 86–106, 2022. @article{Selezneva2022, Control of involuntary orienting of attention toward new but task-irrelevant events is essential to successfully perform a task. We investigated top-down control of involuntary orienting of attention caused by task-irrelevant novel sounds embedded in a sequence of repeated standard sounds in 7–9-year-old children (N = 30) and in an adult control group (N = 30). The type of sound was announced by visual cues, which were correct in 80% of the trials. We co-registered sound-related pupil dilation responses (PDR), the attention-related component P3a in the EEG and performance. Task-irrelevant novel sounds evoked increased amplitudes of the PDR and the P3a and prolonged reaction times in both age groups. In children only, invalidly cued novel sounds evoked larger PDR amplitudes than validly cued novel sounds, while this cue effect was not observed for standard sounds. In both age groups, P3a amplitudes in the centro-parietal region were reduced to the correctly cued compared to the incorrectly cued novel sounds, indicating top-down control of orienting of attention. The reaction time prolongation to both validly and invalidly cued novel sounds were similar in both age groups. These findings demonstrate that children are capable of reducing the orienting of attention and evaluation triggered by task-irrelevant sounds by using probabilistic cues. Children's pupil results indicate a high sensitivity of pupil dynamics to cue-related top-down influences on novel sound processing, emphasizing the utility of pupillometry in developmental research. |
Matthew K. Robison; Nathaniel T. Diede; Jessica Nicosia; B. Hunter Ball; Julie M. Bugg A multimodal analysis of sustained attention in younger and older adults Journal Article In: Psychology and Aging, vol. 37, no. 3, pp. 307–325, 2022. @article{Robison2022, Age-related cognitive decline has been attributed to processing speed differences, as well as differences in executive control and response inhibition. However, recent research has shown that healthy older adults have intact, if not superior, sustained attention abilities compared to younger adults. The present study used a combination of reaction time (RT), thought probes, and pupillometry to measure sustained attention in samples of younger and older adults. The RT data revealed that, while slightly slower overall, older adults sustained their attention to the task better than younger adults, and did not show a vigilance decrement. Older adults also reported fewer instances of task-unrelated thoughts and reported feeling more motivated and alert than younger adults, despite finding the task more demanding. Additionally, older adults showed larger, albeit laterpeaking, task-evoked pupillary responses (TEPRs), corroborating the behavioral and self-report data. Finally, older adults did not show a shallowing of TEPRs across time, corroborating the finding that their RTs also did not change across time. The present findings are interpreted in light of processing speed theory, resourcedepletion theories of vigilance, and recent neurological theories of cognitive aging. |
Paula Ríos-López; Andreas Widmann; Aurélie Bidet-Caulet; Nicole Wetzel The effect of background speech on attentive sound processing: A pupil dilation study Journal Article In: International Journal of Psychophysiology, vol. 174, pp. 47–56, 2022. @article{RiosLopez2022, Listening to task-irrelevant speech while performing a cognitive task can involuntarily deviate our attention and lead to decreases in performance. One explanation for the impairing effect of irrelevant speech is that semantic processing can consume attentional resources. In the present study, we tested this assumption by measuring performance in a non-linguistic attentional task while participants were exposed to meaningful (native) and non-meaningful (foreign) speech. Moreover, based on the tight relation between pupillometry and attentional processes, we also registered changes in pupil diameter size to quantify the effect of meaningfulness upon attentional allocation. To these aims, we recruited 41 native German speakers who had neither received formal instruction in French nor had extensive informal contact with this language. The focal task consisted of an auditory oddball task. Participants performed a duration discrimination task containing frequently repeated standard sounds and rarely presented deviant sounds while a story was read in German or (non-meaningful) French in the background. Our results revealed that, whereas effects of language meaningfulness on attention were not detectable at the behavioural level, participants' pupil dilated more in response to the sounds of the auditory task when background speech was played in non-meaningful French compared to German, independent of sound type. In line with the initial hypothesis, this suggested that semantic processing of the native language required attentional resources, which lead to fewer resources devoted to the processing of the sounds of the focal task. Our results highlight the potential of the pupil dilation response for the investigation of subtle cognitive processes that might not surface when only behaviour is measured. |
Jasmine Pan; Michaela Klímová; Joseph T. McGuire; Sam Ling Arousal-based pupil modulation is dictated by luminance Journal Article In: Scientific Reports, vol. 12, no. 1, pp. 1–11, 2022. @article{Pan2022, Pupillometry has become a standard measure for assessing arousal state. However, environmental factors such as luminance, a primary dictator of pupillary responses, often vary across studies. To what degree does luminance interact with arousal-driven pupillary changes? Here, we parametrically assessed luminance-driven pupillary responses across a wide-range of luminances, while concurrently manipulating cognitive arousal using auditory math problems of varying difficulty. At the group-level, our results revealed that the modulatory effect of cognitive arousal on pupil size interacts multiplicatively with luminance, with the largest effects occurring at low and mid-luminances. However, at the level of individuals, there were qualitatively distinct individual differences in the modulatory effect of cognitive arousal on luminance-driven pupillary responses. Our findings suggest that pupillometry as a measure for assessing arousal requires more careful consideration: there are ranges of luminance levels that are more ideal in observing pupillary differences between arousal conditions than others. |
Drew J. McLaughlin; Maggie E. Zink; Lauren Gaunt; Brent Spehar; Kristin J. Van Engen; Mitchell S. Sommers; Jonathan E. Peelle Pupillometry reveals cognitive demands of lexical competition during spoken word recognition in young and older adults Journal Article In: Psychonomic Bulletin & Review, vol. 29, no. 1, pp. 1–13, 2022. @article{McLaughlin2022, In most contemporary activation-competition frameworks for spoken word recognition, candidate words compete against phonological “neighbors” with similar acoustic properties (e.g., “cap” vs. “cat”). Thus, recognizing words with more competitors should come at a greater cognitive cost relative to recognizing words with fewer competitors, due to increased demands for selecting the correct item and inhibiting incorrect candidates. Importantly, these processes should operate even in the absence of differences in accuracy. In the present study, we tested this proposal by examining differences in processing costs associated with neighborhood density for highly intelligible items presented in quiet. A second goal was to examine whether the cognitive demands associated with increased neighborhood density were greater for older adults compared with young adults. Using pupillometry as an index of cognitive processing load, we compared the cognitive demands associated with spoken word recognition for words with many or fewer neighbors, presented in quiet, for young (n = 67) and older (n = 69) adult listeners. Growth curve analysis of the pupil data indicated that older adults showed a greater evoked pupil response for spoken words than did young adults, consistent with increased cognitive load during spoken word recognition. Words from dense neighborhoods were marginally more demanding to process than words from sparse neighborhoods. There was also an interaction between age and neighborhood density, indicating larger effects of density in young adult listeners. These results highlight the importance of assessing both cognitive demands and accuracy when investigating the mechanisms underlying spoken word recognition. |
Joel T. Martin; Annalise H. Whittaker; Stephen J. Johnston Pupillometry and the vigilance decrement: Task-evoked but not baseline pupil measures reflect declining performance in visual vigilance tasks Journal Article In: European Journal of Neuroscience, vol. 55, no. 3, pp. 778–799, 2022. @article{Martin2022, Baseline and task-evoked pupil measures are known to reflect the activity of the nervous system's central arousal mechanisms. With the increasing availability, affordability and flexibility of video-based eye tracking hardware, these measures may one day find practical application in real-time biobehavioural monitoring systems to assess performance or fitness for duty in tasks requiring vigilant attention. But real-world vigilance tasks are predominantly visual in their nature and most research in this area has taken place in the auditory domain. Here, we explore the relationship between pupil size—both baseline and task-evoked—and behavioural performance measures in two novel vigilance tasks requiring visual target detection: (1) a traditional vigilance task involving prolonged, continuous and uninterrupted performance (n = 28) and (2) a psychomotor vigilance task (n = 25). In both tasks, behavioural performance and task-evoked pupil responses declined as time spent on task increased, corroborating previous reports in the literature of a vigilance decrement with a corresponding reduction in task-evoked pupil measures. Also in line with previous findings, baseline pupil size did not show a consistent relationship with performance measures. Our data offer novel insights into the complex interplay of brain systems involved in vigilant attention and question the validity of the assumption that baseline (prestimulus) pupil size and task-evoked (poststimulus) pupil measures reflect the tonic and phasic firing modes of the locus coeruleus. |
Kangjoo Lee; Corey Horien; David O'Connor; Bronwen Garand-Sheridan; Fuyuze Tokoglu; Dustin Scheinost; Evelyn M. R. Lake; R. Todd Constable Arousal impacts distributed hubs modulating the integration of brain functional connectivity Journal Article In: NeuroImage, vol. 258, pp. 1–17, 2022. @article{Lee2022c, Even when subjects are at rest, it is thought that brain activity is organized into distinct brain states during which reproducible patterns are observable. Yet, it is unclear how to define or distinguish different brain states. A potential source of brain state variation is arousal, which may play a role in modulating functional interactions between brain regions. Here, we use simultaneous resting state functional magnetic resonance imaging (fMRI) and pupillometry to study the impact of arousal levels indexed by pupil area on the integration of large-scale brain networks. We employ a novel sparse dictionary learning-based method to identify hub regions participating in between-network integration stratified by arousal, by measuring k-hubness, the number (k) of functionally overlapping networks in each brain region. We show evidence of a brain-wide decrease in between-network integration and inter-subject variability at low relative to high arousal, with differences emerging across regions of the frontoparietal, default mode, motor, limbic, and cerebellum networks. State-dependent changes in k-hubness relate to the actual patterns of network integration within these hubs, suggesting a brain state transition from high to low arousal characterized by global synchronization and reduced network overlaps. We demonstrate that arousal is not limited to specific brain areas known to be directly associated with arousal regulation, but instead has a brain-wide impact that involves high-level between-network communications. Lastly, we show a systematic change in pairwise fMRI signal correlation structures in the arousal state-stratified data, and demonstrate that the choice of global signal regression could result in different conclusions in conventional graph theoretical analysis and in the analysis of k-hubness when studying arousal modulations. Together, our results suggest the presence of global and local effects of pupil-linked arousal modulations on resting state brain functional connectivity. |
Paul A. Keene; Megan T. DeBettencourt; Edward Awh; Edward K. Vogel Pupillometry signatures of sustained attention and working memory Journal Article In: Attention, Perception, and Psychophysics, vol. 84, pp. 2472–2482, 2022. @article{Keene2022, There exists an intricate relationship between attention and working memory. Recent work has further established that attention and working memory fluctuate synchronously, by tightly interleaving sustained attention and working memory tasks. This work has raised many open questions about physiological signatures underlying these behavioral fluctuations. Across two experiments, we explore pupil dynamics using real-time triggering in conjunction with an interleaved sustained attention and working memory task. In Experiment 1, we use behavioral real-time triggering and replicate recent findings from our lab (deBettencourt et al., 2019) that sustained attention fluctuates concurrently with the number of items maintained in working memory. Furthermore, highly attentive moments, detected via behavior, also exhibited larger pupil sizes. In Experiment 2, we develop a novel real-time pupil-triggering technique to track pupil size fluctuations in real time and trigger working memory probes. We show that this pupil triggering procedure reveals differences in sustained attention, as indexed by response time. These experiments reflect methodological advances in real-time triggering and further disentangle the relationship among general arousal, sustained attention, and working memory. |
Michael A. Johns; Paola E. Dussias Comparing single-word insertions and multi-word alternations in bilingual speech: Insights from pupillometry Journal Article In: Languages, vol. 7, pp. 1–19, 2022. @article{Johns2022, Prominent sociolinguistic theories of language mixing have posited that single-word insertions of one language into the other are the result of a distinct process than multi-word alternations between two languages given that the former overwhelmingly surface morphosyntactically integrated into the surrounding language. To date, this distinction has not been tested in comprehension. The present study makes use of pupillometry to examine the online processing of single-word insertions and multi-word alternations by highly proficient Spanish-English bilinguals in Puerto Rico. Participants heard sentences containing target noun/adjective pairs (1) in unilingual Spanish, (2) where the Spanish noun was replaced with its English translation equivalent, followed by a Spanish post-nominal adjective, and (3) where both the noun and adjective appeared in English with the adjective occurring in the English pre-nominal position. Both types of language mixing elicit larger pupillary responses when compared to unilingual Spanish speech, though the magnitude of this difference depends on the grammatical gender of the target noun. Importantly, single-word insertions and multi-word alternations did not differ from one another. Taken together, these findings suggest that morphosyntactic integration is not the defining feature of single-word insertions, at least in comprehension, and that the comprehension system is tuned to the distributional properties of bilingual speech. |
Novera Istiqomah; Yuta Suzuki; Yuya Kinzuka; Tetsuto Minami; Shigeki Nakauchi Anisotropy in the peripheral visual field based on pupil response to the glare illusion Journal Article In: Heliyon, vol. 8, no. 6, pp. 1–7, 2022. @article{Istiqomah2022, Visual-field (VF) anisotropy has been investigated in terms of spatial resolution of attention, spatial frequency, and semantic processing. Brightness perception has also been reported to vary between VFs. However, the influence of VF anisotropy on brightness perception using pupillometry has not been investigated. The present study measured participants' pupil size during glare illusion, in which converging luminance gradients evoke brightness enhancement and a glowing impression on the central white area of the stimulus, and halo stimuli, in which the same physical brightness of the glare illusion is used with a diverging luminance pattern. The results revealed greater stimulus-evoked pupillary dilation and glare-related dilated pupil reduction in the upper VF (UVF) compared with other VFs and halo-related pupillary changes, respectively. The stimulus-evoked pupillary dilation was affected by poor contrast sensitivity. However, owing to the superior cognitive bias formed by statistical regularity in natural scene processing of the glare illusion in the UVF, we found reduced pupillary dilation compared with the response to halo stimuli and the response from other VFs. These findings offer valuable insight into a method to reduce the potential glare effect of any VF anisotropy induced by the glare effect experienced in daily vision. An important practical implication of our study may be in informing the design of applications aimed at improving nighttime driving behavior. We also believe that our study makes a significant contribution to the literature because it offers valuable insights on VF anisotropy using evidence from pupillometry and the glare illusion. |
Josephine M. Groot; Gábor Csifcsák; Sven Wientjes; Birte U. Forstmann; Matthias Mittner Catching wandering minds with tapping fingers: Neural and behavioral insights into task-unrelated cognition Journal Article In: Cerebral Cortex, vol. 32, pp. 4447–4463, 2022. @article{Groot2022, When the human mind wanders, it engages in episodes during which attention is focused on self-generated thoughts rather than on external task demands. Although the sustained attention to response task is commonly used to examine relationships between mind wandering and executive functions, limited executive resources are required for optimal task performance. In the current study, we aimed to investigate the relationship between mind wandering and executive functions more closely by employing a recently developed finger-tapping task to monitor fluctuations in attention and executive control through task performance and periodical experience sampling during concurrent functional magnetic resonance imaging (fMRI) and pupillometry. Our results show that mind wandering was preceded by increases in finger-tapping variability, which was correlated with activity in dorsal and ventral attention networks. The entropy of random finger-tapping sequences was related to activity in frontoparietal regions associated with executive control, demonstrating the suitability of this paradigm for studying executive functioning. The neural correlates of behavioral performance, pupillary dynamics, and self-reported attentional state diverged, thus indicating a dissociation between direct and indirect markers of mind wandering. Together, the investigation of these relationships at both the behavioral and neural level provided novel insights into the identification of underlying mechanisms of mind wandering. |
Léon Franzen; Amanda Cabugao; Bianca Grohmann; Karine Elalouf; Aaron P. Johnson Individual pupil size changes as a robust indicator of cognitive familiarity differences Journal Article In: PLoS ONE, vol. 17, no. 1, pp. 1–22, 2022. @article{Franzen2022, Cognitive psychology has a long history of using physiological measures, such as pupillometry. However, their susceptibility to confounds introduced by stimulus properties, such as color and luminance, has limited their application. Pupil size measurements, in particular, require sophisticated experimental designs to dissociate relatively small changes in pupil diameter due to cognitive responses from larger ones elicited by changes in stimulus properties or the experimental environment. Here, building on previous research, we present a pupillometry paradigm that adapts the pupil to stimulus properties during the baseline period without revealing stimulus meaning or context by using a pixel-scrambled image mask around an intact image. We demonstrate its robustness in the context of pupillary responses to branded product familiarity. Results show larger average and peak pupil dilation for passively viewed familiar product images and an extended later temporal component representing differences in familiarity across participants (starting around 1400 ms post-stimulus onset). These amplitude differences are present for almost all participants at the single-participant level, and vary somewhat by product category. However, amplitude differences were absent during the baseline period. These findings demonstrate that involuntary pupil size measurements combined with the presented paradigm are successful in dissociating cognitive effects of familiarity from physical stimulus confounds. |
Julia Fietz; Dorothee Pöhlchen; Florian P. Binder; Michael Czisch; Philipp G. Sämann; Victor I. Spoormaker Pupillometry tracks cognitive load and salience network activity in a working memory functional magnetic resonance imaging task Journal Article In: Human Brain Mapping, vol. 43, no. 2, pp. 665–680, 2022. @article{Fietz2022, The diameter of the human pupil tracks working memory processing and is associated with activity in the frontoparietal network. At the same time, recent neuroimaging research has linked human pupil fluctuations to activity in the salience network. In this combined functional magnetic resonance imaging (fMRI)/pupillometry study, we recorded the pupil size of healthy human participants while they performed a blockwise organized working memory task (N-back) inside an MRI scanner in order to monitor the pupil fluctuations associated neural activity during working memory processing. We first confirmed that mean pupil size closely followed working memory load. Combining this with fMRI data, we focused on blood oxygen level dependent (BOLD) correlates of mean pupil size modeled onto the task blocks as a parametric modulation. Interrogating this modulated task regressor, we were able to retrieve the frontoparietal network. Next, to fully exploit the within-block dynamics, we divided the blocks into 1 s time bins and filled these with corresponding pupil change values (first-order derivative of pupil size). We found that pupil change within N-back blocks was positively correlated with BOLD amplitudes in the areas of the salience network (namely bilateral insula, and anterior cingulate cortex). Taken together, fMRI with simultaneous measurement of pupil parameters constitutes a valuable tool to dissect working memory subprocesses related to both working memory load and salience of the presented stimuli. |
Prateek Dhamija; Allison Wong; Asaf Gilboa Early auditory event related potentials distinguish higher-order from first-order aversive conditioning Journal Article In: Frontiers in Behavioral Neuroscience, vol. 16, pp. 1–14, 2022. @article{Dhamija2022, Stimuli in reality rarely co-occur with primary reward or punishment to allow direct associative learning of value. Instead, value is thought to be inferred through complex higher-order associations. Rodent research has demonstrated that the formation and maintenance of first-order and higher-order associations are supported by distinct neural substrates. In this study, we explored whether this pattern of findings held true for humans. Participants underwent first-order and subsequent higher-order conditioning using an aversive burst of white noise or neutral tone as the unconditioned stimuli. Four distinct tones, initially neutral, served as first-order and higher-order conditioned stimuli. Autonomic and neural responses were indexed by pupillometry and evoked response potentials (ERPs) respectively. Conditioned aversive values of first-order and higher-order stimuli led to increased autonomic responses, as indexed by pupil dilation. Distinct temporo-spatial auditory evoked response potentials were elicited by first-order and high-order conditioned stimuli. Conditioned first-order responses peaked around 260 ms and source estimation suggested a primary medial prefrontal and amygdala source. Conversely, conditioned higher-order responses peaked around 120 ms with an estimated source in the medial temporal lobe. Interestingly, pupillometry responses to first-order conditioned stimuli were diminished after higher order training, possibly signifying concomitant incidental extinction, while responses to higher-order stimuli remained. This suggests that once formed, higher order associations are at least partially independent of first order conditioned representations. This experiment demonstrates that first-order and higher-order conditioned associations have distinct neural signatures, and like rodents, the medial temporal lobe may be specifically involved with higher-order conditioning. |
Annabell Coors; Monique M. B. Breteler; Ulrich Ettinger Processing speed, but not working memory or global cognition, is associated with pupil diameter during fixation Journal Article In: Psychophysiology, vol. 59, pp. 1–13, 2022. @article{Coors2022, Mean pupil size during fixation has been suggested to reflect interindividual differences in working memory and fluid intelligence. However, due to small samples with limited age range (17–35 years) and suboptimal light conditions in previous studies, these associations are still controversial and it is unclear whether they are observed at older ages. Therefore, we assessed whether interindividual differences in cognitive performance are reflected in pupil diameter during fixation and whether these associations are age-dependent. We analyzed pupillometry and cognition data of 4560 individuals aged 30–95 years of the community-based Rhineland Study. Pupillometry data were extracted from a one-minute fixation task. The cognitive test battery included tests of oculomotor control, working memory, episodic verbal memory, processing speed, executive function, and crystallized intelligence. For data analysis, we used multivariable regression models. Working memory and global cognition were not associated with pupil diameter during fixation. Better processing speed performance was associated with larger pupil diameter during fixation. Associations between cognition and pupil diameter during fixation hardly varied with age, but pupil diameter during fixation declined linearly with age (adjusted decline: 0.33 mm per 10 years of age). There were no significant sex differences in pupil size. We conclude that interindividual differences in mean pupil diameter during fixation may partly reflect interindividual differences in the speed of processing and response generation. We could not confirm that interindividual differences in working memory and fluid intelligence are reflected in pupil size during fixation; however, our sample differed in age range from previous studies. |
Emily A. Burg; Tanvi D. Thakkar; Ruth Y. Litovsky Interaural speech asymmetry predicts bilateral speech intelligibility but not listening effort in adults with bilateral cochlear implants Journal Article In: Frontiers in Neuroscience, vol. 16, pp. 1–13, 2022. @article{Burg2022, Introduction: Bilateral cochlear implants (BiCIs) can facilitate improved speech intelligibility in noise and sound localization abilities compared to a unilateral implant in individuals with bilateral severe to profound hearing loss. Still, many individuals with BiCIs do not benefit from binaural hearing to the same extent that normal hearing (NH) listeners do. For example, binaural redundancy, a speech intelligibility benefit derived from having access to duplicate copies of a signal, is highly variable among BiCI users. Additionally, patients with hearing loss commonly report elevated listening effort compared to NH listeners. There is some evidence to suggest that BiCIs may reduce listening effort compared to a unilateral CI, but the limited existing literature has not shown this consistently. Critically, no studies to date have investigated this question using pupillometry to quantify listening effort, where large pupil sizes indicate high effort and small pupil sizes indicate low effort. Thus, the present study aimed to build on existing literature by investigating the potential benefits of BiCIs for both speech intelligibility and listening effort. Methods: Twelve BiCI adults were tested in three listening conditions: Better Ear, Poorer Ear, and Bilateral. Stimuli were IEEE sentences presented from a loudspeaker at 0° azimuth in quiet. Participants were asked to repeat back the sentences, and responses were scored by an experimenter while changes in pupil dilation were measured. Results: On average, participants demonstrated similar speech intelligibility in the Better Ear and Bilateral conditions, and significantly worse speech intelligibility in the Poorer Ear condition. Despite similar speech intelligibility in the Better Ear and Bilateral conditions, pupil dilation was significantly larger in the Bilateral condition. Discussion: These results suggest that the BiCI users tested in this study did not demonstrate binaural redundancy in quiet. The large interaural speech asymmetries demonstrated by participants may have precluded them from obtaining binaural redundancy, as shown by the inverse relationship between the two variables. Further, participants did not obtain a release from effort when listening with two ears versus their better ear only. Instead, results indicate that bilateral listening elicited increased effort compared to better ear listening, which may be due to poor integration of asymmetric inputs. |
Doug J. K. Barrett; David Souto; Michael Pilling; David M. Baguley An exploratory investigation of pupillometry as a measure of tinnitus intrusiveness on a test of auditory short-term memory Journal Article In: Ear & Hearing, vol. 43, no. 5, pp. 1540–1548, 2022. @article{Barrett2022, Objectives: The purpose of the current study was to investigate the potential of pupillometry to provide an objective measure of competition between tinnitus and external sounds during a test of auditory short-term memory. Design: Twelve participants with chronic tinnitus and twelve control participants without tinnitus took part in the study. Pretest sessions used an adaptive method to estimate listeners' frequency discrimination threshold on a test of delayed pitch discrimination for pure tones. Target and probe tones were presented at 72 dB SPL and centered on 750 Hz±2 semitones with an additional jitter of 5 to 20 Hz. Test sessions recorded baseline pupil diameter and task-related pupillary responses (TEPRs) during three blocks of delayed pitch discrimination trials. The difference between target and probe tones was set to the individual's frequency detection threshold for 80% response-accuracy. Listeners with tinnitus also completed the Tinnitus Handicap Inventory (THI). Linear mixed effects procedures were applied to examine changes in baseline pupil diameter and TEPRs associated with group (tinnitus versus control), block (1 to 3) and their interaction. The association between THI scores and maximum TEPRs was assessed using simple linear regression. Results: Patterns of baseline pupil dilation across trials diverged in listeners with tinnitus and controls. For controls, baseline pupil dilation remained constant across blocks. For listeners with tinnitus, baseline pupil dilation increased on blocks 2 and 3 compared with block 1. TEPR amplitudes were also larger in listeners with tinnitus than controls. Linear mixed effects models yielded a significant group by block interaction for baseline pupil diameter and a significant main effect of group on maximum TEPR amplitudes. Regression analyses yielded a significant association between THI scores and TEPR amplitude in listeners with tinnitus. Conclusions: Our data indicate measures of baseline pupil diameter, and TEPRs are sensitive to competition between tinnitus and external sounds during a test of auditory short-term memory. This result suggests pupillometry can provide an objective measure of intrusion in tinnitus. Future research will be required to establish whether our findings generalize to listeners across a full range of tinnitus severity. |
Mattis Appelqvist-Dalton; James P. Wilmott; Mingjian He; Andrea Megela Simmons Time perception in film is modulated by sensory modality and arousal Journal Article In: Attention, Perception, and Psychophysics, vol. 84, no. 3, pp. 926–942, 2022. @article{AppelqvistDalton2022, Considerable research has shown that the perception of time can be distorted subjectively, but little empirical work has examined what factors affect time perception in film, a naturalistic multimodal stimulus. Here, we explore the effect of sensory modality, arousal, and valence on how participants estimate durations in film. Using behavioral ratings combined with pupillometry in a within-participants design, we analyzed responses to and duration estimates of film clips in three experimental conditions: audiovisual (containing music and sound effects), visual (without music and sound effects), and auditory (music and sound effects without a visual scene). Participants viewed clips from little-known nature documentaries, fiction, animation, and experimental films. They were asked to judge clip duration and to report subjective arousal and valence, as their pupil sizes were recorded. Data were analyzed using linear mixed-effects models. Results reveal duration estimates varied between experimental conditions. Clip durations were judged to be shorter than actual durations in all three conditions, with visual-only clips perceived as longer (i.e., less distorted in time) than auditory-only and audiovisual clips. High levels of Composite Arousal (an average of self-reported arousal and pupil size changes) were correlated with longer (more accurate) estimates of duration, particularly in the audiovisual modality. This effect may reflect stimulus complexity or greater cognitive engagement. Increased ratings of valence were correlated with longer estimates of duration. The use of naturalistic, complex stimuli such as film can enhance our understanding of the psychology of time perception. |
Andreas Alexandersen; Gábor Csifcsák; Josephine Groot; Matthias Mittner In: Neuroimage: Reports, vol. 2, no. 3, pp. 1–19, 2022. @article{Alexandersen2022, Mind wandering (MW) is a mental phenomenon humans experience daily. Yet, we lack a complete understanding of the neural basis of this pervasive mental state. Over the past decade there has been an increase in publications using transcranial direct current stimulation (tDCS) to modulate the propensity to mind wander, but findings are diverse, and a satisfactory conclusion is missing. Recently, Boayue et al. (2020) reported successful reduction of mind wandering using high-definition tDCS (HD-tDCS) over the dorsolateral prefrontal cortex, providing preliminary evidence for the efficacy of HD-tDCS in interfering with mind wandering. The current study is a high-powered, pre-registered direct replication attempt of the effect found by Boayue et al. (2020). In addition, we investigated whether the effects of HD-tDCS on mind wandering would be prolonged and assessed the underlying processes of mind wandering using electroencephalography (EEG) and pupillometry during a finger-tapping random sequence generation task that requires the use of executive resources. We failed to find any evidence of the original effect of reduced MW during and after stimulation. When combining our data with the data from Boayue et al. (2020), the original effect of reduced MW caused by HD-tDCS disappeared. In addition, we observed increased occipital alpha power as task duration increased and increased midfrontal theta power preceding response patterns signaling high executive function use. Finally, tonic and phasic pupil size decreased as task duration increased yet, phasic responses were increased, while tonic responses were reduced preceding reports of MW. Additionally phasic pupil size also showed a tendency to be increased during periods of high executive function use. Importantly, none of the EEG or pupil measures were modulated by HD-tDCS. We conclude that HD-tDCS over the dorsolateral prefrontal cortex does not affect MW propensity and its neural signatures. Furthermore, we recommend that previously reported effects of tDCS on mind wandering and other cognitive functions should only be accepted after a successful pre-registered replication. |
Miriam Acquafredda; Paola Binda; Claudia Lunghi Attention cueing in rivalry: Insights from pupillometry Journal Article In: eNeuro, vol. 9, no. 3, pp. 1–12, 2022. @article{Acquafredda2022, We used pupillometry to evaluate the effects of attention cueing on perceptual bi-stability, as reported by adult human observers. Perceptual alternations and pupil diameter were measured during two forms of rivalry, generated by presenting a white and a black disk to the two eyes (binocular rivalry) or splitting the disks between eyes (interocular grouping rivalry). In line with previous studies, we found that subtle pupil size modula- tions (;0.05 mm) tracked alternations between exclusive dominance phases of the black or white disk. These pupil responses were larger for perceptually stronger stimuli: presented to the dominant eye or with physically higher luminance contrast. However, cueing of endogenous attention to one of the rivaling percepts did not affect pupil modulations during exclusive dominance phases. This was observed despite the reliable effects of endogenous attention on perceptual dominance, which shifted in favor of the cued percept by ;10%. The results were comparable for binocular and interocular grouping rivalry. Cueing only had a marginal modulatory effect on pupil size during mixed percepts in binocular rivalry. This may suggest that, rather than acting by modulating perceptual strength, endogenous attention primarily acts during periods of unresolved competition, which is compatible with attention being automatically directed to the rivaling stimuli during periods of exclusive dominance and thereby sustaining perceptual alternations. |
Keiyu Niikuni; Ming Wang; Michiru Makuuchi; Masatoshi Koizumi; Sachiko Kiyama Pupil dilation reflects emotional arousal via poetic language Journal Article In: Perceptual and Motor Skills, pp. 1–18, 2022. @article{Niikuni2022, We investigated pupillary responses to the world's shortest fixed verses, Japanese haiku as aesthetic poetry (AP) and senryu as comic poetry (CP), in comparison with non-poetry control stimuli (NP) comprised of slogans that had the same rhythm patterns. Native Japanese speakers without literary training listened to these stimuli while we recorded their pupil diameters. We found that participants' pupils were significantly dilated for CP compared to NP in an early time window. While AP also evoked larger dilations than NP, the latency for AP-related pupil dilation was relatively long. Thus, lay people experience quick and intense arousal in response to funny and humorous words, while aesthetic properties of words may also elicit intense but slower changes in listeners' arousal levels, presumably because they evoke more implicit and subtle emotional effects. This study is the first to provide evidence that poetic language elicits human pupillary dilation. A better understanding of the cognitive and neural substrates for the sensitive awareness of pleasures expressed via poetic language will provide insights for improving mental and physical health. Hence, pupillometry can act as a useful convenient measurement to delineate the sympathetic activation of emotional contexts via language. |
Stuart B. Murray; Tomislav D. Zbozinek; Michelle Craske; Reza Tadayonnejad; Michael Strober; Ausaf A. Bari; John P. O'Doherty; Jamie D. Feusner Neural, physiological, and psychological markers of appetitive conditioning in anorexia nervosa: a study protocol Journal Article In: Journal of Eating Disorders, vol. 10, no. 1, pp. 1–11, 2022. @article{Murray2022b, Background: Anorexia nervosa (AN) is a chronic and disabling psychiatric condition characterized by low hedonic drive towards food, and is thought to be inclusive of altered dimensions of reward processing. Whether there exists a fundamental aberrancy in the capacity to acquire and maintain de novo hedonic associations—a critical component of hedonic responding—has never been studied in AN. Methods: This multi-modal study will employ a 2-day Pavlovian appetitive conditioning paradigm to interrogate the (1) acquisition, (2) extinction, (3) spontaneous recovery and (4) reinstatement of appetitive learning in adolescents and young adults with AN. Participants will be 30 currently ill, underweight individuals with AN; 30 weight-restored individuals with AN; and 30 age-matched healthy controls, all aged 12–22 years. All subjects will undergo clinical assessment, followed by the 2-day appetitive conditioning task during which fMRI, pupillometry, heart rate deceleration, and subjective ratings will be acquired. Discussion: This study will be the first to interrogate appetitive conditioning in AN—a disorder characterized by altered hedonic responding to food. Results will help establish objective biomarkers of appetitive conditioning in AN and lay the groundwork for developing novel lines of treatment for AN and other psychiatric disorders involving diminished ability to experience pleasure and reward. Trial registration: Pending. Intended registry: Clinicaltrials.gov. |
Laura Müller-Pinzler; Nora Czekalla; Annalina V. Mayer; Alexander Schröder; David S. Stolz; Frieder M. Paulus; Sören Krach Neurocomputational mechanisms of affected beliefs Journal Article In: Communications Biology, vol. 5, no. 1, pp. 1–16, 2022. @article{MuellerPinzler2022, The feedback people receive on their behavior shapes the process of belief formation and self-efficacy in mastering a particular task. However, the neural and computational mechanisms of how the subjective value of self-efficacy beliefs, and the corresponding affect, influence the learning process remain unclear. We investigated these mechanisms during self-efficacy belief formation using fMRI, pupillometry, and computational modeling, and by analyzing individual differences in affective experience. Biases in the formation of self-efficacy beliefs were associated with affect, pupil dilation, and neural activity within the anterior insula, amygdala, ventral tegmental area/ substantia nigra, and mPFC. Specifically, neural and pupil responses mapped the valence of the prediction errors in correspondence with individuals' experienced affective states and learning biases during self-efficacy belief formation. Together with the functional connectivity dynamics of the anterior insula within this network, our results provide evidence for neural and computational mechanisms of how we arrive at affected beliefs. |
2021 |
Kyung Yoo; Jeongyeol Ahn; Sang-Hun Lee The confounding effects of eye blinking on pupillometry, and their remedy Journal Article In: PLoS ONE, vol. 16, no. 12, pp. 1–32, 2021. @article{Yoo2021, Pupillometry, thanks to its strong relationship with cognitive factors and recent advancements in measuring techniques, has become popular among cognitive or neural scientists as a tool for studying the physiological processes involved in mental or neural processes. Despite this growing popularity of pupillometry, the methodological understanding of pupillometry is limited, especially regarding potential factors that may threaten pupillary measurements' validity. Eye blinking can be a factor because it frequently occurs in a manner dependent on many cognitive components and induces a pulse-like pupillary change consisting of constriction and dilation with substantive magnitude and length. We set out to characterize the basic properties of this “blink-locked pupillary response (BPR),” including the shape and magnitude of BPR and their variability across subjects and blinks, as the first step of studying the confounding nature of eye blinking. Then, we demonstrated how the dependency of eye blinking on cognitive factors could confound, via BPR, the pupillary responses that are supposed to reflect the cognitive states of interest. By building a statistical model of how the confounding effects of eye blinking occur, we proposed a probabilistic-inference algorithm of de-confounding raw pupillary measurements and showed that the proposed algorithm selectively removed BPR and enhanced the statistical power of pupillometry experiments. Our findings call for attention to the presence and confounding nature of BPR in pupillometry. The algorithm we developed here can be used as an effective remedy for the confounding effects of BPR on pupillometry. |
Peter S. Whitehead; Younis Mahmoud; Paul Seli; Tobias Egner Mind wandering at encoding, but not at retrieval, disrupts one-shot stimulus-control learning Journal Article In: Attention, Perception, and Psychophysics, vol. 83, no. 7, pp. 2968–2982, 2021. @article{Whitehead2021, The one-shot pairing of a stimulus with a specific cognitive control process, such as task switching, can bind the two together in memory. The episodic control-binding hypothesis posits that the formation of temporary stimulus-control bindings, which are held in event-files supported by episodic memory, can guide the contextually appropriate application of cognitive control. Across two experiments, we sought to examine the role of task-focused attention in the encoding and implementation of stimulus-control bindings in episodic event-files. In Experiment 1, we obtained self-reports of mind wandering during encoding and implementation of stimulus-control bindings. Results indicated that, whereas mind wandering during the implementation of stimulus-control bindings does not decrease their efficacy, mind wandering during the encoding of these control-state associations interferes with their successful deployment at a later point. In Experiment 2, we complemented these results by using trial-by-trial pupillometry to measure attention, again demonstrating that attention levels at encoding predict the subsequent implementation of stimulus-control bindings better than attention levels at implementation. These results suggest that, although encoding stimulus-control bindings in episodic memory requires active attention and engagement, once encoded, these bindings are automatically deployed to guide behavior when the stimulus recurs. These findings expand our understanding of how cognitive control processes are integrated into episodic event files. |
Christoph J. Völter; Ludwig Huber Dogs ' looking times and pupil dilation response reveal expectations about contact causality Journal Article In: Biology Letters, vol. 17, pp. 1–5, 2021. @article{Voelter2021, Contact causality is one of the fundamental principles allowing us to make sense of our physical environment. From an early age, humans perceive spatio-temporally contiguous launching events as causal. Surprisingly little is known about causal perception in non-human animals, particularly outside the primate order. Violation-of-expectation paradigms in combination with eye-tracking and pupillometry have been used to study physical expectations in human infants. In the current study, we establish this approach for dogs (Canis familiaris). We presented dogs with realistic three-dimensional animations of launching events with contact (regular launching event) or without contact between the involved objects. In both conditions, the objects moved with the same timing and kinematic properties. The dogs tracked the object movements closely throughout the study but their pupils were larger in the no-contact condition and they looked longer at the object initiating the launch after the no-contact event compared to the contact event. We conclude that dogs have implicit expectations about contact causality. |
Nathan Van der Stoep; M. J. Van der Smagt; C. Notaro; Z. Spock; M. Naber The additive nature of the human multisensory evoked pupil response Journal Article In: Scientific Reports, vol. 11, pp. 1–12, 2021. @article{VanderStoep2021, Pupillometry has received increased interest for its usefulness in measuring various sensory processes as an alternative to behavioural assessments. This is also apparent for multisensory investigations. Studies of the multisensory pupil response, however, have produced conflicting results. Some studies observed super-additive multisensory pupil responses, indicative of multisensory integration (MSI). Others observed additive multisensory pupil responses even though reaction time (RT) measures were indicative of MSI. Therefore, in the present study, we investigated the nature of the multisensory pupil response by combining methodological approaches of previous studies while using supra-threshold stimuli only. In two experiments we presented auditory and visual stimuli to observers that evoked a(n) (onset) response (be it constriction or dilation) in a simple detection task and a change detection task. In both experiments, the RT data indicated MSI as shown by race model inequality violation. Still, the multisensory pupil response in both experiments could best be explained by linear summation of the unisensory pupil responses. We conclude that the multisensory pupil response for supra-threshold stimuli is additive in nature and cannot be used as a measure of MSI, as only a departure from additivity can unequivocally demonstrate an interaction between the senses. |
Chiara Tortelli; Marco Turi; David C. Burr; Paola Binda Objective pupillometry shows that perceptual styles covary with autistic-like personality traits Journal Article In: eLife, vol. 10, pp. 1–13, 2021. @article{Tortelli2021, 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. |
Chiara Tortelli; Marco Turi; David C. Burr; Paola Binda Pupillary responses obey Emmert's law and co-vary with autistic traits Journal Article In: Journal of Autism and Developmental Disorders, vol. 51, no. 8, pp. 2908–2919, 2021. @article{Tortelli2021a, We measured the pupil response to a light stimulus subject to a size illusion and found that stimuli perceived as larger evoke a stronger pupillary response. The size illusion depends on combining retinal signals with contextual 3D information; contextual processing is thought to vary across individuals, being weaker in individuals with stronger autistic traits. Consistent with this theory, autistic traits correlated negatively with the magnitude of pupil modulations in our sample of neurotypical adults; however, psychophysical measurements of the illusion did not correlate with autistic traits, or with the pupil modulations. This shows that pupillometry provides an accurate objective index of complex perceptual processes, particularly useful for quantifying interindividual differences, and potentially more informative than standard psychophysical measures. |
Anthony Tapper; David Gonzalez; Mina Nouredanesh; Ewa Niechwiej-Szwedo In: Vision Research, vol. 184, pp. 43–51, 2021. @article{Tapper2021, Research shows that concussions cause long-term deficits in executive functions when tested using challenging tasks with high cognitive load. The neurophysiological mechanism(s) associated with executive dysfunction are not well understood. Pupillometry provides a non-invasive index of arousal and cognitive load; therefore, the current study investigated whether pupillometry could help explain the persistent deficits in dual-task performance in individuals with a history of concussion (n = 14) compared to controls (n = 13). Participants were tested using a computerized Corsi block task which increased in difficulty as a function of set size (i.e., number of blocks to be remembered) and task condition (i.e., performed alone and concurrently with an auditory task). Pupil size was measured during the initial fixation prior to the Corsi task to assess arousal level, and during the encoding phase to assess task evoked pupil response. Results showed that: 1) in contrast to the control group, pupil size was not modulated by task condition in the concussed group indicating that arousal level was similar in the single and dual task; 2) task evoked pupil dilation increased as a function of set size in the single task in both groups, 3) in contrast to the control group, those with a history of concussion had similar pupil size during the single and dual task conditions. One interpretation of these results is that individuals with a history of concussion exert greater effort when performing relatively easier tasks, and they reach capacity limits when the cognitive load is lower in comparison to non-concussed individuals. In conclusion, pupillometry may provide insight into persisting deficits in executive functions following concussion(s). |
Jack W. Silcox; Brennan R. Payne The costs (and benefits) of effortful listening on context processing: A simultaneous electrophysiology, pupillometry, and behavioral study Journal Article In: Cortex, vol. 142, pp. 296–316, 2021. @article{Silcox2021, There is an apparent disparity between the fields of cognitive audiology and cognitive electrophysiology as to how linguistic context is used when listening to perceptually challenging speech. To gain a clearer picture of how listening effort impacts context use, we conducted a pre-registered study to simultaneously examine electrophysiological, pupillometric, and behavioral responses when listening to sentences varying in contextual constraint and acoustic challenge in the same sample. Participants (N = 44) listened to sentences that were highly constraining and completed with expected or unexpected sentence-final words (“The prisoners were planning their escape/party”) or were low-constraint sentences with unexpected sentence-final words (“All day she thought about the party”). Sentences were presented either in quiet or with +3 dB SNR background noise. Pupillometry and EEG were simultaneously recorded and subsequent sentence recognition and word recall were measured. While the N400 expectancy effect was diminished by noise, suggesting impaired real-time context use, we simultaneously observed a beneficial effect of constraint on subsequent recognition memory for degraded speech. Importantly, analyses of trial-to-trial coupling between pupil dilation and N400 amplitude showed that when participants' showed increased listening effort (i.e., greater pupil dilation), there was a subsequent recovery of the N400 effect, but at the same time, higher effort was related to poorer subsequent sentence recognition and word recall. Collectively, these findings suggest divergent effects of acoustic challenge and listening effort on context use: while noise impairs the rapid use of context to facilitate lexical semantic processing in general, this negative effect is attenuated when listeners show increased effort in response to noise. However, this effort-induced reliance on context for online word processing comes at the cost of poorer subsequent memory. |
Adi Shechter; David L. Share Keeping an eye on effort: A pupillometric investigation of effort and effortlessness in visual word recognition Journal Article In: Psychological Science, vol. 32, no. 1, pp. 80–95, 2021. @article{Shechter2021a, Rapid and seemingly effortless word recognition is a virtually unquestioned characteristic of skilled reading, yet the definition and operationalization of the concept of cognitive effort have proven elusive. We investigated the cognitive effort involved in oral and silent word reading using pupillometry among adults (Experiment 1 |
Omer Sharon; Firas Fahoum; Yuval Nir Transcutaneous vagus nerve stimulation in humans induces pupil dilation and attenuates alpha oscillations Journal Article In: Journal of Neuroscience, vol. 41, no. 2, pp. 320–330, 2021. @article{Sharon2021, Vagus nerve stimulation (VNS) is widely used to treat drug-resistant epilepsy and depression. While the precise mechanisms mediating its long-term therapeutic effects are not fully resolved, they likely involve locus coeruleus (LC) stimulation via the nucleus of the solitary tract, which receives afferent vagal inputs. In rats, VNS elevates LC firing and forebrain noradrenaline levels, whereas LC lesions suppress VNS therapeutic efficacy. Noninvasive transcutaneous VNS (tVNS) uses electrical stimulation that targets the auricular branch of the vagus nerve at the cymba conchae of the ear. However, the extent to which tVNS mimics VNS remains unclear. Here, we investigated the short-term effects of tVNS in healthy human male volunteers (n = 24), using high-density EEG and pupillometry during visual fixation at rest. We compared short (3.4 s) trials of tVNS to sham electrical stimulation at the earlobe (far from the vagus nerve branch) to control for somatosensory stimulation. Although tVNS and sham stimulation did not differ in subjective intensity ratings, tVNS led to robust pupil dilation (peaking 4-5 s after trial onset) that was significantly higher than following sham stimulation. We further quantified, using parallel factor analysis, how tVNS modulates idle occipital alpha (8-13Hz) activity identified in each participant. We found greater attenuation of alpha oscillations by tVNS than by sham stimulation. This demonstrates that tVNS reliably induces pupillary and EEG markers of arousal beyond the effects of somatosensory stimulation, thus supporting the hypothesis that tVNS elevates noradrenaline and other arousal-promoting neuromodulatory signaling, and mimics invasive VNS. |
Christoph Scheffel; Sven-Thomas Graupner; Anne Gärtner; Josephine Zerna; Alexander Strobel; Denise Dörfel Effort beats effectiveness in emotion regulation choice: Differences between suppression and distancing in subjective and physiological measures Journal Article In: Psychophysiology, vol. 58, no. 11, pp. e13908, 2021. @article{Scheffel2021, Emotion regulation (ER) can be implemented by different strategies which differ in their capacity to alter emotional responding. What all strategies have in common is that cognitive control must be exercised in order to implement them. The aim of the present preregistered study was to investigate whether the two ER strategies, expressive suppression and distancing, require different amounts of cognitive effort and whether effort is associated with personality traits. Effort was assessed subjectively via ratings and objectively via pupillometry and heart period. In two studies |
Dorothee Pöhlchen; Marthe Priouret; Miriam S. Kraft; Florian P. Binder; Deniz A. Gürsel; Götz Berberich; Kathrin Koch; Victor I. Spoormaker In: Frontiers in Psychiatry, vol. 12, pp. 730742, 2021. @article{Poehlchen2021, Obsessive-compulsive disorder (OCD) is characterized by recurrent, persistent thoughts and repetitive behaviors causing stress and anxiety. In the associative learning model of OCD, mechanisms of fear extinction are supposed to partly underlie symptom development, maintenance and treatment of OCD, proposing that OCD patients suffer from rigid memory associations and inhibitory learning deficits. To test these assumptions, previous studies have used skin conductance and subjective ratings as readouts in fear conditioning paradigms, finding impaired fear extinction learning, impaired fear extinction recall or no differences between individuals with OCD and healthy controls. Against this heterogeneous background, we tested fear acquisition and extinction in 37 OCD patients and 56 healthy controls, employing skin conductance as well as pupillometry and startle electromyography. Extinction recall was also included in a subsample. We did not observe differences between groups in any of the task phases, except a trend toward higher startle amplitudes during extinction for OCD. Overall, sensitive readouts such as pupillometry and startle responses did not provide evidence for moderate-to-large inhibitory learning deficits using classical fear conditioning, challenging the assumption of generically impaired extinction learning and memory in OCD. |
Benjamin A. Parris; Nabil Hasshim; Zoltan Dienes In: European Journal of Neuroscience, vol. 53, no. 8, pp. 2819–2834, 2021. @article{Parris2021, The mechanisms underpinning the apparently remarkable levels of cognitive and behavioural control following hypnosis and hypnotic suggestion are poorly understood. Numerous independent studies have reported that Stroop interference can be reduced following a post-hypnotic suggestion that asks participants to perceive words as if made up of characters from a foreign language. This effect indicates that frontal executive functions can be more potent than is generally accepted and has been described as resulting from top-down control not normally voluntarily available. We employed eye tracking and pupillometry to investigate whether the effect results from voluntary visuo-attentional strategies (subtly looking away from the word to prevent optimal word processing), reduced response conflict but not overall conflict, Stroop effects being pushed from response selection to response execution (response durations) or increased proactive effortful control given enhanced contextual motivation (as indexed via pupil dilation). We replicated the reduction in Stroop interference following the suggestion despite removing any trials on which eye movements were not consistent with optimal word processing. Our data were inconclusive with regards to conflict type affected by the suggestion in the latency data, although preserved semantic conflict was evident in the pupil data. There was also no evidence of Stroop effects on response durations. However, we show that baseline-corrected pupil sizes were larger following the suggestion indicating the socio-cognitive context and experimental demands motivate participants to marshal greater effortful control. |
Verónica Mäki-Marttunen Pupil-based states of brain integration across cognitive states Journal Article In: Neuroscience, vol. 471, pp. 61–71, 2021. @article{MaekiMarttunen2021, Arousal is a potent mechanism that provides the brain with functional flexibility and adaptability to external conditions. Within the wake state, arousal levels driven by activity in the neuromodulatory systems are related to specific signatures of neural activation and brain synchrony. However, direct evidence is still lacking on the varying effects of arousal on macroscopic brain characteristics and across a variety of cognitive states in humans. Using a concurrent fMRI-pupillometry approach, we used pupil size as a proxy for arousal and obtained patterns of brain integration associated with increasing arousal levels. We carried out this analysis on resting-state data and data from two attentional tasks implicating different cognitive processes. We found that an increasing level of arousal was related to a state of increased brain integration. This effect was prominent in the salience, visual and default-mode networks in all conditions, while other regions showed task-specificity. Increased integration in the salience network was also related to faster pupil dilation in the two attentional tasks. Furthermore, task performance was related to arousal level, with lower accuracy at higher level of arousal. Taken together, our study provides evidence in humans for pupil size as an index of brain network state, and supports the role of arousal as a switch that drives brain coordination in specific brain regions according to the cognitive state. |
Wouter Kruijne; Christian N. L. Olivers; Hedderik Rijn Neural repetition suppression modulates time perception: Evidence from electrophysiology and pupillometry Journal Article In: Journal of Cognitive Neuroscience, vol. 33, no. 7, pp. 1230–1252, 2021. @article{Kruijne2021, Human time perception is malleable and subject to many biases. For example, it has repeatedly been shown that stimuli that are physically intense or that are unexpected seem to last longer. Two competing hypotheses have been proposed to account for such biases: One states that these temporal illusions are the result of increased levels of arousal that speeds up neural clock dynamics, whereas the alternative “magnitude coding” account states that the magnitude of sensory responses causally modulates perceived durations. Common experimental paradigms used to study temporal biases cannot dissociate between these accounts, as arousal and sensory magnitude covary and modulate each other. Here, we present two temporal discrimination experiments where two flashing stimuli demarcated the start and end of a to-be-timed interval. These stimuli could be either in the same or a different location, which led to different sensory responses because of neural repetition suppression. Crucially, changes and repetitions were fully predictable, which allowed us to explore effects of sensory response magnitude without changes in arousal or surprise. Intervals with changing markers were perceived as lasting longer than those with repeating markers. We measured EEG (Experiment 1) and pupil size (Experiment 2) and found that temporal perception was related to changes in ERPs (P2) and pupil constriction, both of which have been related to responses in the sensory cortex. Conversely, correlates of surprise and arousal (P3 amplitude and pupil dilation) were unaffected by stimulus repetitions and changes. These results demonstrate, for the first time, that sensory magnitude affects time perception even under constant levels of arousal. |
Elif Canseza Kaplan; Anita E. Wagner; Paolo Toffanin; Deniz Başkent Do musicians and non-musicians differ in speech-on-speech processing? Journal Article In: Frontiers in Psychology, vol. 12, pp. 623787, 2021. @article{Kaplan2021, Earlier studies have shown that musically trained individuals may have a benefit in adverse listening situations when compared to non-musicians, especially in speech-on-speech perception. However, the literature provides mostly conflicting results. In the current study, by employing different measures of spoken language processing, we aimed to test whether we could capture potential differences between musicians and non-musicians in speech-on-speech processing. We used an offline measure of speech perception (sentence recall task), which reveals a post-task response, and online measures of real time spoken language processing: gaze-tracking and pupillometry. We used stimuli of comparable complexity across both paradigms and tested the same groups of participants. In the sentence recall task, musicians recalled more words correctly than non-musicians. In the eye-tracking experiment, both groups showed reduced fixations to the target and competitor words' images as the level of speech maskers increased. The time course of gaze fixations to the competitor did not differ between groups in the speech-in-quiet condition, while the time course dynamics did differ between groups as the two-talker masker was added to the target signal. As the level of two-talker masker increased, musicians showed reduced lexical competition as indicated by the gaze fixations to the competitor. The pupil dilation data showed differences mainly in one target-to-masker ratio. This does not allow to draw conclusions regarding potential differences in the use of cognitive resources between groups. Overall, the eye-tracking measure enabled us to observe that musicians may be using a different strategy than non-musicians to attain spoken word recognition as the noise level increased. However, further investigation with more fine-grained alignment between the processes captured by online and offline measures is necessary to establish whether musicians differ due to better cognitive control or sound processing. |
Michael A. Johns; Paola E. Dussias Of revistas and magacínes: Lexical competition in the online processing of established loanwords Journal Article In: Journal of Second Language Studies, vol. 4, no. 2, pp. 375–411, 2021. @article{Johns2021, The transfer of words from one language to another is ubiquitous in many of the world's languages. While loanwords have a rich literature in the fields of historical linguistics, language contact, and sociolinguistics, little work has been done examining how loanwords are processed by bilinguals with knowledge of both the source and recipient languages. The present study uses pupillometry to compare the online processing of established loanwords in Puerto Rican Spanish to native Spanish words by highly proficient Puerto Rican Spanish-English bilinguals. Established loanwords elicited a significantly larger pupillary response than native Spanish words, with the pupillary response modulated by both the frequency of the loanword itself and of the native Spanish counterpart. These findings suggest that established loanwords are processed differently than native Spanish words and compete with their native equivalents, potentially due to both intra- and inter-lingual effects of saliency. |
Mega B. Herlambang Id; Fokie Cnossen; Niels A. Taatgen The effects of intrinsic motivation on mental fatigue Journal Article In: PLoS ONE, vol. 16, no. 1, pp. e0243754, 2021. @article{Id2021, There have been many studies attempting to disentangle the relation between motivation and mental fatigue. Mental fatigue occurs after performing a demanding task for a prolonged time, and many studies have suggested that motivation can counteract the negative effects of mental fatigue on task performance. To complicate matters, most mental fatigue studies looked exclusively at the effects of extrinsic motivation but not intrinsic motivation. Individu- als are said to be extrinsically motivated when they perform a task to attain rewards and avoid punishments, while they are said to be intrinsically motivated when they do for the pleasure of doing the activity. To assess whether intrinsic motivation has similar effects as extrinsic motivation, we conducted an experiment using subjective, performance, and physi- ological measures (heart rate variability and pupillometry). In this experiment, 28 partici- pants solved Sudoku puzzles on a computer for three hours, with a cat video playing in the corner of the screen. The experiment consisted of 14 blocks with two alternating conditions: low intrinsic motivation and high intrinsic motivation. The main results showed that irrespec- tive of condition, participants reported becoming fatigued over time. They performed better, invested more mental effort physiologically, and were less distracted in high-level than in low-level motivation blocks. The results suggest that similarly to extrinsic motivation, time- on-task effects are modulated by the level of intrinsic motivation: With high intrinsic motiva- tion, people can maintain their performance over time as they seem willing to invest more effort as time progresses than in low intrinsic motivation. |