All EyeLink Eye Tracker Publications
All 13,000+ peer-reviewed EyeLink research publications up until 2024 (with some early 2025s) are listed below by year. You can search the publications library using keywords such as Visual Search, Smooth Pursuit, Parkinson’s, etc. You can also search for individual author names. Eye-tracking studies grouped by research area can be found on the solutions pages. If we missed any EyeLink eye-tracking papers, please email us!
2022 |
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. |
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. |
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, & 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, & 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. |
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, & 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 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. |
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. |
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. |
Serena K. Mon; Mira Nencheva; Francesca M. M. Citron; Casey Lew-Williams; Adele E. Goldberg Conventional metaphors elicit greater real-time engagement than literal paraphrases or concrete sentences Journal Article In: Journal of Memory and Language, vol. 121, pp. 104285, 2021. @article{Mon2021,Conventional metaphors (e.g., a firm grasp on an idea) are extremely common. A possible explanation for their ubiquity is that they are more engaging, evoking more focused attention, than their literal paraphrases (e.g., a good understanding of an idea). To evaluate whether, when, and why this may be true, we created a new database of 180 English sentences consisting of conventional metaphors, literal paraphrases, and concrete descriptions (e.g., a firm grip on a doorknob). Extensive norming matched differences across sentence types in complexity, plausibility, emotional valence, intensity, and familiarity of the key phrases. Then, using pupillometry to study the time course of metaphor processing, we predicted that metaphors would elicit greater event-evoked pupil dilation compared to other sentence types. Results confirmed the predicted increase beginning at the onset of the key phrase and lasting seconds beyond the end of the sentence. When metaphorical and literal sentences were compared directly in survey data, participants judged metaphorical sentences to convey “richer meaning,” but not more information. We conclude that conventional metaphors are more engaging than literal paraphrases or concrete sentences in a way that is irreducible to difficulty or ease, amount of information, short-term lexical access, or downstream inferences. |
Alice Milne; Sijia Zhao; Christina Tampakaki; Gabriela Bury; Maria Chait Sustained pupil responses are modulated by predictability of auditory sequences Journal Article In: Journal of Neuroscience, vol. 41, no. 28, pp. 6116–6127, 2021. @article{Milne2021,The brain is highly sensitive to auditory regularities and exploits the predictable order of sounds in many situations, from parsing complex auditory scenes, to the acquisition of language. To understand the impact of stimulus predictability on perception, it is important to determine how the detection of predictable structure influences processing and attention. Here we use pupillometry to gain insight into the effect of sensory regularity on arousal. Pupillometry is a commonly used measure of salience and processing effort, with more perceptually salient or perceptually demanding stimuli consistently associated with larger pupil diameters. In two experiments we tracked human listeners' pupil dynamics while they listened to sequences of 50ms tone pips of different frequencies. The order of the tone pips was either random, contained deterministic (fully predictable) regularities (experiment 1 |
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. |
Isabell Hubert Lyall; Juhani Järvikivi Individual differences in political ideology and disgust sensitivity affect real-time spoken language comprehension Journal Article In: Frontiers in Psychology, vol. 12, pp. 699071, 2021. @article{HubertLyall2021,Individuals' moral views have been shown to affect their event-related potentials (ERP) response to spoken statements, and people's political ideology has been shown to guide their sentence completion behavior. Using pupillometry, we asked whether political ideology and disgust sensitivity affect online spoken language comprehension. 60 native speakers of English listened to spoken utterances while their pupil size was tracked. Some of those utterances contained grammatical errors, semantic anomalies, or socio-cultural violations, statements incongruent with existing gender stereotypes and perceived speaker identity, such as “I sometimes buy my bras at Hudson's Bay,” spoken by a male speaker. An individual's disgust sensitivity is associated with the Behavioral Immune System, and may be correlated with socio-political attitudes, for example regarding out-group stigmatization. We found that more disgust-sensitive individuals showed greater pupil dilation with semantic anomalies and socio-cultural violations. However, political views differently affected the processing of the two types of violations: whereas more conservative listeners showed a greater pupil response to socio-cultural violations, more progressive listeners engaged more with semantic anomalies, but this effect appeared much later in the pupil record. |
Marcus Grueschow; Nico Stenz; Hanna Thörn; Ulrike Ehlert; Jan Breckwoldt; Monika Brodmann Maeder; Aristomenis K. Exadaktylos; Roland Bingisser; Christian C. Ruff; Birgit Kleim Real-world stress resilience is associated with the responsivity of the locus coeruleus Journal Article In: Nature Communications, vol. 12, pp. 2275, 2021. @article{Grueschow2021,Individuals may show different responses to stressful events. Here, we investigate the neurobiological basis of stress resilience, by showing that neural responsitivity of the noradrenergic locus coeruleus (LC-NE) and associated pupil responses are related to the subsequent change in measures of anxiety and depression in response to prolonged real-life stress. We acquired fMRI and pupillometry data during an emotional-conflict task in medical residents before they underwent stressful emergency-room internships known to be a risk factor for anxiety and depression. The LC-NE conflict response and its functional coupling with the amygdala was associated with stress-related symptom changes in response to the internship. A similar relationship was found for pupil-dilation, a potential marker of LC-NE firing. Our results provide insights into the noradrenergic basis of conflict generation, adaptation and stress resilience. |
Josephine M. Groot; Nya M. Boayue; Gábor Csifcsák; Wouter Boekel; René Huster; Birte U. Forstmann; Matthias Mittner Probing the neural signature of mind wandering with simultaneous fMRI-EEG and pupillometry Journal Article In: NeuroImage, vol. 224, pp. 117412, 2021. @article{Groot2021,Mind wandering reflects the shift in attentional focus from task-related cognition driven by external stimuli toward self-generated and internally-oriented thought processes. Although such task-unrelated thoughts (TUTs) are pervasive and detrimental to task performance, their underlying neural mechanisms are only modestly understood. To investigate TUTs with high spatial and temporal precision, we simultaneously measured fMRI, EEG, and pupillometry in healthy adults while they performed a sustained attention task with experience sampling probes. Features of interest were extracted from each modality at the single-trial level and fed to a support vector machine that was trained on the probe responses. Compared to task-focused attention, the neural signature of TUTs was characterized by weaker activity in the default mode network but elevated activity in its anticorrelated network, stronger functional coupling between these networks, widespread increase in alpha, theta, delta, but not beta, frequency power, predominantly reduced amplitudes of late, but not early, event-related potentials, and larger baseline pupil size. Particularly, information contained in dynamic interactions between large-scale cortical networks was predictive of transient changes in attentional focus above other modalities. Together, our results provide insight into the spatiotemporal dynamics of TUTs and the neural markers that may facilitate their detection. |
Jan Grenzebach; Thomas G. G. Wegner; Wolfgang Einhäuser; Alexandra Bendixen Pupillometry in auditory multistability Journal Article In: PLoS ONE, vol. 16, no. 6, pp. e0230039, 2021. @article{Grenzebach2021,In multistability, a constant stimulus induces alternating perceptual interpretations. For many forms of visual multistability, the transition from one interpretation to another ("perceptual switch") is accompanied by a dilation of the pupil. Here we ask whether the same holds for auditory multistability, specifically auditory streaming. Two tones were played in alternation, yielding four distinct interpretations: the tones can be perceived as one integrated percept (single sound source), or as segregated with either tone or both tones in the foreground. We found that the pupil dilates significantly around the time a perceptual switch is reported ("multistable condition"). When participants instead responded to actual stimulus changes that closely mimicked the multistable perceptual experience ("replay condition"), the pupil dilated more around such responses than in multistability. This still held when data were corrected for the pupil response to the stimulus change as such. Hence, active responses to an exogeneous stimulus change trigger a stronger or temporally more confined pupil dilation than responses to an endogenous perceptual switch. In another condition, participants randomly pressed the buttons used for reporting multistability. In Study 1, this "random condition"failed to sufficiently mimic the temporal pattern of multistability. By adapting the instructions, in Study 2 we obtained a response pattern more similar to the multistable condition. In this case, the pupil dilated significantly around the random button presses. Albeit numerically smaller, this pupil response was not significantly different from the multistable condition. While there are several possible explanations-related, e.g., to the decision to respond-this underlines the difficulty to isolate a purely perceptual effect in multistability. Our data extend previous findings from visual to auditory multistability. They highlight methodological challenges in interpreting such data and suggest possible approaches to meet them, including a novel stimulus to simulate the experience of perceptual switches in auditory streaming. |
Steven M. Gillespie; Ian J. Mitchell; Anthony R. Beech; Pia Rotshtein Processing of emotional faces in sexual offenders with and without child victims: An eye-tracking study with pupillometry Journal Article In: Biological Psychology, vol. 163, pp. 108141, 2021. @article{Gillespie2021,Socio-affective dysfunction is a risk-factor for sexual offense recidivism. However, it remains unknown whether men who have sexually offended with and without child victims show differences in eye scan paths and autonomic responsivity while viewing facial expressions of emotion. We examined differences in accuracy of emotion recognition, eye movements, and pupil dilation responses between sex offenders with child victims, sex offenders without child victims, and a group of non-offenders living in the community. Sex offenders without child victims looked for longer at the eyes than sex offenders with child victims and non-offenders. Men without child victims also scored higher for psychopathy linked disinhibition, and these traits were associated with looking longer at the eyes of afraid faces. We found no evidence for group differences in accuracy, visual attention to the mouth, or pupil dilation responses. Our findings have implications for understanding the nature of socio-affective dysfunction in sexual offenders. |
Wendel M. Friedl; Andreas Keil Aversive conditioning of spatial position sharpens neural population-level tuning in visual cortex and selectively alters alpha-band activity Journal Article In: Journal of Neuroscience, vol. 41, no. 26, pp. 5723–5733, 2021. @article{Friedl2021,Processing capabilities for many low-level visual features are experientially malleable, aiding sighted organisms in adapting to dynamic environments. Explicit instructions to attend a specific visual field location influence retinotopic visuocortical activity, amplifying responses to stimuli appearing at cued spatial positions. It remains undetermined both how such prioritization affects surrounding nonprioritized locations, and if a given retinotopic spatial position can attain enhanced cortical representation through experience rather than instruction. The current report examined visuocortical response changes as human observers (N = 51, 19 male) learned, through differential classical conditioning, to associate specific screen locations with aversive outcomes. Using dense-array EEG and pupillometry, we tested the preregistered hypotheses of either sharpening or generalization around an aversively associated location following a single conditioning session. Competing hypotheses tested whether mean response changes would take the form of a Gaussian (generalization) or difference-of-Gaussian (sharpening) distribution over spatial positions, peaking at the viewing location paired with a noxious noise. Occipital 15 Hz steady-state visual evoked potential responses were selectively heightened when viewing aversively paired locations and displayed a nonlinear, difference-of-Gaussian profile across neighboring locations, consistent with suppressive surround modulation of nonprioritized positions. Measures of alpha-band (8-12 Hz) activity were differentially altered in anterior versus posterior locations, while pupil diameter exhibited selectively heightened responses to noise-paired locations but did not evince differences across the nonpaired locations. These results indicate that visuocortical spatial representations are sharpened in response to location-specific aversive conditioning, while top-down influences indexed by alpha-power reduction exhibit posterior generalization and anterior sharpening. |
Gerardo Fernández; Mario A. Parra Oculomotor behaviors and integrative memory functions in the Alzheimer's Clinical Syndrome Journal Article In: Journal of Alzheimer's Disease, vol. 82, no. 3, pp. 1033–1044, 2021. @article{Fernandez2021,Background: Biological information drawn from eye-tracking metrics is providing evidence regarding drivers of cognitive decline in Alzheimer's disease. In particular, pupil size has proved useful to investigate cognitive performance during online activities. Objective: To investigate the oculomotor correlates of impaired performance of patients with mild Alzheimer's Clinical Syndrome (ACS) on a recently developed memory paradigm, namely the Short-Term Memory Binding Test (STMBT). Methods: We assessed a sample of eighteen healthy controls (HC) and eighteen patients with a diagnosis of mild ACS with the STMBT while we recorded their oculomotor behaviors using pupillometry and eye-tracking. Results: As expected, a group (healthy controls versus ACS) by condition (Unbound Colours versus Bound Colours) interaction was found whereby behavioral group differences were paramount in the Bound Colours condition. Healthy controls' pupils dilated significantly more in the Bound Colours than in the Unbound Colours condition, a discrepancy not observed in ACS patients. Furthermore, ROC analysis revealed the abnormal pupil behaviors distinguished ACS patients from healthy controls with values of sensitivity and specify of 100%, thus outperforming both recognition scores and gaze duration. Conclusion: The biological correlates of Short-Term Memory Binding impairments appear to involve a network much wider than we have thought to date, which expands across cortical and subcortical structures. We discuss these findings focusing on their implications for our understanding of neurocognitive phenotypes in the preclinical stages of Alzheimer's disease and potential development of cognitive biomarkers that can support ongoing initiatives to prevent dementia. |
J. C. F. Winter; S. M. Petermeijer; L. Kooijman; D. Dodou Replicating five pupillometry studies of Eckhard Hess Journal Article In: International Journal of Psychophysiology, vol. 165, pp. 145–205, 2021. @article{Winter2021,Several papers by Eckhard Hess from the 1960s and 1970s report that the pupils dilate or constrict according to the interest value, arousing content, or mental demands of visual stimuli. However, Hess mostly used small sample sizes and undocumented luminance control. In a first experiment (N = 182) and a second preregistered experiment (N = 147), we replicated five studies of Hess using modern equipment. Our experiments (1) did not support the hypothesis of gender differences in pupil diameter change with respect to baseline (PC) when viewing stimuli of different interest value, (2) showed that solving more difficult multiplications yields a larger PC in the seconds before providing an answer and a larger maximum PC, but a smaller PC at a fixed time after the onset of the multiplication, (3) did not support the hypothesis that participants' PC mimics the pupil diameter in a pair of schematic eyes but not in single-eyed or three-eyed stimuli, (4) did not support the hypothesis of gender differences in PC when watching a video of a male trying to escape a mob, and (5) supported the hypothesis that arousing words yield a higher PC than non-arousing words. Although we did not observe consistent gender differences in PC, additional analyses showed gender differences in eye movements towards erogenous zones. Furthermore, PC strongly correlated with the luminance of the locations where participants looked. Overall, our replications confirm Hess's findings that pupils dilate in response to mental demands and stimuli of an arousing nature. Hess's hypotheses regarding pupil mimicry and gender differences in pupil dilation did not replicate. |
Kevin Silva Castanheira; Myle LoParco; A. Ross Otto Task-evoked pupillary responses track effort exertion: Evidence from task-switching Journal Article In: Cognitive, Affective, & Behavioral Neuroscience, vol. 21, no. 3, pp. 592–606, 2021. @article{daSilvaCastanheira2021,A spate of research has examined how individuals regulate effortful processing in service of goal-directed behaviors. One key challenge in developing an account of this regulation is quantifying the momentary amount of cognitive effort exerted by an individual in service of their goals. A growing body of literature has suggested using task-evoked pupil dilations as a potential psychophysiological index of cognitive effort; however, it remains unclear whether pupil diameter indexes effort exertion or merely reflects task load, as both are tightly intertwined. Here, we attempt to disentangle these disparate accounts of pupil diameter by leveraging individual differences in executive function (as measured by Stroop interference) and a motivational manipulation (i.e., monetary incentives) while participants complete a task-switching paradigm. In line with both the effort and demand accounts, we observed larger task-evoked pupillary responses (TEPRs) for trials in which there was a task switch versus a task repetition. Additionally, we found that larger phasic pupillary responses at baseline (without reward incentives) predicted smaller switch costs. Mirroring this pattern, individual differences in reward-induced switch cost reductions were predicted by reward-induced increases in phasic pupil diameter. Finally, we observed that the interrelationship between effort and pupil diameter at baseline was modulated by individual differences in Stroop interference costs. Together, these findings provide support for an effort account of TEPRs, and suggest that pupillometry is a viable index of cognitive effort. |
Sarah Colby; Bob McMurray Cognitive and physiological measures of listening effort during degraded speech perception: Relating dual-task and pupillometry paradigms Journal Article In: Journal of Speech, Language, and Hearing Research, vol. 64, no. 9, pp. 3627–3652, 2021. @article{Colby2021,Purpose: Listening effort is quickly becoming an important metric for assessing speech perception in less-than-ideal situations. However, the relationship between the construct of listening effort and the measures used to assess it remains unclear. We compared two measures of listening effort: a cognitive dual task and a physiological pupillometry task. We sought to investigate the relationship between these measures of effort and whether engaging effort impacts speech accuracy. Method: In Experiment 1, 30 participants completed a dual task and a pupillometry task that were carefully matched in stimuli and design. The dual task consisted of a spoken word recognition task and a visual match-to-sample task. In the pupillometry task, pupil size was monitored while participants completed a spoken word recognition task. Both tasks presented words at three levels of listening difficulty (unmodified, eight-channel vocoding, and four-channel vocoding) and provided response feedback on every trial. We refined the pupillometry task in Experiment 2 (n = 31); crucially, participants no longer received response feedback. Finally, we ran a new group of subjects on both tasks in Experiment 3 (n = 30). Results: In Experiment 1, accuracy in the visual task decreased with increased signal degradation in the dual task, but pupil size was sensitive to accuracy and not vocoding condition. After removing feedback in Experiment 2, changes in pupil size were predicted by listening condition, suggesting the task was now sensitive to engaged effort. Both tasks were sensitive to listening difficulty in Experiment 3, but there was no relationship between the tasks and neither task predicted speech accuracy. Conclusions: Consistent with previous work, we found little evidence for a relationship between different measures of listening effort. We also found no evidence that effort predicts speech accuracy, suggesting that engaging more effort does not lead to improved speech recognition. Cognitive and physiological measures of listening effort are likely sensitive to different aspects of the construct of listening effort. |
Emily A. Burg; Tanvi Thakkar; Taylor Fields; Sara M. Misurelli; Stefanie E. Kuchinsky; Joseph Roche; Daniel J. Lee; Ruth Y. Litovsky Systematic comparison of trial exclusion criteria for pupillometry data analysis in individuals with single-sided deafness and normal hearing Journal Article In: Trends in Hearing, vol. 25, pp. 1–17, 2021. @article{Burg2021,The measurement of pupil dilation has become a common way to assess listening effort. Pupillometry data are subject to artifacts, requiring highly contaminated data to be discarded from analysis. It is unknown how trial exclusion criteria impact experimental results. The present study examined the effect of a common exclusion criterion, percentage of blinks, on speech intelligibility and pupil dilation measures in 9 participants with single-sided deafness (SSD) and 20 participants with normal hearing. Participants listened to and repeated sentences in quiet or with speech maskers. Pupillometry trials were processed using three levels of blink exclusion criteria: 15%, 30%, and 45%. These percentages reflect a threshold for missing data points in a trial, where trials that exceed the threshold are excluded from analysis. Results indicated that pupil dilation was significantly greater and intelligibility was significantly lower in the masker compared with the quiet condition for both groups. Across-group comparisons revealed that speech intelligibility in the SSD group decreased significantly more than the normal hearing group from quiet to masker conditions, but the change in pupil dilation was similar for both groups. There was no effect of blink criteria on speech intelligibility or pupil dilation results for either group. However, the total percentage of blinks in the masker condition was significantly greater than in the quiet condition for the SSD group, which is consistent with previous studies that have found a relationship between blinking and task difficulty. This association should be carefully considered in future experiments using pupillometry to gauge listening effort. |
Nicolai D. Ayasse; Alana J. Hodson; Arthur Wingfield The principle of least effort and comprehension of spoken sentences by younger and older adults Journal Article In: Frontiers in Psychology, vol. 12, pp. 629464, 2021. @article{Ayasse2021,There is considerable evidence that listeners' understanding of a spoken sentence need not always follow from a full analysis of the words and syntax of the utterance. Rather, listeners may instead conduct a superficial analysis, sampling some words and using presumed plausibility to arrive at an understanding of the sentence meaning. Because this latter strategy occurs more often for sentences with complex syntax that place a heavier processing burden on the listener than sentences with simpler syntax, shallow processing may represent a resource conserving strategy reflected in reduced processing effort. This factor may be even more important for older adults who as a group are known to have more limited working memory resources. In the present experiment, 40 older adults (Mage = 75.5 years) and 20 younger adults (Mage = 20.7) were tested for comprehension of plausible and implausible sentences with a simpler subject-relative embedded clause structure or a more complex object-relative embedded clause structure. Dilation of the pupil of the eye was recorded as an index of processing effort. Results confirmed greater comprehension accuracy for plausible than implausible sentences, and for sentences with simpler than more complex syntax, with both effects amplified for the older adults. Analysis of peak pupil dilations for implausible sentences revealed a complex three-way interaction between age, syntactic complexity, and plausibility. Results are discussed in terms of models of sentence comprehension, and pupillometry as an index of intentional task engagement. |
Naila Ayala; Ewa Niechwiej-Szwedo Effects of blocked vs. interleaved administration mode on saccade preparatory set revealed using pupillometry Journal Article In: Experimental Brain Research, vol. 239, no. 1, pp. 245–255, 2021. @article{Ayala2021a,Eye movements have been used extensively to assess information processing and cognitive function. However, significant variability in saccade performance has been observed, which could arise from methodological variations across different studies. For example, prosaccades and antisaccades have been studied using either a blocked or interleaved design, which has a significant influence on error rates and latency. This is problematic as it makes it difficult to compare saccade performance across studies and may limit the ability to use saccades as a behavioural assay to assess neurocognitive function. Thus, the current study examined how administration mode influences saccade related preparatory activity by employing pupil size as a non-invasive proxy for neural activity related to saccade planning and execution. Saccade performance and pupil dynamics were examined in eleven participants as they completed pro- and antisaccades in blocked and interleaved paradigms. Results showed that administration mode significantly modulated saccade performance and preparatory activity. Reaction times were longer for both pro- and antisaccades in the interleaved condition, compared to the blocked condition (p < 0.05). Prosaccade pupil dilations were larger in the interleaved condition (p < 0.05), while antisaccade pupil dilations did not significantly differ between administration modes. Additionally, ROC analysis provided preliminary evidence that pupil size can effectively predict saccade directional errors prior to saccade onset. We propose that task-evoked pupil dilations reflect an increase in preparatory activity for prosaccades and the corresponding cognitive demands associated with interleaved administration mode. Overall, the results highlight the importance that administration mode plays in the design of neurocognitive tasks. |
Naila Ayala; Matthew Heath Pupillometry reveals the role of arousal in a postexercise benefit to executive function Journal Article In: Brain Sciences, vol. 11, no. 8, pp. 1–10, 2021. @article{Ayala2021,A single bout of aerobic exercise improves executive function; however, the mechanism(s) underlying this improvement remains unclear. Here, we employed a 20-min bout of aerobic exercise, and at pre-and immediate post-exercise sessions examined executive function via pro-(i.e., saccade to veridical target location) and anti-saccade (i.e., saccade mirror symmetrical to a target) performance and pupillometry metrics. Notably, tonic and phasic pupillometry responses in oculomotor control provided a framework to determine the degree that arousal and/or executive resource recruitment influence behavior. Results demonstrated a pre-to post-exercise decrease in pro-and anti-saccade reaction times (p = 0.01) concurrent with a decrease and increase in tonic baseline pupil size and task-evoked pupil dilations, respectively (ps < 0.03). Such results demonstrate that an exercise-induced improvement in saccade performance is related to an executive-mediated “shift” in physiological and/or psychological arousal, supported by the locus coeruleus norepinephrine system to optimize task engagement. |
Thomas Armstrong; Sara Federman; Kari Hampson; Owen Crabtree; Bunmi O. Olatunji Fear learning in veterans with combat-related PTSD is linked to anxiety sensitivity: Evidence from self-report and pupillometry Journal Article In: Behavior Therapy, vol. 52, no. 1, pp. 149–161, 2021. @article{Armstrong2021,Several studies have observed heightened Pavlovian fear conditioning in PTSD. However, it is unclear how fear conditioning in PTSD is related to risk factors for the disorder, such as anxiety sensitivity. Fifty-one combat-exposed veterans (20 with PTSD, 31 without PTSD) completed a differential fear conditioning task in which one colored rectangle (CS +) predicted a loud scream (US), whereas a different colored rectangle (CS-) predicted no US. Veterans with PTSD were characterized by greater anxiety to the CS + but not the CS- during acquisition and extinction, and greater US expectancy during the CS + but not the CS- at extinction. Also, veterans with PTSD had greater pupil dilation to both CSs at extinction, but not at acquisition. Anxiety sensitivity was correlated with anxiety and US expectancy in response to the CS +, but not the CS-, at both acquisition and extinction, and also with pupil diameter to both the CS + and CS- at extinction. Nearly all of these relations held when covarying for PTSD symptoms and trait anxiety. These findings suggest that increased fear conditioning in PTSD may be related to elevated anxiety sensitivity. |
Sara Alhanbali; Kevin J. Munro; Piers Dawes; Peter J. Carolan; Rebecca E. Millman In: International Journal of Audiology, vol. 60, no. 10, pp. 762–772, 2021. @article{Alhanbali2021,Objective: Pupillometry is sensitive to cognitive resource allocation and has been used as a potential measure of listening-related effort and fatigue. We investigated associations between peak pupil diameter, pre-stimulus pupil diameter, performance on a listening task, and the dimensionality of self-reported outcomes (task-related listening effort and fatigue). Design: Pupillometry was recorded while participants performed a speech-in-noise task. Participants rated their experience of listening effort and fatigue using the NASA-Task Load Index (NASA-TLX) and the Visual Analogue Scale of Fatigue (VAS-F), respectively. The dimensionality of the NASA-TLX and the VAS-F was investigated using factor analysis. Study sample: 82 participants with either normal hearing or aided hearing impairment (age range: 55–85 years old, 43 male). Results: Hierarchal linear regression analyses suggested that pre-stimulus pupil diameter predicts a dimension of self-reported fatigue, which we interpreted as tiredness/drowsiness, and listening task performance when controlling for hearing level and age: Larger pre-stimulus pupil diameter was associated with less tiredness/drowsiness and better task performance. Conclusion: Pre-stimulus pupil diameter is a potential index of listening fatigue associated with speech processing in challenging listening conditions. To our knowledge, this is the first investigation of the associations between pre-stimulus pupil diameter and self-reported ratings of listening effort and fatigue. |
2020 |
Lisa Wirz; Lars Schwabe Prioritized attentional processing: Acute stress, memory and stimulus emotionality facilitate attentional disengagement Journal Article In: Neuropsychologia, vol. 138, pp. 107334, 2020. @article{Wirz2020,Rapid attentional orienting toward relevant stimuli and efficient disengagement from irrelevant stimuli are critical for survival. Here, we examined the roles of memory processes, emotional arousal and acute stress in attentional disengagement. To this end, 64 healthy participants encoded negative and neutral facial expressions and, after being exposed to a stress or control manipulation, performed an attention task in which they had to disengage from these previously encoded as well as novel face stimuli. During the attention task, electroencephalography (EEG) and pupillometry data were recorded. Our results showed overall faster reaction times after acute stress and when participants had to disengage from emotionally negative or old facial expressions. Further, pupil dilations were larger in response to neutral faces. During disengagement, our EEG data revealed a reduced N2pc amplitude when participants disengaged from neutral compared to negative facial expressions when these were not presented before, as well as earlier onset latencies for the N400f (for disengagement from negative and old faces), the N2pc, and the LPP (for disengagement from negative faces). In addition, early visual processing of negative faces, as reflected in the P1 amplitude, was enhanced specifically in stressed participants. Our findings indicate that attentional disengagement is improved for negative and familiar stimuli and that stress facilitates not only attentional disengagement but also emotional processing in general. Together, these processes may represent important mechanisms enabling efficient performance and rapid threat detection. |
Nicole Wetzel; Wolfgang Einhäuser; Andreas Widmann Picture-evoked changes in pupil size predict learning success in children Journal Article In: Journal of Experimental Child Psychology, vol. 192, pp. 1–18, 2020. @article{Wetzel2020,Episodic memory, the ability to remember past events in time and place, develops during childhood. Much knowledge about the underlying neuronal mechanisms has been gained from methods not suitable for children. We applied pupillometry to study memory encoding and recognition mechanisms. Children aged 8 and 9 years (n = 24) and adults (n = 24) studied a set of visual scenes to later distinguish them from new pictures. Children performed worse than adults, demonstrating immature episodic memory. During memorization, picture-related changes in pupil diameter predicted later successful recognition. This prediction effect was also observed on a single-trial level. During retrieval, novel pictures showed stronger pupil constriction than familiar pictures in both age groups. The statistically independent effects of objective familiarity (previously presented pictures) versus subjective familiarity (pictures evaluated as familiar independent of the prior presentation) suggest dissociable underlying brain mechanisms. In addition, we isolated principal components of the picture-related pupil response that were differently affected by the memorization and retrieval effects. Results are discussed in the context of the maturation of the medial temporal lobe and prefrontal networks. Our results demonstrate the dissociation of distinct contributions to episodic memory with a psychophysiological method that is suitable for a wide age spectrum. |
Max Schneider; Immanuel G. Elbau; Teachawidd Nantawisarakul; Dorothee Pöhlchen; Tanja Brückl; Michael Czisch; Philipp G. Saemann; Michael D. Lee; Elisabeth B. Binder; Victor I. Spoormaker Pupil dilation during reward anticipation is correlated to depressive symptom load in patients with major depressive disorder Journal Article In: Brain Sciences, vol. 10, no. 12, pp. 1–15, 2020. @article{Schneider2020a,Depression is a debilitating disorder with high prevalence and socioeconomic cost, but the brain-physiological processes that are altered during depressive states are not well understood. Here, we build on recent findings in macaques that indicate a direct causal relationship between pupil dilation and anterior cingulate cortex mediated arousal during anticipation of reward. We translated these findings to human subjects with concomitant pupillometry/fMRI in a sample of unmedicated participants diagnosed with major depression and healthy controls. We could show that the upregulation and maintenance of arousal in anticipation of reward was disrupted in patients in a symptom-load dependent manner. We could further show that the failure to maintain reward anticipatory arousal showed state-marker properties, as it tracked the load and impact of depressive symptoms independent of prior diagnosis status. Further, group differences of anticipatory arousal and continuous correlations with symptom load were not traceable only at the level of pupillometric responses, but were mirrored also at the neural level within salience network hubs. The upregulation and maintenance of arousal during reward anticipation is a novel translational and well-traceable process that could prove a promising gateway to a physiologically informed patient stratification and targeted interventions. |
Daniel J. Schad; Michael A. Rapp; Maria Garbusow; Stephan Nebe; Miriam Sebold; Elisabeth Obst; Christian Sommer; Lorenz Deserno; Milena Rabovsky; Eva Friedel; Nina Romanczuk-Seiferth; Hans Ulrich Wittchen; Ulrich S. Zimmermann; Henrik Walter; Philipp Sterzer; Michael N. Smolka; Florian Schlagenhauf; Andreas Heinz; Peter Dayan; Quentin J. M. M. Huys Dissociating neural learning signals in human sign- and goal-trackers Journal Article In: Nature Human Behaviour, vol. 4, no. 2, pp. 201–214, 2020. @article{Schad2020,Individuals differ in how they learn from experience. In Pavlovian conditioning models, where cues predict reinforcer delivery at a different goal location, some animals—called sign-trackers—come to approach the cue, whereas others, called goal-trackers, approach the goal. In sign-trackers, model-free phasic dopaminergic reward-prediction errors underlie learning, which renders stimuli ‘wanted'. Goal-trackers do not rely on dopamine for learning and are thought to use model-based learning. We demonstrate this double dissociation in 129 male humans using eye-tracking, pupillometry and functional magnetic resonance imaging informed by computational models of sign- and goal-tracking. We show that sign-trackers exhibit a neural reward prediction error signal that is not detectable in goal-trackers. Model-free value only guides gaze and pupil dilation in sign-trackers. Goal-trackers instead exhibit a stronger model-based neural state prediction error signal. This model-based construct determines gaze and pupil dilation more in goal-trackers. |
Jamie Reilly; Bonnie Zuckerman; Alexandra Kelly; Maurice Flurie; Sagar Rao Neuromodulation of cursing in American English: A combined tDCS and pupillometry study Journal Article In: Brain and Language, vol. 206, pp. 1–8, 2020. @article{Reilly2020,Many neurological disorders are associated with excessive and/or uncontrolled cursing. The right prefrontal cortex has long been implicated in a diverse range of cognitive processes that underlie the propensity for cursing, including non-propositional language representation, emotion regulation, theory of mind, and affective arousal. Neurogenic cursing often poses significant negative social consequences, and there is no known behavioral intervention for this communicative disorder. We examined whether right vs. left lateralized prefrontal neurostimultion via tDCS could modulate taboo word production in neurotypical adults. We employed a pre/post design with a bilateral frontal electrode montage. Half the participants received left anodal and right cathodal stimulation; the remainder received the opposite polarity stimulation at the same anatomical loci. We employed physiological (pupillometry) and behavioral (reaction time) dependent measures as participants read aloud taboo and non-taboo words. Pupillary responses demonstrated a crossover reaction, suggestive of modulation of phasic arousal during cursing. Participants in the right anodal condition showed elevated pupil responses for taboo words post stimulation. In contrast, participants in the right cathodal condition showed relative dampening of pupil responses for taboo words post stimulation. We observed no effects of stimulation on response times. We interpret these findings as supporting modulation of right hemisphere affective arousal that disproportionately impacts taboo word processing. We discuss alternate accounts of the data and future applications to neurological disorders. |
Antonella Pomè; Paola Binda; Guido Marco Cicchini; David C. Burr Pupillometry correlates of visual priming, and their dependency on autistic traits Journal Article In: Journal of Vision, vol. 20, no. 3, pp. 1–12, 2020. @article{Pome2020,In paradigms of visual search where the search feature (say color) can change from trial to trials, responses are faster for trials where the search color is repeated than when it changes. This is a clear example of "priming" of attention. Here we test whether the priming effects can be revealed by pupillometry, and also whether they are related to autistic-like personality traits, as measured by the Autism-Spectrum Quotient (AQ). We repeated Maljkovic and Nakayama's (1994) classic priming experiment, asking subjects to identify rapidly the shape of a singleton target defined by color. As expected, reaction times were faster when target color repeated, and the effect accumulated over several trials; but the magnitude of the effect did not correlate with AQ. Reaction times were also faster when target position was repeated, again independent of AQ. Presentation of stimuli caused the pupil to dilate, and the magnitude of dilation was greater for switched than repeated trials. This effect did not accumulate over trials, and did not correlate with the reaction times difference, suggesting that the two indexes measure independent aspects of the priming phenomenon. Importantly, the amplitude of pupil modulation correlated negatively with AQ, and was significant only for those participants with low AQ. The results confirm that pupillometry can track perceptual and attentional processes, and furnish useful information unobtainable from standard psychophysics, including interesting dependencies on personality traits. |
Dorothee Pöhlchen; Laura Leuchs; Florian P. Binder; Borbala Blaskovich; Taechawidd Nantawisarakul; Pavlos Topalidis; Tanja M. Brückl; Seth D. Norrholm; Tanja Jovanovic; Victor I. Spoormaker; Elisabeth B. Binder; Michael Czisch; Angelika Erhardt; Norma C. Grandi; Sanja Ilic-Cocic; Susanne Lucae; Philipp Sämann; Alina Tontsch No robust differences in fear conditioning between patients with fear-related disorders and healthy controls Journal Article In: Behaviour Research and Therapy, vol. 129, pp. 1–10, 2020. @article{Poehlchen2020,Fear conditioning and extinction serve as a dominant model for the development and maintenance of pathological anxiety, particularly for phasic fear to specific stimuli or situations. The validity of this model would be supported by differences in the physiological or subjective fear response between patients with fear-related disorders and healthy controls, whereas the model's validity would be questioned by a lack of such differences. We derived pupillometry, skin conductance response and startle electromyography as well as unconditioned stimulus expectancy in a two-day fear acquisition, immediate extinction and recall task and compared an unmedicated group of patients (n = 73) with phobias or panic disorder and a group of patients with posttraumatic stress disorder (PTSD |
Nick B. Pandža; Ian Phillips; Valerie P. Karuzis; Polly O'Rourke; Stefanie E. Kuchinsky Neurostimulation and pupillometry: New directions for learning and research in applied linguistics Journal Article In: Annual Review of Applied Linguistics, vol. 40, pp. 56–77, 2020. @article{Pandza2020,This paper begins by discussing new trends in the use of neurostimulation techniques in cognitive science and learning research, as well as the nascent research on their application in second language learning. To illustrate this, an experiment designed to investigate the impact of transcutaneous vagus nerve stimulation (tVNS), which is delivered via earbuds, on how learners process and learn Mandarin tones is reported. Pupillometry, which is an index of cognitive effort, is explained and illustrated as one way to assess the impact of tVNS. Participants in the study were native English speakers, naïve to tone languages, pseudorandomly assigned to active or control conditions, while balancing for nonlinguistic pitch ability and musical experience. Their performance after tVNS was assessed using a range of more traditional language outcome measures, including accuracy and reaction times from lexical recognition and recall tasks and was triangulated with pupillometry during word-learning to help understand the mechanism through which tVNS operates. Findings are discussed in light of the literatures on lexical tone learning, cognitive effort, and neurostimulation, including specific benefits for learners of tone languages. Recommendations are made for future work on the increasingly popular area of neurostimulation for the field of applied linguistics in the 40th anniversary issue of ARAL. |
Satoshi Nakakoga; Hiroshi Higashi; Junya Muramatsu; Shigeki Nakauchi; Tetsuto Minami Asymmetrical characteristics of emotional responses to pictures and sounds: Evidence from pupillometry Journal Article In: PLoS ONE, vol. 15, no. 4, pp. e0230775, 2020. @article{Nakakoga2020,In daily life, our emotions are often elicited by a multimodal environment, mainly visual and auditory stimuli. Therefore, it is crucial to investigate the symmetrical characteristics of emotional responses to pictures and sounds. In this study, we aimed to elucidate the relationship of attentional states to emotional unimodal stimuli (pictures or sounds) and emotional responses by measuring the pupil diameter, which reflects the emotional arousal associated with increased sympathetic activity. Our hypothesis was that the emotional responses to both the image and sound stimuli are symmetrical: emotion might be suppressed when attentional resources are allocated to another stimulus of the same modality as the emotional stimulus-such as a dot presented at the same time as an emotional image, and a beep sound presented at the same time as an emotional sound. In our two experiments, data for 24 participants were analyzed for a pupillary response. In experiment 1, we investigated the relationship of the attentional state with emotional visual stimuli (International Affective Picture System) and emotional responses by using pupillometry. We set four task conditions to modulate the attentional state (emotional task, no task, visual detection task, and auditory detection task). We observed that the velocity of pupillary dilation was faster during the presentation of emotionally arousing pictures compared to that of neutral ones, regardless of the valence of the pictures. Importantly, this effect was not dependent on the task condition. In experiment 2, we investigated the relationship of the attentional state with emotional auditory sounds (International Affective Digitized Sounds) and emotional responses. We observed a trend towards a significant interaction between the stimulus and the task conditions with regard to the velocity of pupillary dilation. In the emotional and auditory detection tasks, the velocity of pupillary dilation was faster with positive and neutral sounds than negative sounds. However, there were no significant differences between the no task and visual detection task conditions. Taken together, the current data reveal that different pupillary responses were elicited to emotional visual and auditory stimuli, at least in the point that there is no attentional effect to emotional responses to visual stimuli, despite both experiments being sufficiently controlled to be of symmetrical experimental design. |
Drew J. McLaughlin; Kristin J. Van Engen Task-evoked pupil response for accurately recognized accented speech Journal Article In: The Journal of the Acoustical Society of America, vol. 147, no. 2, pp. EL151–EL156, 2020. @article{McLaughlin2020,Unfamiliar second-language (L2) accents present a common challenge to speech understanding. However, the extent to which accurately recognized unfamiliar L2-accented speech imposes a greater cognitive load than native speech remains unclear. The current study used pupillometry to assess cognitive load for native English listeners during the per- ception of intelligible Mandarin Chinese-accented English and American-accented English. Results showed greater pupil response (indicating greater cognitive load) for the unfamiliar L2-accented speech. These findings indicate that the mismatches between unfamiliar L2- accented speech and native listeners' linguistic representations impose greater cognitive load even when recognition accuracy is at ceiling. |
Kevin P. Madore; Anna M. Khazenzon; Cameron W. Backes; Jiefeng Jiang; Melina R. Uncapher; Anthony M. Norcia; Anthony D. Wagner Memory failure predicted by attention lapsing and media multitasking Journal Article In: Nature, vol. 587, no. 7832, pp. 87–91, 2020. @article{Madore2020,With the explosion of digital media and technologies, scholars, educators and the public have become increasingly vocal about the role that an ‘attention economy' has in our lives1. The rise of the current digital culture coincides with longstanding scientific questions about why humans sometimes remember and sometimes forget, and why some individuals remember better than others2–6. Here we examine whether spontaneous attention lapses—in the moment7–12, across individuals13–15 and as a function of everyday media multitasking16–19—negatively correlate with remembering. Electroencephalography and pupillometry measures of attention20,21 were recorded as eighty young adults (mean age, 21.7 years) performed a goal-directed episodic encoding and retrieval task22. Trait-level sustained attention was further quantified using task-based23 and questionnaire measures24,25. Using trial-to-trial retrieval data, we show that tonic lapses in attention in the moment before remembering, assayed by posterior alpha power and pupil diameter, were correlated with reductions in neural signals of goal coding and memory, along with behavioural forgetting. Independent measures of trait-level attention lapsing mediated the relationship between neural assays of lapsing and memory performance, and between media multitasking and memory. Attention lapses partially account for why we remember or forget in the moment, and why some individuals remember better than others. Heavier media multitasking is associated with a propensity to have attention lapses and forget. |
Russell A. Cohen Hoffing; Nina Lauharatanahirun; Daniel E. Forster; Javier O. Garcia; Jean M. Vettel; Steven M. Thurman Dissociable mappings of tonic and phasic pupillary features onto cognitive processes involved in mental arithmetic Journal Article In: PLoS ONE, vol. 15, no. 3, pp. e0230517, 2020. @article{Hoffing2020,Pupil size modulations have been used for decades as a window into the mind, and several pupillary features have been implicated in a variety of cognitive processes. Thus, a general challenge facing the field of pupillometry has been understanding which pupil features should be most relevant for explaining behavior in a given task domain. In the present study, a longitudinal design was employed where participants completed 8 biweekly sessions of a classic mental arithmetic task for the purposes of teasing apart the relationships between tonic/phasic pupil features (baseline, peak amplitude, peak latency) and two task-related cognitive processes including mental processing load (indexed by math question difficulty) and decision making (indexed by response times). We used multi-level modeling to account for individual variation while identifying pupil-to-behavior relationships at the single-trial and between-session levels. We show a dissociation between phasic and tonic features with peak amplitude and latency (but not baseline) driven by ongoing task-related processing, whereas baseline was driven by state-level effects that changed over a longer time period (i.e. weeks). Finally, we report a dissociation between peak amplitude and latency whereby amplitude reflected surprise and processing load, and latency reflected decision making times. |
Eugen Fischer; Paul E. Engelhardt Lingering stereotypes: Salience bias in philosophical argument Journal Article In: Mind and Language, vol. 35, no. 4, pp. 415–439, 2020. @article{Fischer2020,Many philosophical thought experiments and arguments involve unusual cases. We present empirical reasons to doubt the reliability of intuitive judgments and conclusions about such cases. Inferences and intuitions prompted by verbal case descriptions are influenced by routine comprehension processes which invoke stereotypes. We build on psycholinguistic findings to determine conditions under which the stereotype associated with the most salient sense of a word predictably supports inappropriate inferences from descriptions of unusual (stereotype-divergent) cases. We conduct an experiment that combines plausibility ratings with pupillometry to document this “salience bias.” We find that under certain conditions, competent speakers automatically make stereotypical inferences they know to be inappropriate. |
Mojgan Farahani; Vijay Parsa; Björn Herrmann; Mason Kadem; Ingrid Johnsrude; Philip C. Doyle An auditory-perceptual and pupillometric study of vocal strain and listening effort in adductor spasmodic dysphonia Journal Article In: Applied Sciences, vol. 10, no. 17, pp. 5907, 2020. @article{Farahani2020,This study evaluated ratings of vocal strain and perceived listening effort by normal hearing participants while listening to speech samples produced by talkers with adductor spasmodic dysphonia (AdSD). In addition, objective listening effort was measured through concurrent pupillometry to determine whether listening to disordered voices changed arousal as a result of emotional state or cognitive load. Recordings of the second sentence of the "Rainbow Passage" produced by talkers with varying degrees of AdSD served as speech stimuli. Twenty naïve young adult listeners perceptually evaluated these stimuli on the dimensions of vocal strain and listening effort using two separate visual analogue scales. While making the auditory-perceptual judgments, listeners' pupil characteristics were objectively measured in synchrony with the presentation of each voice stimulus. Data analyses revealed moderate-to-high inter- and intra-rater reliability. A significant positive correlation was found between the ratings of vocal strain and listening effort. In addition, listeners displayed greater peak pupil dilation (PPD) when listening to more strained and effortful voice samples. Findings from this study suggest that when combined with an auditory-perceptual task, non-volitional physiologic changes in pupil response may serve as an indicator of listening and cognitive effort or arousal. |
Camilla E. J. Elphick; Graham E. Pike; Graham J. Hole You can believe your eyes: Measuring implicit recognition in a lineup with pupillometry Journal Article In: Psychology, Crime and Law, vol. 26, no. 1, pp. 67–92, 2020. @article{Elphick2020,As pupil size is affected by cognitive processes, we investigated whether it could serve as an independent indicator of target recognition in lineups. Participants saw a simulated crime video, followed by two viewings of either a target-present or target-absent video lineup while pupil size was measured with an eye-tracker. Participants who made correct identifications showed significantly larger pupil sizes when viewing the target compared with distractors. Some participants were uncertain about their choice of face from the lineup, but nevertheless showed pupillary changes when viewing the target, suggesting covert recognition of the target face had occurred. The results suggest that pupillometry might be a useful aid in assessing the accuracy of an eyewitness' identification. |
Daniel S. Drew; Kinan Muhammed; Fahd Baig; Mark Kelly; Youssuf Saleh; Nagaraja Sarangmat; David Okai; Michele Hu; Sanjay Manohar; Masud Husain Dopamine and reward hypersensitivity in Parkinson's disease with impulse control disorder Journal Article In: Brain, vol. 143, no. 8, pp. 2502–2518, 2020. @article{Drew2020,Impulse control disorders in Parkinson's disease are common neuropsychiatric complications associated with dopamine replacement therapy. Some patients treated with dopamine agonists develop pathological behaviours, such as gambling, compulsive eating, shopping, or disinhibited sexual behaviours, which can have a severe impact on their lives and that of their families. In this study we investigated whether hypersensitivity to reward might contribute to these pathological behaviours and how this is influenced by dopaminergic medication. We asked participants to shift their gaze to a visual target as quickly as possible, in order to obtain reward. Critically, the reward incentive on offer varied over trials. Motivational effects were indexed by pupillometry and saccadic velocity, and patients were tested ON and OFF dopaminergic medication, allowing us to measure the effect of dopaminergic medication changes on reward sensitivity. Twenty-three Parkinson's disease patients with a history of impulse control disorders were compared to 26 patients without such behaviours, and 31 elderly healthy controls. Intriguingly, behavioural apathy was reported alongside impulsivity in the majority of patients with impulse control disorders. Individuals with impulse control disorders also exhibited heightened sensitivity to exogenous monetary rewards cues both ON and OFF (overnight withdrawal) dopamine medication, as indexed by pupillary dilation in anticipation of reward. Being OFF dopaminergic medication overnight did not modulate pupillary reward sensitivity in impulse control disorder patients, whereas in control patients reward sensitivity was significantly reduced when OFF dopamine. These effects were independent of cognitive impairment or total levodopa equivalent dose. Although dopamine agonist dose did modulate pupillary responses to reward, the pattern of results was replicated even when patients with impulse control disorders on dopamine agonists were excluded from the analysis. The findings suggest that hypersensitivity to rewards might be a contributing factor to the development of impulse control disorders in Parkinson's disease. However, there was no difference in reward sensitivity between patient groups when ON dopamine medication, suggesting that impulse control disorders may not emerge simply because of a direct effect of dopaminergic drug level on reward sensitivity. The pupillary reward sensitivity measure described here provides a means to differentiate, using a physiological measure, Parkinson's disease patients with impulse control disorder from those who do not experience such symptoms. Moreover, follow-up of control patients indicated that increased pupillary modulation by reward can be predictive of the risk of future emergence of impulse control disorders and may thereby provide the potential for early identification of patients who are more likely to develop these symptoms. |
Violet A. Brown; Drew J. McLaughlin; Julia F. Strand; Kristin J. Van Engen Rapid adaptation to fully intelligible nonnative-accented speech reduces listening effort Journal Article In: Quarterly Journal of Experimental Psychology, vol. 73, no. 9, pp. 1431–1443, 2020. @article{Brown2020b,In noisy settings or when listening to an unfamiliar talker or accent, it can be difficult to understand spoken language. This difficulty typically results in reductions in speech intelligibility, but may also increase the effort necessary to process the speech even when intelligibility is unaffected. In this study, we used a dual-task paradigm and pupillometry to assess the cognitive costs associated with processing fully intelligible accented speech, predicting that rapid perceptual adaptation to an accent would result in decreased listening effort over time. The behavioural and physiological paradigms provided converging evidence that listeners expend greater effort when processing nonnative- relative to native-accented speech, and both experiments also revealed an overall reduction in listening effort over the course of the experiment. Only the pupillometry experiment, however, revealed greater adaptation to nonnative- relative to native-accented speech. An exploratory analysis of the dual-task data that attempted to minimise practice effects revealed weak evidence for greater adaptation to the nonnative accent. These results suggest that even when speech is fully intelligible, resolving deviations between the acoustic input and stored lexical representations incurs a processing cost, and adaptation may attenuate this cost. |
Giulia Borghini; Valerie Hazan Effects of acoustic and semantic cues on listening effort during native and non-native speech perception Journal Article In: The Journal of the Acoustical Society of America, vol. 147, no. 6, pp. 3783–3794, 2020. @article{Borghini2020,Relative to native listeners, non-native listeners who are immersed in a second language environment experience increased listening effort and reduced ability to successfully perform an additional task while listening. Previous research demonstrated that listeners can exploit a variety of intelligibility-enhancing cues to cope with adverse listening conditions. However, little is known about the implications of those speech perception strategies for listening effort. The current research aims to investigate by means of pupillometry how listening effort is modulated in native and non-native listeners by the availability of semantic context and acoustic enhancements during the comprehension of spoken sentences. For this purpose, semantic plausibility and speaking style were manipulated both separately and in combination during a speech perception task in noise. The signal to noise ratio was individually adjusted for each participant in order to target 50% intelligibility level. Behavioural results indicated that native and non-native listeners were equally able to fruitfully exploit both semantic and acoustic cues to aid their comprehension. Pupil data indicated that listening effort was reduced for both groups of listeners when acoustic enhancements were available, while the presence of a plausible semantic context did not lead to a reduction in listening effort. |
Naila Ayala; Matthew Heath Executive dysfunction after a sport-related concussion is independent of task-based symptom burden Journal Article In: Journal of Neurotrauma, vol. 37, pp. 2558–2568, 2020. @article{Ayala2020,A sport-related concussion (SRC) results in short- and long-term deficits in oculomotor control; however, it is unclear whether this change reflects executive dysfunction and/or a performance decrement due to an increase in task-based symptom burden. Here, individuals with a SRC - and age- and sex-matched controls - completed an antisaccade task (i.e., saccade mirror-symmetrical to a target) during the early (initial assessment: <=12 days) and later (follow-up assessment: < 30 days) stages of recovery. Antisaccades were used because they require top-down executive control and exhibit performance decrements following a SRC. Reaction time (RT) and directional errors were included with pupillometry because pupil size in the antisaccade task has been shown to provide a neural proxy for executive control. In addition, the Sport-Concussion Assessment Tool (SCAT-5) symptom checklist was completed prior to and after each oculomotor assessment to identify a possible task-based increase in symptomology. The SRC group yielded longer initial assessment RTs, more directional errors and larger task-evoked pupil dilations (TEPD) than the control group. At the follow-up assessment, RTs for the SRC and control group did not reliably differ; however, the former demonstrated more directional errors and larger TEPDs. SCAT-5 symptom severity scores did not vary from the pre- to post-oculomotor evaluation for either initial or follow-up assessments. Accordingly, a SRC imparts a persistent executive dysfunction to oculomotor planning independent of a task-based increase in symptom burden. These findings evince that antisaccades serve as an effective tool to identify subtle executive deficits during the early and later stages of SRC recovery. |
2019 |
Felicia Zhang; Sagi Jaffe-Dax; Robert C. Wilson; Lauren L. Emberson Prediction in infants and adults: A pupillometry study Journal Article In: Developmental Science, vol. 22, no. 4, pp. 1–9, 2019. @article{Zhang2019h,Adults use both bottom-up sensory inputs and top-down signals to generate predictions about future sensory inputs. Infants have also been shown to make predictions with simple stimuli and recent work has suggested top-down processing is available early in infancy. However, it is unknown whether this indicates that top-down prediction is an ability that is continuous across the lifespan or whether an infant's ability to predict is different from an adult's, qualitatively or quantitatively. We employed pupillometry to provide a direct comparison of prediction abilities across these disparate age groups. Pupil dilation response (PDR) was measured in 6-month olds and adults as they completed an identical implicit learning task designed to help learn associations between sounds and pictures. We found significantly larger PDR for visual omission trials (i.e. trials that violated participants' predictions without the presentation of new stimuli to control for bottom-up signals) compared to visual present trials (i.e. trials that confirmed participants' predictions) in both age groups. Furthermore, a computational learning model that is closely linked to prediction error (Rescorla-Wagner model) demonstrated similar learning trajectories suggesting a continuity of predictive capacity and learning across the two age groups. |
Peter Vincent; Thomas Parr; David Benrimoh; Karl J. Friston With an eye on uncertainty: Modelling pupillary responses to environmental volatility Journal Article In: PLOS Computational Biology, vol. 15, no. 7, pp. e1007126, 2019. @article{Vincent2019,Living creatures must accurately infer the nature of their environments. They do this despite being confronted by stochastic and context sensitive contingencies—and so must constantly update their beliefs regarding their uncertainty about what might come next. In this work, we examine how we deal with uncertainty that evolves over time. This prospective uncertainty (or imprecision) is referred to as volatility and has previously been linked to noradrenergic signals that originate in the locus coeruleus. Using pupillary dilatation as a measure of central noradrenergic signalling, we tested the hypothesis that changes in pupil diameter reflect inferences humans make about environmental volatility. To do so, we collected pupillometry data from participants presented with a stream of numbers. We generated these numbers from a process with varying degrees of volatility. By measuring pupillary dilatation in response to these stimuli—and simulating the inferences made by an ideal Bayesian observer of the same stimuli—we demonstrate that humans update their beliefs about environmental contingencies in a Bayes optimal way. We show this by comparing general linear (convolution) models that formalised competing hypotheses about the causes of pupillary changes. We found greater evidence for models that included Bayes optimal estimates of volatility than those without. We additionally explore the interaction between different causes of pupil dilation and suggest a quantitative approach to characterising a person's prior beliefs about volatility. |
Megan H. Papesh; Juan D. Guevara Pinto Spotting rare items makes the brain “blink” harder: Evidence from pupillometry Journal Article In: Attention, Perception, & Psychophysics, vol. 81, no. 8, pp. 2635–2647, 2019. @article{Papesh2019,In many visual search tasks (e.g., cancer screening, airport baggage inspections), the most serious search targets occur infrequently. As an ironic side effect, when observers finally encounter important objects (e.g., a weapon in baggage), they often fail to notice them, a phenomenon known as the low-prevalence effect (LPE). Although many studies have investigated LPE search errors, we investigated the attentional consequences of successful rare target detection. Using an attentional blink paradigm, we manipulated how often observers encountered the first serial target (T1), then measured its effects on their ability to detect a following target (T2). Across two experiments, we show that the LPE is more than just an inflated miss rate: When observers successfully detected rare targets, they were less likely to spot subsequent targets. Using pupillometry to index locus-coeruleus (LC) mediated attentional engagement, Experiment 2 confirmed that an LC refractory period mediates the attentional blink (`Nieuwenhuis, Gilzenrat, Holmes, & Cohen, 2005, Journal of Experimental Psychology: General, 134[3], 291–307), and that these effects emerge relatively quickly following T1 onset. Moreover, in both behavioral and pupil analyses, we found that detecting low-prevalence targets exacerbates the LC refractory period. Consequences for theories of the LPE are discussed. |
Sijia Zhao; Gabriela Bury; Alice Milne; Maria Chait Pupillometry as an objective measure of sustained attention in young and older listeners Journal Article In: Trends in Hearing, vol. 23, pp. 1–21, 2019. @article{Zhao2019a,The ability to sustain attention on a task-relevant sound-source whilst avoiding distraction from other concurrent sounds is fundamental to listening in crowded environments. To isolate this aspect of hearing we designed a paradigm that continuously measured behavioural and pupillometry responses during 25-second-long trials in young (18-35 yo) and older (63-79 yo) participants. The auditory stimuli consisted of a number (1, 2 or 3) of concurrent, spectrally distinct tone streams. On each trial, participants detected brief silent gaps in one of the streams whilst resisting distraction from the others. Behavioural performance demonstrated increasing difficulty with time-on-task and with number/proximity of distractor streams. In young listeners (N=20), pupillometry revealed that pupil diameter (on the group and individual level) was dynamically modulated by instantaneous task difficulty such that periods where behavioural performance revealed a strain on sustained attention, were also accompanied by increased pupil diameter. Only trials on which participants performed successfully were included in the pupillometry analysis. Therefore, the observed effects reflect consequences of task demands as opposed to failure to attend.In line with existing reports, we observed global changes to pupil dynamics in the older group, including decreased pupil diameter, a limited dilation range, and reduced temporal variability. However, despite these changes, the older group showed similar effects of attentive tracking to those observed in the younger listeners. Overall, our results demonstrate that pupillometry can be a reliable and time-sensitive measure of the effort associated with attentive tracking over long durations in both young and (with some caveats) older listeners. |
Felicia Zhang; Lauren L. Emberson Opposing timing constraints severely limit the use of pupillometry to investigate visual statistical learning Journal Article In: Frontiers in Psychology, vol. 10, pp. 1792, 2019. @article{Zhang2019c,Majority of visual statistical learning (VSL) research uses only offline measures, collected after the familiarization phase (i.e. learning) has occurred. Offline measures have revealed a lot about the extent of statistical learning (SL) but less is known about the learning mechanisms that support VSL. Studies have shown that prediction can be a potential learning mechanism for VSL, but it is difficult to examine the role of prediction in VSL using offline measures alone. Pupil diameter is a promising online measure to index prediction in VSL because it can be collected during learning, requires no overt action or task and can be used in a wide-range of populations (e.g., infants and adults). Furthermore, pupil diameter has already been used to investigate processes that are part of prediction such as prediction error and updating. While the properties of pupil diameter have the potentially to powerfully expand studies in VSL, through a series of three experiments, we find that the two are not compatible with each other. Our results revealed that pupil diameter, used to index prediction, is not related to offline measures of learning. We also found that pupil differences that appear to be a result of prediction, are actually a result of where we chose to baseline instead. Ultimately, we conclude that the fast-paced nature of VSL paradigms make it incompatible with the slow nature of pupil change. Therefore, our findings suggest pupillometry should not be used to investigate learning mechanisms in fast-paced VSL tasks. |
Chad C. Williams; Mitchel Kappen; Cameron D. Hassall; Bruce Wright; Olave E. Krigolson Thinking theta and alpha: Mechanisms of intuitive and analytical reasoning Journal Article In: NeuroImage, vol. 189, pp. 574–580, 2019. @article{Williams2019,Humans have a unique ability to engage in different modes of thinking. Intuitive thinking (coined System 1, see Kahneman, 2011) is fast, automatic, and effortless whereas analytical thinking (coined System 2) is slow, contemplative, and effortful. We extend seminal pupillometry research examining these modes of thinking by using electroencephalography (EEG) to decipher their respective underlying neural mechanisms. We demonstrate that System 1 thinking is characterized by an increase in parietal alpha EEG power reflecting autonomic access to long-term memory and a release of attentional resources whereas System 2 thinking is characterized by an increase in frontal theta EEG power indicative of the engagement of cognitive control and working memory processes. Consider our results in terms of an example - a child may need cognitive control and working memory when contemplating a mathematics problem yet an adullt can drive a car with little to no attention by drawing on easily accessed memories. Importantly, the unravelling of intuitive and analytical thinking mechanisms and their neural signatures will provide insight as to how different modes of thinking drive our everyday lives. |
Leonhard Waschke; Sarah Tune; Jonas Obleser Local cortical desynchronization and pupil-linked arousal differentially shape brain states for optimal sensory performance Journal Article In: eLife, vol. 8, pp. 1–27, 2019. @article{Waschke2019,Instantaneous brain states have consequences for our sensation, perception, and behaviour. Fluctuations in arousal and neural desynchronization likely pose perceptually relevant states. However, their relationship and their relative impact on perception is unclear. We here show that, at the single-trial level in humans, local desynchronization in sensory cortex (expressed as time-series entropy) versus pupil- linked arousal differentially impact perceptual processing. While we recorded electroencephalography (EEG) and pupillometry data, stimuli of a demanding auditory discrimination task were presented into states of high or low desynchronization of auditory cortex via a real-time closed-loop setup. Desynchronization and arousal distinctly influenced stimulus-evoked activity and shaped behaviour displaying an inverted u-shaped relationship: States of intermediate desynchronization elicited minimal response bias and fastest responses, while states of intermediate arousal gave rise to highest response sensitivity. Our results speak to a model in which independent states of local desynchronization and global arousal jointly optimise sensory processing and performance. |
Amy T. Walsh; David Carmel; Gina M. Grimshaw Reward elicits cognitive control over emotional distraction: Evidence from pupillometry Journal Article In: Cognitive, Affective, & Behavioral Neuroscience, vol. 19, no. 3, pp. 537–554, 2019. @article{Walsh2019,Attention is biased toward emotional stimuli, even when they are irrelevant to current goals. Motivation, elicited by performance-contingent reward, reduces behavioural emotional distraction. In emotionally neutral contexts, reward is thought to encourage use of a proactive cognitive control strategy, altering anticipatory attentional settings to more effectively suppress distractors. The current preregistered study investigates whether a similar proactive shift occurs even when distractors are highly arousing emotional images. We monitored pupil area, an online measure of both cognitive and emotional processing, to examine how reward influences the time course of control. Participants (n = 110) identified a target letter flanking an irrelevant central image. Images were meaningless scrambles on 75% of trials; on the remaining 25%, they were intact positive (erotic), negative (mutilation), or neutral images. Half the participants received financial rewards for fast and accurate performance, while the other half received no performance-contingent reward. Emotional distraction was greater than neutral distraction, and both were attenuated by reward. Consistent with behavioural findings, pupil dilation was greater following emotional than neutral distractors, and dilation to intact distractors (regardless of valence) was decreased by reward. Although reward did not enhance tonic pupil dilation (an index of sustained proactive control), exploratory analyses showed that reward altered the time course of control—eliciting a sharp, rapid, increase in dilation immediately preceding stimulus onset (reflecting dynamic use of anticipatory control), that extended until well after stimulus offset. These findings suggest that reward alters the time course of control by encouraging proactive preparation to rapidly disengage from emotional distractors. |
Anita E. Wagner; Leanne Nagels; Paolo Toffanin; Jane M. Opie; Deniz Başkent Individual variations in effort: Assessing pupillometry for the hearing impaired Journal Article In: Trends in Hearing, vol. 23, 2019. @article{Wagner2019,Assessing effort in speech comprehension for hearing-impaired (HI) listeners is important, as effortful processing of speech can limit their hearing rehabilitation. We examined the measure of pupil dilation in its capacity to accommodate the heterogeneity that is present within clinical populations by studying lexical access in users with sensorineural hearing loss, who perceive speech via cochlear implants (CIs). We compared the pupillary responses of 15 experienced CI users and 14 age-matched normal-hearing (NH) controls during auditory lexical decision. A growth curve analysis was applied to compare the responses between the groups. NH listeners showed a coherent pattern of pupil dilation that reflects the task demands of the experimental manipulation and a homogenous time course of dilation. CI listeners showed more variability in the morphology of pupil dilation curves, potentially reflecting variable sources of effort across individuals. In follow-up analyses, we examined how speech perception, a task that relies on multiple stages of perceptual analyses, poses multiple sources of increased effort for HI listeners, wherefore we might not be measuring the same source of effort for HI as for NH listeners. We argue that interindividual variability among HI listeners can be clinically meaningful in attesting not only the magnitude but also the locus of increased effort. The understanding of individual variations in effort requires experimental paradigms that (a) differentiate the task demands during speech comprehension, (b) capture pupil dilation in its time course per individual listeners, and (c) investigate the range of individual variability present within clinical and NH populations. |
Rozemarijn S. Kleef; Claudi L. H. Bockting; Evelien Valen; André Aleman; Jan Bernard C. Marsman; Marie José Tol In: BMC Psychiatry, vol. 19, pp. 1–11, 2019. @article{Kleef2019,Background: Major Depressive Disorder (MDD) is a psychiatric disorder with a highly recurrent character, making prevention of relapse an important clinical goal. Preventive Cognitive Therapy (PCT) has been proven effective in preventing relapse, though not for every patient. A better understanding of relapse vulnerability and working mechanisms of preventive treatment may inform effective personalized intervention strategies. Neurocognitive models of MDD suggest that abnormalities in prefrontal control over limbic emotion-processing areas during emotional processing and regulation are important in understanding relapse vulnerability. Whether changes in these neurocognitive abnormalities are induced by PCT and thus play an important role in mediating the risk for recurrent depression, is currently unclear. In the Neurocognitive Working Mechanisms of the Prevention of Relapse In Depression (NEWPRIDE) study, we aim to 1) study neurocognitive factors underpinning the vulnerability for relapse, 2) understand the neurocognitive working mechanisms of PCT, 3) predict longitudinal treatment effects based on pre-treatment neurocognitive characteristics, and 4) validate the pupil dilation response as a marker for prefrontal activity, reflecting emotion regulation capacity and therapy success. Methods: In this randomized controlled trial, 75 remitted recurrent MDD (rrMDD) patients will be included. Detailed clinical and cognitive measurements, fMRI scanning and pupillometry will be performed at baseline and three-month follow-up. In the interval, 50 rrMDD patients will be randomized to eight sessions of PCT and 25 rrMDD patients to a waiting list. At baseline, 25 healthy control participants will be additionally included to objectify cross-sectional residual neurocognitive abnormalities in rrMDD. After 18 months, clinical assessments of relapse status are performed to investigate which therapy induced changes predict relapse in the 50 patients allocated to PCT. Discussion: The present trial is the first to study the neurocognitive vulnerability factors underlying relapse and mediating relapse prevention, their value for predicting PCT success and whether pupil dilation acts as a valuable marker in this regard. Ultimately, a deeper understanding of relapse prevention could contribute to the development of better targeted preventive interventions. Trial registration: Trial registration: Netherlands Trial Register, August 18, 2015, trial number NL5219. |
Laura Leuchs; Max Schneider; Victor I. Spoormaker Measuring the conditioned response: A comparison of pupillometry, skin conductance, and startle electromyography Journal Article In: Psychophysiology, vol. 56, no. 1, pp. e13283, 2019. @article{Leuchs2019,In human fear conditioning studies, different physiological readouts can be used to track conditioned responding during fear learning. Commonly employed readouts such as skin conductance responses (SCR) or startle responses have in recent years been complemented by pupillary readouts, but to date it is unknown how pupillary readouts relate to other measures of the conditioned response. To examine differences and communalities among pupil responses, SCR, and startle responses, we simultaneously recorded pupil diameter, skin conductance, and startle electromyography in 47 healthy subjects during fear acquisition, extinction, and a recall test on 2 consecutive days. The different measures correlated only weakly, displaying most prominent differences in their response patterns during fear acquisition. Whereas SCR and startle responses habituated, pupillary measures did not. Instead, they increased in response to fear conditioned stimuli and most closely followed ratings of unconditioned stimulus (US) expectancy. Moreover, we observed that startle-induced pupil responses showed stimulus discrimination during fear acquisition, suggesting a fear potentiation of the auditory pupil reflex. We conclude that different physiological outcome measures of the conditioned response inform about different cognitive-affective processes during fear learning, with pupil responses being least affected by physiological habituation and most closely following US expectancy. |
Robert Jagiello; Ulrich Pomper; Makoto Yoneya; Sijia Zhao; Maria Chait Rapid brain sesponses to familiar vs. unfamiliar music-an EEG and pupillometry study Journal Article In: Scientific Reports, vol. 9, pp. 15570, 2019. @article{Jagiello2019,Human listeners exhibit marked sensitivity to familiar music, perhaps most readily revealed by popular "name that tune" games, in which listeners often succeed in recognizing a familiar song based on extremely brief presentation. In this work, we used electroencephalography (EEG) and pupillometry to reveal the temporal signatures of the brain processes that allow differentiation between a familiar, well liked, and unfamiliar piece of music. In contrast to previous work, which has quantified gradual changes in pupil diameter (the so-called "pupil dilation response"), here we focus on the occurrence of pupil dilation events. This approach is substantially more sensitive in the temporal domain and allowed us to tap early activity with the putative salience network. Participants (N = 10) passively listened to snippets (750 ms) of a familiar, personally relevant and, an acoustically matched, unfamiliar song, presented in random order. A group of control participants (N = 12), who were unfamiliar with all of the songs, was also tested. We reveal a rapid differentiation between snippets from familiar and unfamiliar songs: Pupil responses showed greater dilation rate to familiar music from 100-300 ms post-stimulus-onset, consistent with a faster activation of the autonomic salience network. Brain responses measured with EEG showed a later differentiation between familiar and unfamiliar music from 350 ms post onset. Remarkably, the cluster pattern identified in the EEG response is very similar to that commonly found in the classic old/new memory retrieval paradigms, suggesting that the recognition of brief, randomly presented, music snippets, draws on similar processes. |
Katja Herrmann; Andreas Sprenger; Leoni Baumung; Daniel Alvarez-Fischer; Alexander Münchau; Valerie Brandt Help or hurt? How attention modulates tics under different conditions Journal Article In: Cortex, vol. 120, pp. 471–482, 2019. @article{Herrmann2019,Tourette syndrome is a neuropsychiatric developmental disorder, characterized by tics that are often preceded by an increasingly uncomfortable urge to move. Tic frequency can increase when patients pay attention to their tics, if tics are not suppressed. This study investigates how attentions modulates urge intensity, tic frequency and arousal during free ticcing and tic suppression. Tic frequency (video recording), urge intensity (rating scale) and pupil width (pupillometry as a measure of arousal) were assessed in 23 patients with Tourette syndrome (mean age 33.48 ± 12.37; 14 male) during five attention conditions: 1) baseline, 2) watching own tics in a live video-feedback, 3) watching own tics in a previously recorded video, 4) thinking about situations that can trigger tics and 5) thinking about specific, non-tic related stimuli (distraction condition) during: a) free ticcing and b) tic suppression tic states. Urge intensity and tic frequency increased in the free ticcing condition when patients viewed their own tics live and when they thought about tic-triggering situations. In the tic suppression condition, tic frequency increased when patients watched a video of their tics, thought about their tics or were distracted. Pupil width increased significantly during the live feedback and the video condition compared to baseline in both tic states. Paying attention to own tics can be detrimental when tics are not suppressed. In contrast, paying attention to other stimuli appears detrimental when tics are suppressed, as would be the case during most current behavioural therapy techniques. However, results point to high emotional arousal and patients feeling uncomfortable when seeing themselves tic. The results also suggest that urge intensity is modulated by changes in attention in the same manner as tics and may drive change in tic frequency during free ticcing. |
Dorothea Hämmerer; Philipp Schwartenbeck; Maria Gallagher; Thomas Henry Benedict FitzGerald; Emrah Düzel; Raymond Joseph Dolan Older adults fail to form stable task representations during model-based reversal inference Journal Article In: Neurobiology of Aging, vol. 74, pp. 90–100, 2019. @article{Haemmerer2019,Older adults struggle in dealing with changeable and uncertain environments across several cognitive domains. This has been attributed to difficulties in forming adequate task representations that help navigate uncertain environments. Here, we investigate how, in older adults, inadequate task representations impact on model-based reversal learning. We combined computational modeling and pupillometry during a novel model-based reversal learning task, which allowed us to isolate the relevance of task representations at feedback evaluation. We find that older adults overestimate the changeability of task states and consequently are less able to converge on unequivocal task representations through learning. Pupillometric measures and behavioral data show that these unreliable task representations in older adults manifest as a reduced ability to focus on feedback that is relevant for updating task representations, and as a reduced metacognitive awareness in the accuracy of their actions. Instead, the data suggested older adults' choice behavior was more consistent with a guidance by uninformative feedback properties such as outcome valence. Our study highlights that an inability to form adequate task representations may be a crucial factor underlying older adults' impaired model-based inference. |
Steven M. Gillespie; Pia Rotshtein; Harriet Chapman; Emmie Brown; Anthony R. Beech; Ian J. Mitchell Pupil reactivity to emotional faces among convicted violent offenders: The role of psychopathic traits Journal Article In: Journal of Abnormal Psychology, vol. 128, no. 6, pp. 622–632, 2019. @article{Gillespie2019,Psychopathy is characteristically associated with impairments in recognizing others' facial expressions of emotion, and there is some evidence that these difficulties are specific to the callousness features of the disorder. However, it remains unclear whether these difficulties are accompanied by reductions in autonomic reactivity when viewing others' emotional expressions, and whether these impairments are particular to expressions showing another's distress or are more pervasive across different emotional expressions. In this study, 73 adult male prisoners with histories of serious sexual or violent offenses- who ranged across the psychopathy continuum-completed a facial emotion recognition task. For the first time in a convicted offender sample, we used pupillometry techniques to measure changes in the pupil dilation response, a measure of sympathetic autonomic arousal to affective stimuli. We found that the callousness features of psychopathy were related to impaired recognition of fearful faces. Strikingly, we also showed that increasing callousness was associated with a reduction in the pupil dilation response and that this was pervasive across different emotional expressions. Our results highlight a potential role of the locus coeruleus-noradrenaline system in the pathophysiology of psychopathy and demonstrate the potential of the pupillary response as a technique for understanding attention- emotion interactions in psychopathy. |
Abraham I. J. Gajardo; Samuel Madariaga; Pedro E. Maldonado Autonomic nervous system assessment by pupillary response as a potential biomarker for cardiovascular risk: A pilot study Journal Article In: Journal of Clinical Neuroscience, vol. 59, pp. 41–46, 2019. @article{Gajardo2019,Background: Cardiovascular risk (CVR) biomarkers are of increasing interest because of their potential utility in management of cardiovascular diseases. The activity of the autonomic nervous system (ANS) is known to be highly correlated with CVR and therefore, is a putative biomarker. Common ANS measurement tools have several technological limitations and high-variance signals. The pupillary responses (PR) is controlled by both components of the ANS, and recent advances in pupillometry are making this measurement, easy and reliable. Thus, PR assessment could become a useful clinical tool to measure the ANS modulation and its relation to CVR. Here, we aimed to evaluate differences in PR between low CVR and moderate/high CVR individuals. Methods: We performed a cross-sectional study. We recruited voluntaries with low CVR (group 1 |
Daniel L. Bowling; Pablo Graf Ancochea; Michael J. Hove; W. Tecumseh Fitch Pupillometry of groove: Evidence for noradrenergic arousal in the link between music and movement Journal Article In: Frontiers in Neuroscience, vol. 12, pp. 1039, 2019. @article{Bowling2019,The capacity to entrain motor action to rhythmic auditory stimulation is highly developed in humans and extremely limited in our closest relatives. Investigating the mechanisms underlying this capacity thus represents an opportunity to advance our understanding of human evolution. An important aspect of auditory-motor entrainment is that not all forms of rhythmic stimulation motivate movement to the same degree. This variation is captured by the concept of musical groove: high-groove music stimulates a strong desire for movement, whereas low-groove music does not. Here, we utilize this difference to investigate the neurophysiological basis of our capacity for auditory-motor entrainment. In a series of three experiments we examine pupillary responses to musical stimuli varying in groove. Our results show stronger pupil dilation in response to (1) high- vs. low-groove music, (2) high vs. low spectral content, and (3) syncopated vs. straight drum patterns. We additionally report evidence for consistent sex differences in music-induced pupillary responses, with males exhibiting larger differences between responses, but females exhibiting stronger responses overall. These results imply that the biological link between movement and auditory rhythms in our species is supported by the capacity of high-groove music to stimulate arousal in the central and peripheral nervous system, presumably via highly conserved noradrenergic mechanisms. |
Sara Alhanbali; Piers Dawes; Rebecca E. Millman; Kevin J. Munro Measures of listening effort are multidimensional Journal Article In: Ear & Hearing, vol. 40, no. 5, pp. 1084–1097, 2019. @article{Alhanbali2019,OBJECTIVES: Listening effort can be defined as the cognitive resources required to perform a listening task. The literature on listening effort is as confusing as it is voluminous: measures of listening effort rarely correlate with each other and sometimes result in contradictory findings. Here, we directly compared simultaneously recorded multimodal measures of listening effort. After establishing the reliability of the measures, we investigated validity by quantifying correlations between measures and then grouping-related measures through factor analysis. DESIGN: One hundred and sixteen participants with audiometric thresholds ranging from normal to severe hearing loss took part in the study (age range: 55 to 85 years old, 50.3% male). We simultaneously measured pupil size, electroencephalographic alpha power, skin conductance, and self-report listening effort. One self-report measure of fatigue was also included. The signal to noise ratio (SNR) was adjusted at 71% criterion performance using sequences of 3 digits. The main listening task involved correct recall of a random digit from a sequence of six presented at a SNR where performance was around 82 to 93%. Test-retest reliability of the measures was established by retesting 30 participants 7 days after the initial session. RESULTS: With the exception of skin conductance and the self-report measure of fatigue, interclass correlation coefficients (ICC) revealed good test-retest reliability (minimum ICC: 0.71). Weak or nonsignificant correlations were identified between measures. Factor analysis, using only the reliable measures, revealed four underlying dimensions: factor 1 included SNR, hearing level, baseline alpha power, and performance accuracy; factor 2 included pupillometry; factor 3 included alpha power (during speech presentation and during retention); factor 4 included self-reported listening effort and baseline alpha power. CONCLUSIONS: The good ICC suggests that poor test reliability is not the reason for the lack of correlation between measures. We have demonstrated that measures traditionally used as indicators of listening effort tap into multiple underlying dimensions. We therefore propose that there is no "gold standard" measure of listening effort and that different measures of listening effort should not be used interchangeably. When choosing method(s) to measure listening effort, the nature of the task and aspects of increased listening demands that are of interest should be taken into account. The findings of this study provide a framework for understanding and interpreting listening effort measures. |
2018 |
Matthew B. Winn; Ashley N. Moore In: Trends in Hearing, vol. 22, 2018. @article{Winn2018,Contextual cues can be used to improve speech recognition, especially for people with hearing impairment. However, previous work has suggested that when the auditory signal is degraded, context might be used more slowly than when the signal is clear. This potentially puts the hearing-impaired listener in a dilemma of continuing to process the last sentence when the next sentence has already begun. This study measured the time course of the benefit of context using pupillary responses to high- and low-context sentences that were followed by silence or various auditory distractors (babble noise, ignored digits, or attended digits). Participants were listeners with cochlear implants or normal hearing using a 12-channel noise vocoder. Context-related differences in pupil dilation were greater for normal hearing than for cochlear implant listeners, even when scaled for differences in pupil reactivity. The benefit of context was systematically reduced for both groups by the presence of the later-occurring sounds, including virtually complete negation when sentences were followed by another attended utterance. These results challenge how we interpret the benefit of context in experiments that present just one utterance at a time. If a listener uses context to ‘‘repair'' part of a sentence, and later-occurring auditory stimuli interfere with that repair process, the benefit of context might not survive outside the idealized laboratory or clinical environment. Elevated listening effort in hearing-impaired listeners might therefore result not just from poor auditory encoding but also inefficient use of context and prolonged processing of misperceived utterances competing with perception of incoming speech. |