EyeLink EEG / fNIRS / TMS Publications
All EyeLink EEG, fNIRS, and TMS research publications (with concurrent eye tracking) up until 2024 (with early 2025s) are listed below by year. You can search the publications using keywords such as P300, Gamma band, NIRS, etc. You can also search for individual author names. If we missed any EyeLink EEG, fNIRS, or TMS articles, please email us!
2025 |
Masataka Yano; Keiyu Niikuni; Ruri Shimura; Natsumi Funasaki; Masatoshi Koizumi Producing non-basic word orders in (in)felicitous contexts: Evidence from pupillometry and functional near-infrared spectroscopy (fNIRS) Journal Article In: Language, Cognition and Neuroscience, vol. 40, no. 1, pp. 1–22, 2025. @article{Yano2025, The present study examined why speakers of languages with flexible word orders are more likely to use syntactically complex non-basic word orders when they provide discourse-given information earlier in sentences. This may be because they are more efficient for speakers to produce (the Speaker Economy Hypothesis). Alternatively, speakers may produce them to help listeners understand sentences more efficiently (the Listener Economy Hypothesis), given that previous studies showed that the processing of non-basic word orders was facilitated when the felicitous context was provided (i.e. a displaced object refers to discourse-given information). We addressed this issue by conducting a picture-description experiment, in which participants uttered sentences with syntactically basic Subject-Object-Verb (SOV) or non-basic Object-Subject-Verb (OSV) in felicitous or infelicitous contexts while cognitive load was tracked using pupillometry and functional near-infrared spectroscopy. The results showed that the felicitous context facilitated the filler-gap dependency formation of OSVs in production, supporting the Speaker Economy Hypothesis. |
Jing Zhu; Yuanlong Li; Changlin Yang; Hanshu Cai; Xiaowei Li; Bin Hu Transformer-based fusion model for mild depression recognition with EEG and pupil area signals Journal Article In: Medical and Biological Engineering & Computing, pp. 1–17, 2025. @article{Zhu2025, Early detection and treatment are crucial for the prevention and treatment of depression; compared with major depression, current researches pay less attention to mild depression. Meanwhile, analysis of multimodal biosignals such as EEG, eye movement data, and magnetic resonance imaging provides reliable technical means for the quantitative analysis of depression. However, how to effectively capture relevant and complementary information between multimodal data so as to achieve efficient and accurate depression recognition remains a challenge. This paper proposes a novel Transformer-based fusion model using EEG and pupil area signals for mild depression recognition. We first introduce CSP into the Transformer to construct single-modal models of EEG and pupil data and then utilize attention bottleneck to construct a mid-fusion model to facilitate information exchange between the two modalities; this strategy enables the model to learn the most relevant and complementary information for each modality and only share the necessary information, which improves the model accuracy while reducing the computational cost. Experimental results show that the accuracy of the EEG and pupil area signals of single-modal models we constructed is 89.75% and 84.17%, the precision is 92.04% and 95.21%, the recall is 89.5% and 71%, the specificity is 90% and 97.33%, the F1 score is 89.41% and 78.44%, respectively, and the accuracy of mid-fusion model can reach 93.25%. Our study demonstrates that the Transformer model can learn the long-term time-dependent relationship between EEG and pupil area signals, providing an idea for designing a reliable multimodal fusion model for mild depression recognition based on EEG and pupil area signals. |
Hao Zhang; Yiqing Hu; Yang Li; Shuangyu Zhang; Xiao Li Li; Chenguang Zhao Simultaneous dataset of brain, eye and hand during visuomotor tasks Journal Article In: Scientific Data, vol. 12, no. 1, pp. 1–15, 2025. @article{Zhang2025a, Visuomotor integration is a complex skill set encompassing many fundamental abilities, such as visual search, attention monitoring, and motor control. To explore the dynamic interplay between visual inputs and motor outputs, it is necessary to simultaneously record multiple brain activities with high temporal and spatial resolution, as well as to record implicit and explicit behaviors. However, there is a lack of public datasets that provide simultaneous multiple modalities during a visual-motor task. Functional near-infrared spectroscopy and electroencephalography to record brain activity simultaneously facilitate more precise capture of the complex visuomotor of brain mechanisms. Additionally, by employing a combined eye movement and manual response, it is possible to fully evaluate the effects of visuomotor outputs from implicit and explicit dimensions. We recorded whole-brain EEG (34 electrodes) and fNIRS (44 channels) covering the frontal and parietal cortex along with eye movements, behavior sampling, and operant behavior. The dataset underwent rigorous synchronization, quality control to highlight the effectiveness of our experiments and to demonstrate the high quality of our multimodal data framework. |
Taishen Zeng; Longxia Lou; Zhifang Liu; Zhijun Zhang Age-related depreciation in predictive processing during Chinese reading: Insights from fixation-related potentials Journal Article In: Current Psychology, no. 2004, pp. 1–11, 2025. @article{Zeng2025a, To overcome methodological deficiencies in previous eye-tracking and event-related potentials (ERP) studies, the fixa- tion-related potential (FRP) approach was used to investigate how aging affects predictive processing in silent Chinese free-view reading. Forty older and 42 young adults participated in the experiment. All of them reported good reading abilities and none suffered from physical, mental, or cognitive diseases. The older participants were over 60 years of age (62.670 ± 3.018), and they did not differ from the younger group in the schooling years (11.43 vs. 12.10 |
Taishen Zeng; Longxia Lou; Zhi-Fang Liu; Chaoyang Chen; Zhijun Zhang Coregistration of eye movements and EEG reveals frequency effects of words and their constituent characters in natural silent Chinese reading Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–14, 2025. @article{Zeng2025, We conducted two experiments to examine the lexical and sub-lexical processing of Chinese two-character words in reading. We used a co-registration electroencephalogram (EEG) for the first fixation on target words. In Experiment 1, whole-word occurrence frequency and initial constituent character frequency were orthogonally manipulated, while in Experiment 2, whole-word occurrence frequency and end constituent character frequency were orthogonally manipulated. Results showed that word frequency facilitated eye-tracking measures, while initial and end character frequencies inhibited them. Classical word frequency effects on N170 and N400 in the posterior region and reversed word frequency effects over the anterior region were consistently observed in both experiments. Experiment 1 revealed an inhibiting effect of initial character frequency on anterior N170. In Experiment 2, interaction between end-character frequency and word frequency showed reliable effects on anterior N170 and N400. These results demonstrate both facilitating and inhibiting word frequency effects, along with inhibiting effects of character frequency and that word frequency moderates the inhibiting effects of end constituent character frequency during natural silent Chinese reading. |
Magdalena Gruner; Andreas Widmann; Stefan Wöhner; Erich Schröger; Jörg D. Jescheniak Semantic context effects in picture and sound naming: Evidence from event-related potentials and pupillometric data Journal Article In: Journal of cognitive neuroscience, vol. 37, no. 2, pp. 443–463, 2025. @article{Gruner2025, When a picture is repeatedly named in the context of semantically related pictures (homogeneous context), responses are slower than when the picture is repeatedly named in the context of unrelated pictures (heterogeneous context). This semantic interference effect in blocked-cyclic naming plays an important role in devising theories of word production. Wöhner, Mädebach, and Jescheniak [Wöhner, S., Mädebach, A., & Jescheniak, J. D. Naming pictures and sounds: Stimulus type affects semantic context effects. Journal of Experimental Psychology: Human Perception and Performance, 47, 716-730, 2021] have shown that the effect is substantially larger when participants name environmental sounds than when they name pictures. We investigated possible reasons for this difference, using EEG and pupillometry. The behavioral data replicated Wöhner and colleagues. ERPs were more positive in the homogeneous compared with the heterogeneous context over central electrode locations between 140-180 msec and 250-350 msec for picture naming and between 250 and 350 msec for sound naming, presumably reflecting semantic interference during semantic and lexical processing. The later component was of similar size for pictures and sounds. ERPs were more negative in the homogeneous compared with the heterogeneous context over frontal electrode locations between 400 and 600 msec only for sounds. The pupillometric data showed a stronger pupil dilation in the homogeneous compared with the heterogeneous context only for sounds. The amplitudes of the late ERP negativity and pupil dilation predicted naming latencies for sounds in the homogeneous context. The latency of the effects indicates that the difference in semantic interference between picture and sound naming arises at later, presumably postlexical processing stages closer to articulation. We suggest that the processing of the auditory stimuli interferes with phonological response preparation and self-monitoring, leading to enhanced semantic interference. |
Ivan Iotzov; Lucas C. Parra Effects of noise and reward on pupil size and electroencephalographic speech tracking in a word-detection task Journal Article In: European Journal of Neuroscience, vol. 61, pp. 1–12, 2025. @article{Iotzov2025, Speech is hard to understand when there is background noise. Speech intelligibility and listening effort both affect our ability to understand speech, but the relative contribution of these factors is hard to disentangle. Previous studies suggest that speech intelligibility could be assessed with EEG speech tracking and listening effort via pupil size. However, these measures may be confounded, because poor intelligibility may require a larger effort. To address this, we developed a novel word-detection paradigm that allows for a rapid behavioural assessment of speech processing. In this paradigm, words appear on the screen during continuous speech, similar to closed captioning. In two listening experiments with a total of 51 participants, we manipulated intelligibility by changing signal-to-noise ratios (SNRs) and modulated effort by varying monetary reward. Increasing SNR improved detection performance along with EEG speech tracking. Additionally, we find that pupil size increases with increased SNR. Surprisingly, when we modulated both reward and SNR, we found that reward modulated only pupil size, whereas SNR modulated only EEG speech tracking. We interpret this as the effects of arousal and listening effort on pupil size and of intelligibility on EEG speech tracking. The experimental paradigm |
Rossella Breveglieri; Riccardo Brandolani; Stefano Diomedi; Markus Lappe; Claudio Galletti; Patrizia Fattori Role of the medial posterior parietal cortex in orchestrating attention and reaching Journal Article In: The Journal of Neuroscience, vol. 45, no. 1, pp. 1–11, 2025. @article{Breveglieri2025, The interplay between attention, alertness, and motor planning is crucial for our manual interactions. To investigate the neural bases of this interaction and challenge the views that attention cannot be disentangled from motor planning, we instructed human volunteers of both sexes to plan and execute reaching movements while attending to the target, while attending elsewhere, or without constraining attention. We recorded reaction times to reach initiation and pupil diameter and interfered with the functions of the medial posterior parietal cortex (mPPC) with online repetitive transcranial magnetic stimulation to test the causal role of this cortical region in the interplay between spatial attention and reaching. We found that mPPC plays a key role in the spatial association of reach planning and covert attention. Moreover, we have found that alertness, measured by pupil size, is a good predictor of the promptness of reach initiation only if we plan a reach to attended targets, and mPPC is causally involved in this coupling. Different from previous understanding, we suggest that mPPC is neither involved in reach planning per se, nor in sustained covert attention in the absence of a reach plan, but it is specifically involved in attention functional to reaching. |
Cemre Baykan; Alexander C. Schütz Electroencephalographic responses to the number of objects in partially occluded and uncovered scenes Journal Article In: Journal of Nognitive neuroscience, vol. 37, no. 1, pp. 227–238, 2025. @article{Baykan2025, Perceptual completion is ubiquitous when estimating properties such as the shape, size, or number of objects in partially occluded scenes. Behavioral experiments showed that the number of hidden objects is underestimated in partially occluded scenes compared with an estimation based on the density of visible objects and the amount of occlusion. It is still unknown at which processing level this (under)estimation of the number of hidden objects occurs. We studied this question using a passive viewing task in which observers viewed a game board that was initially partially occluded and later was uncovered to reveal its hidden parts. We simultaneously measured the electroencephalographic responses to the partially occluded board presentation and its uncovering. We hypothesized that if the underestimation is a result of early sensory processing, it would be observed in the activities of P1 and N1, whereas if it is because of higher level processes such as expectancy, it would be reflected in P3 activities. Our data showed that P1 amplitude increased with numerosity in both occluded and uncovered states, indicating a link between P1 and simple stimulus features. The N1 amplitude was highest when both the initially visible and uncovered areas of the board were completely filled with game pieces, suggesting that the N1 component is sensitive to the overall Gestalt. Finally, we observed that P3 activity was reduced when the density of game pieces in the uncovered parts matched the initially visible parts, implying a relationship between the P3 component and expectation mismatch. Overall, our results suggest that inferences about the number of hidden items are reflected in high-level processing. |
Caleb Stone; Jason B. Mattingley; Dragan Rangelov Neural mechanisms of metacognitive improvement under speed pressure Journal Article In: Communications Biology, vol. 8, pp. 1–12, 2025. @article{Stone2025, The ability to accurately monitor the quality of one's choices, or metacognition, improves under speed pressure, possibly due to changes in post-decisional evidence processing. Here, we investigate the neural processes that regulate decision-making and metacognition under speed pressure using time- resolved analyses of brain activity recorded using electroencephalography. Participants performed a motion discrimination task under short and long response deadlines and provided a metacognitive rating following each response. Behaviourally, participants were faster, less accurate, and showed superior metacognition with short deadlines. These effects were accompanied by a larger centro- parietal positivity (CPP), a neural correlate of evidence accumulation. Crucially, post-decisional CPP amplitude was more strongly associated with participants' metacognitive ratings following errors under short relative to long responsedeadlines. Our results suggest that superior metacognition under speed pressure may stem from enhanced metacognitive readout of post-decisional evidence. |
Yi Hsuan Chang; Rachel Yep; Chin An Wang In: Psychophysiology, vol. 62, pp. 1–22, 2025. @article{Chang2025, Pupil size is a non-invasive index for autonomic arousal mediated by the locus coeruleus–norepinephrine (LC-NE) system. While pupil size and its derivative (velocity) are increasingly used as indicators of arousal, limited research has investigated the relationships between pupil size and other well-known autonomic responses. Here, we simultaneously recorded pupillometry, heart rate, skin conductance, pulse wave amplitude, and respiration signals during an emotional face–word Stroop task, in which task-evoked (phasic) pupil dilation correlates with LC-NE responsivity. We hypothesized that emotional conflict and valence would affect pupil and other autonomic responses, and trial-by-trial correlations between pupil and other autonomic responses would be observed during both tonic and phasic epochs. Larger pupil dilations, higher pupil size derivative, and lower heart rates were observed in the incongruent condition compared to the congruent condition. Additionally, following incongruent trials, the congruency effect was reduced, and arousal levels indexed by previous-trial pupil dilation were correlated with subsequent reaction times. Furthermore, linear mixed models revealed that larger pupil dilations correlated with higher heart rates, higher skin conductance responses, higher respiration amplitudes, and lower pulse wave amplitudes on a trial-by-trial basis. Similar effects were seen between positive and negative valence conditions. Moreover, tonic pupil size before stimulus presentation significantly correlated with all other tonic autonomic responses, whereas tonic pupil size derivative correlated with heart rates and skin conductance responses. These results demonstrate a trial-by-trial relationship between pupil dynamics and other autonomic responses, highlighting pupil size as an effective real-time index for autonomic arousal during emotional conflict and valence processing. |
Gabriela Cruz; María Melcón; Leonardo Sutandi; Matias M. Palva; Satu Palva; Gregor Thut Oscillatory brain activity in the canonical alpha-band conceals distinct mechanisms in attention Journal Article In: The Journal of Neuroscience, vol. 45, no. 1, pp. 1–17, 2025. @article{Cruz2025, Brain oscillations in the alpha-band (8-14 Hz) have been linked to specific processes in attention and perception. In particular, decreases in posterior alpha-amplitude are thought to reflect activation of perceptually relevant brain areas for target engagement, while alpha-amplitude increases have been associated with inhibition for distractor suppression. Traditionally, these alpha-changes have been viewed as two facets of the same process. However, recent evidence calls for revisiting this interpretation. Here, we recorded MEG/EEG in 32 participants (19 females) during covert visuospatial attention shifts (spatial cues) and two control conditions (neutral cue, no-attention cue), while tracking fixational eye movements. In disagreement with a single, perceptually relevant alpha-process, we found the typical alpha-modulations contra- and ipsilateral to the attention focus to be triple dissociated in their timing, topography, and spectral features: Ipsilateral alpha-increases occurred early, over occipital sensors, at a high alpha-frequency (10–14 Hz) and were expressed during spatial attention (alpha spatial cue > neutral cue). In contrast, contralateral alpha-decreases occurred later, over parietal sensors, at a lower alpha-frequency (7–10 Hz) and were associated with attention deployment in general (alpha spatial and neutral cue < no-attention cue). Additionally, the lateralized early alpha-increases but not alpha-decreases during spatial attention coincided in time with directionally biased microsaccades. Overall, this suggests that the attention-related early alpha-increases and late alpha-decreases reflect distinct, likely reflexive versus endogenously controlled attention mechanisms. We conclude that there is more than one perceptually relevant posterior alpha-oscillation, which need to be dissociated for a detailed account of their roles in perception and attention. |
S. Duschek; T. Rainer; P. Piwkowski; J. Vorwerk; L. Riml; U. Ettinger Neural correlates of proactive and reactive control investigated using a novel precued antisaccade paradigm Journal Article In: Psychophysiology, vol. 62, pp. 1–17, 2025. @article{Duschek2025, This ERP study investigated central nervous correlates of proactive and reactive control using a novel precued antisaccade paradigm. Proactive control refers to preparatory processes during anticipation of a behaviorally relevant event; reactive control is activated after such an event to ensure goal attainment. A 64-channel EEG was obtained in 35 subjects; video-based eye tracking was applied for ocular recording. In the task, a target (probe) appeared left or right of the fixation point 1800ms after a visual cue; subjects had to move their gaze to the probe (prosaccade) or its mirror image position (antisaccade). Probes were emotional face expressions; their frame colors instructed task requirements. The cue informed about antisaccade probability (70% vs. 30%) in a trial. High antisaccade probability was associated with larger CNV amplitude than low antisaccade probability. In trials with incongruence between expected and actual task requirements, the probe N2 and P3a amplitudes were larger than in congruent trials. The P3a was smaller for affective than neutral probes. Task accuracy and speed were lower in incongruent trials and varied according to affective probe valence. EEG source imaging suggested origin of the ERPs in the orbitofrontal cortex and superior frontal gyrus. The difference for the CNV indicates greater cortical activity during higher proactive control demands. The larger probe N2 and P3a in incongruent trials reflect greater resource allocation to conflict monitoring and conflict resolution, i.e., reactive control. The influence of probe valence on the P3a suggests reduction of processing capacity due to higher emotional arousal. |
Anna R. Knippenberg; Sabrina Yavari; Gregory P. Strauss Negative auditory hallucinations are associated with increased activation of the defensive motivational system in schizophrenia Journal Article In: Schizophrenia Research: Cognition, vol. 39, pp. 1–6, 2025. @article{Knippenberg2025, Auditory hallucinations (AH) are the most common symptom of psychosis. The voices people hear make comments that are benign or even encouraging, but most often voices are threatening and derogatory. Negative AH are often highly distressing and contribute to suicide risk and violent behavior. Biological mechanisms underlying the valence of voices (i.e., positive, negative, neutral) are not well delineated. In the current study, we examined whether AH voice valence was associated with increased activation of the Defensive Motivational System, as indexed by central and autonomic system response to unpleasant stimuli. Data were evaluated from two studies that used a common symptom rating instrument, the Psychotic Symptom Rating Scale (PSY-RATS), to measure AH valence. Participants included outpatients diagnosed with SZ. Tasks included: Study 1: Trier Social Stress Task while heart rate was recorded via electrocardiography (N = 27); Study 2: Passive Viewing Task while participants were exposed to pleasant, unpleasant, and neutral images from the International Affective Picture System (IAPS) library while eye movements, pupil dilation, and electroencephalography were recorded (N = 25). Results indicated that negative voice content was significantly associated with: 1) increased heart rate during an acute social stressor, 2) increased pupil dilation to unpleasant images, 3) higher neural reactivity to unpleasant images, and 4) a greater likelihood of having bottom-up attention drawn to unpleasant stimuli. Findings suggest that negative AH are associated with greater Defensive Motivational System activation in terms of central and autonomic nervous system response. |
Júlio Medeiros; André Bernardes; Ricardo Couceiro; Paulo Oliveira; Henrique Madeira; César Teixeira; Paulo Carvalho Optimal frequency bands for pupillography for maximal correlation with HRV Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–17, 2025. @article{Medeiros2025, Assessing cognitive load using pupillography frequency features presents a persistent challenge due to the lack of consensus on optimal frequency limits. This study aims to address this challenge by exploring pupillography frequency bands and seeking clarity in defining the most effective ranges for cognitive load assessment. From a controlled experiment involving 21 programmers performing software bug inspection, our study pinpoints the optimal low-frequency (0.06-0.29 Hz) and high-frequency (0.29-0.49 Hz) bands. Correlation analysis yielded a geometric mean of 0.238 compared to Heart Rate Variability features, with individual correlations for low-frequency, high-frequency, and their ratio at 0.279, 0.168, and 0.286, respectively. Extending the study to 51 participants, including a different experiment focusing on mental arithmetic tasks, validated the previous findings and further refined bands, maintaining effectiveness with a geometric mean correlation of 0.236 and surpassing common frequency bands reported in the existing literature. This study represents a pivotal step toward converging and establishing a coherent framework for frequency band definition to be used in pupillography analysis. Furthermore, based on this, it also contributes insights into the importance of more integration and adoption of eye-tracking with pupillography technology into authentic software development contexts for cognitive load assessment at a very fine level of granularity. |
2024 |
Güven Kandemir; Chris Olivers Comparing neural correlates of memory encoding and maintenance for foveal and peripheral stimuli Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 9, pp. 1807–1826, 2024. @article{Kandemir2024, Visual working memory is believed to rely on top–down attentional mechanisms that sustain active sensory representations in early visual cortex, a mechanism referred to as sensory recruitment. However, both bottom–up sensory input and top–down attentional modulations thereof appear to prioritize the fovea over the periphery, such that initially peripheral percepts may even be assimilated by foveal processes. This raises the question whether and how visual working memory differs for central and peripheral input. To address this, we conducted a delayed orientation recall task in which an orientation was presented either at the center of the screen or at 15° eccentricity to the left or right. Response accuracy, EEG activity, and gaze position were recorded from 30 participants. Accuracy was slightly but significantly higher for foveal versus peripheral memories. Decoding of EEG recordings revealed a clear dissociation between early sensory and later maintenance signals. Although sensory signals were clearly decodable for foveal stimuli, they were not for peripheral input. In contrast, maintenance signals were equally decodable for both foveal and peripheral memories, suggesting comparable top–down components regardless of eccentricity. Moreover, although memory representations were initially spatially specific and reflected in voltage fluctuations, later during the maintenance period, they generalized across locations, as emerged in alpha oscillations, thus revealing a dynamic transformation within memory from separate sensory traces to what we propose are common output-related codes. Furthermore, the combined absence of reliable decoding of sensory signals and robust presence of maintenance decoding indicates that storage activity patterns as measured by EEG reflect signals beyond primary visual cortex. We discuss the implications for the sensory recruitment hypothesis. |
Golan Karvat; Nir Ofir; Ayelet N. Landau Sensory drive modifies brain dynamics and the temporal integration window Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 4, pp. 614–631, 2024. @article{Karvat2024, Perception is suggested to occur in discrete temporal win-dows, clocked by cycles of neural oscillations. An important testable prediction of this theory is that individuals' peak frequencies of oscillations should correlate with their ability to segregate the appearance of two successive stimuli. An influential study tested this prediction and showed that individual peak frequency of spontaneously occurring alpha (8–12 Hz) corre-lated with the temporal segregation threshold between two successive flashes of light [Samaha, J., & Postle, B. R. The speed of alpha-band oscillations predicts the temporal resolution of visual perception. Current Biology, 25, 2985–2990, 2015]. How-ever, these findings were recently challenged [Buergers, S., & Noppeney, U. The role of alpha oscillations in temporal binding within and across the senses. Nature Human Behaviour, 6, 732– 742, 2022]. To advance our understanding of the link between oscillations and temporal segregation, we devised a novel experimental approach. Rather than relying entirely on spontaneous brain dynamics, we presented a visual grating before the flash stimuli that is known to induce continuous oscillations in the gamma band (45–65 Hz). By manipulating the contrast of the grating, we found that high contrast induces a stronger gamma response and a shorter temporal segregation threshold, compared to low-contrast trials. In addition, we used a novel tool to characterize sustained oscillations and found that, for half of the participants, both the low-and high-contrast gratings were accompanied by a sustained and phase-locked alpha oscil-lation. These participants tended to have longer temporal segregation thresholds. Our results suggest that visual stimulus drive, reflected by oscillations in specific bands, is related to the temporal resolution of visual perception. |
Samet Kılıç; Eser Sendesen; Filiz Aslan; Nurhan Erbil; Özgür Aydın; Didem Türkyılmaz Investigating sensitivity to auditory cognition in listening effort assessments: A simultaneous EEG and pupillometry study Journal Article In: Brain and Behavior, vol. 14, no. 11, pp. 1–12, 2024. @article{Kilic2024, Background: It is still not fully explained what kind of cognitive sources the methods used in the assessment of listening effort are more sensitive to and how these measurement results are related to each other. The aim of the study is to ascertain which neural resources crucial for listening effort are most sensitive to objective measurement methods using differently degraded speech stimuli. Methods: A total of 49 individuals between the ages of 19 and 34 with normal hearing participated in the study. In the first stage, simultaneous pupillometry, electroencephalogram (EEG), and single-task paradigm reaction time (RT) measurements were made during the challenging listening and repetition task with noise-vocoded speech. Two speech reception thresholds (SRT) (50% and 80%) for two vocoding conditions (16 and 6 channels) were collected, resulting in 4 conditions. In the second stage, the Rey Auditory Verbal Learning Test (RAVLT) and the test of attention in listening (TAIL) were applied. Stepwise linear regression analyses were conducted to examine the predictors of listening effort measurements. Results: A significant difference was found between 6 and 16 channel stimuli in both pupil dilation change and EEG alpha band power change. In the hardest listening condition, whereas RAVLT scores are significant predictors of pupil dilation change, TAIL scores are significant predictors of EEG alpha power. As the stimulus difficulty decreased, the factors that predicted both EEG and pupillometry results decreased. In the single-task paradigm, a significant regression model could not be obtained at all four difficulty levels. Conclusion: As a result of the study, it was found that the pupil dilation change was more sensitive to auditory memory skills and the EEG alpha power change was more sensitive to auditory attention skills. To our knowledge, this study is the first to investigate the sensitivity of different listening effort measurement methods to auditory cognitive skills. |
Silvia Erika Kober; Guilherme Wood; Sarah Schuster; Christof Körner Do miniature eye movements affect neurofeedback training performance? A combined EEG-eye tracking study Journal Article In: Applied Psychophysiology Biofeedback, vol. 49, no. 2, pp. 313–327, 2024. @article{Kober2024, EEG-based neurofeedback is a prominent method to modulate one's own brain activity in a desired direction. However, the EEG signal can be disturbed by artifacts, e.g., eye movements, which can consequently confound the neurofeedback performance. Involuntary miniature eye movements can be hardly detected by conventional EEG correction methods such as recording the electro-oculogram (EOG) and subtracting EOG activity from the EEG signal. However, such miniature eye movements can influence EEG activity, especially in the Gamma frequency range, enormously. In the present study, we investigated whether power in different EEG frequencies can be effectively modulated by self-control of brain signals during neurofeedback training and/or whether changes in EEG power are provoked by miniature eye movements during the training. To this end, 24 participants performed one session of SMR and one session of Gamma neurofeedback training. Additionally, in each training session sham feedback was performed. An eye tracker was used to detect miniature eye movements (< 1°) during neurofeedback training. About two thirds of the participants were able to increase their SMR power over the course of NF training, while one third was able to increase Gamma power. Generally, miniature eye movements induced a strong Gamma power increase. The number of eye movements also increased numerically over the course of the NF training. However, we did not find a significant relationship with the NF training performance. This is a first indication that miniature saccades do not affect NF training performance, but should not be neglected during NF training. Our results have to be confirmed in future studies. |
Jan Kujala; Sasu Mäkelä; Pauliina Ojala; Jukka Hyönä; Riitta Salmelin Beta- and gamma-band cortico-cortical interactions support naturalistic reading of continuous text Journal Article In: European Journal of Neuroscience, vol. 59, no. 2, pp. 238–251, 2024. @article{Kujala2024, Large-scale integration of information across cortical structures, building on neural connectivity, has been proposed to be a key element in supporting human cognitive processing. In electrophysiological neuroimaging studies of reading, quantification of neural interactions has been limited to the level of isolated words or sentences due to artefacts induced by eye movements. Here, we combined magnetoencephalography recording with advanced artefact rejection tools to investigate both cortico-cortical coherence and directed neural interactions during naturalistic reading of full-page texts. Our results show that reading versus visual scanning of text was associated with wide-spread increases of cortico-cortical coherence in the beta and gamma bands. We further show that the reading task was linked to increased directed neural interactions compared to the scanning task across a sparse set of connections within a wide range of frequencies. Together, the results demonstrate that neural connectivity flexibly builds on different frequency bands to support continuous natural reading. |
Nan Li; Suiping Wang; Florian Kornrumpf; Werner Sommer; Olaf Dimigen Parafoveal and foveal N400 effects in natural reading: A timeline of semantic processing from fixation-related potentials Journal Article In: Psychophysiology, vol. 61, no. 5, pp. 1–25, 2024. @article{Li2024c, The depth at which parafoveal words are processed during reading is an ongoing topic of debate. Recent studies using RSVP-with-flanker paradigms have shown that implausible words within sentences elicit an N400 component while they are still in parafoveal vision, suggesting that the semantics of parafoveal words can be accessed to rapidly update the sentence representation. To study this effect in natural reading, we combined the coregistration of eye movements and EEG with the deconvolution modeling of fixation-related potentials (FRPs) to test whether semantic plausibility is processed parafoveally during Chinese sentence reading. For one target word per sentence, both its parafoveal and foveal plausibility were orthogonally manipulated using the boundary paradigm. Consistent with previous eye movement studies, we observed a delayed effect of parafoveal plausibility on fixation durations that only emerged on the foveal word. Crucially, in FRPs aligned to the pretarget fixation, a clear N400 effect emerged already based on parafoveal plausibility, with more negative voltages for implausible previews. Once participants fixated the target, we again observed an N400 effect of foveal plausibility. Interestingly, this foveal N400 was absent whenever the preview had been implausible, indicating that when a word's (im)plausibility is already processed in parafoveal vision, this information is not revised anymore upon direct fixation. Implausible words also elicited a late positive component (LPC), but exclusively when in foveal vision. Our results not only provide convergent neural and behavioral evidence for the parafoveal uptake of semantic information, but also indicate different contributions of parafoveal versus foveal information toward higher level sentence processing. |
Elisabeth Lindner; Andrea Desantis; Felicia Pei Hsin Cheng; Alexander Gail In: NeuroImage, vol. 297, pp. 1–14, 2024. @article{Lindner2024, Self-initiated sensory action effects are widely assumed to lead to less intense perception and reduced neural responses compared to externally triggered stimuli (sensory attenuation). However, it is unclear if sensory attenuation occurs in all cases of action-effect prediction. Specifically, when predicted action-effects are relevant to determine follow-up actions attenuation could be detrimental. We quantified auditory event-related potentials (ERP) in electroencephalography (EEG) when human participants created two-sound sequences by pressing two keys on a keyboard associated with different pitch, giving rise to identity-specific action-effect prediction after the first keypress. The first sound corresponded to (congruent) or violated (incongruent) the predicted pitch and was either relevant for the selection of the second keypress to correctly complete the sequence (Relevance) or irrelevant (Control Movement), or there was only one keypress and sound (Baseline). We found a diminished P2-timed ERP component in incongruent compared to congruent trials when the sound was relevant for the subsequent action. This effect of action-effect prediction was due to an ERP reduction for incongruent relevant sounds compared to incongruent irrelevant sounds at P2 latencies and correlated negatively with modulations of pupil dilation. Contrary to our expectation, we did not observe an N1 modulation by congruency in any condition. Attenuation of the N1 component seems absent for predicted identity-specific auditory action effects, while P2-timed ERPs as well as pupil size are sensitive to predictability, at least when action effects are relevant for the selection of the next action. Incongruent relevant stimuli thereby take a special place and seem to be subject to attentional modulations and error processing. |
Federico Mancinelli; Juliana K. Sporrer; Vladislav Myrov; Filip Melinscak; Josua Zimmermann; Huaiyu Liu; Dominik R. Bach Dimensionality and optimal combination of autonomic fear-conditioning measures in humans Journal Article In: Behavior Research Methods, vol. 56, no. 6, pp. 6119–6129, 2024. @article{Mancinelli2024, Fear conditioning, also termed threat conditioning, is a commonly used learning model with clinical relevance. Quantification of threat conditioning in humans often relies on conditioned autonomic responses such as skin conductance responses (SCR), pupil size responses (PSR), heart period responses (HPR), or respiration amplitude responses (RAR), which are usually analyzed separately. Here, we investigate whether inter-individual variability in differential conditioned responses, averaged across acquisition, exhibits a multi-dimensional structure, and the extent to which their linear combination could enhance the precision of inference on whether threat conditioning has occurred. In a mega-analytic approach, we re-analyze nine data sets including 256 individuals, acquired by the group of the last author, using standard routines in the framework of psychophysiological modeling (PsPM). Our analysis revealed systematic differences in effect size between measures across datasets, but no evidence for a multidimensional structure across various combinations of measures. We derive the statistically optimal weights for combining the four measures and subsets thereof, and we provide out-of-sample performance metrics for these weights, accompanied by bias-corrected confidence intervals. We show that to achieve the same statistical power, combining measures allows for a relevant reduction in sample size, which in a common scenario amounts to roughly 24%. To summarize, we demonstrate a one-dimensional structure of threat conditioning measures, systematic differences in effect size between measures, and provide weights for their optimal linear combination in terms of maximal retrodictive validity. |
Rocío Mayol-Troncoso; Pablo A. Gaspar; Roberto Verdugo; Juan J. Mariman; Pedro E. Maldonado Fixational eye movements and their associated evoked potentials during natural vision are altered in schizophrenia Journal Article In: Schizophrenia Research: Cognition, vol. 38, pp. 1–9, 2024. @article{MayolTroncoso2024, Background: Visual exploration is abnormal in schizophrenia; however, few studies have investigated the physiological responses during selecting objectives in more ecological scenarios. This study aimed to demonstrate that people with schizophrenia have difficulties observing the prominent elements of an image due to a deficit mechanism of sensory modulation (active sensing) during natural vision. Methods: An electroencephalogram recording with eye tracking data was collected on 18 healthy individuals and 18 people affected by schizophrenia while looking at natural images. These had a prominent color element and blinking produced by changes in image luminance. Results: We found fewer fixations when all images were scanned, late focus on prominent image areas, decreased amplitude in the eye-fixation-related potential, and decreased intertrial coherence in the SCZ group. Conclusions: The decrease in the visual attention response evoked by the prominence of visual stimuli in patients affected by schizophrenia is generated by a reduction in endogenous attention mechanisms to initiate and maintain visual exploration. Further work is required to explain the relationship of this decrease with clinical indicators. |
Xiaolin Mei; Shuyuan Chen; Xinyi Xia; Bo Yang; Yanping Liu Neural correlates for word-frequency effect in Chinese natural reading Journal Article In: Attention, Perception, & Psychophysics, no. April, 2024. @article{Mei2024, Word frequency effect has always been of interest for reading research because of its critical role in exploring mental processing underlying reading behaviors. Access to word frequency information has long been considered an indicator of the beginning of lexical processing and the most sensitive marker for studying when the brain begins to extract semantic information Sereno & Rayner, Brain and Cognition, 42, 78–81, (2000), Trends in Cognitive Sciences, 7, 489–493, (2003). While the word frequency effect has been extensively studied in numerous eye-tracking and traditional EEG research using the RSVP paradigm, there is a lack of corresponding evidence in studies of natural reading. To find the neural correlates of the word frequency effect, we conducted a study of Chinese natural reading using EEG and eye-tracking coregistration to examine the time course of lexical processing. Our results reliably showed that the word frequency effect first appeared in the N200 time window and the bilateral occipitotemporal regions. Additionally, the word frequency effect was reflected in the N400 time window, spreading from the occipital region to the central parietal and frontal regions. Our current study provides the first neural correlates for word-frequency effect in natural Chinese reading so far, shedding new light on understanding lexical processing in natural reading and could serve as an important basis for further reading study when considering neural correlates in a realistic manner. |
David Melcher; Ani Alaberkyan; Chrysi Anastasaki; Xiaoyi Liu; Michele Deodato; Gianluca Marsicano; Diogo Almeida An early effect of the parafoveal preview on post-saccadic processing of English words Journal Article In: Attention, Perception, & Psychophysics, pp. 1–26, 2024. @article{Melcher2024, A key aspect of efficient visual processing is to use current and previous information to make predictions about what we will see next. In natural viewing, and when looking at words, there is typically an indication of forthcoming visual information from extrafoveal areas of the visual field before we make an eye movement to an object or word of interest. This “preview effect” has been studied for many years in the word reading literature and, more recently, in object perception. Here, we integrated methods from word recognition and object perception to investigate the timing of the preview on neural measures of word recognition. Through a combined use of EEG and eye-tracking, a group of multilingual participants took part in a gaze-contingent, single-shot saccade experiment in which words appeared in their parafoveal visual field. In valid preview trials, the same word was presented during the preview and after the saccade, while in the invalid condition, the saccade target was a number string that turned into a word during the saccade. As hypothesized, the valid preview greatly reduced the fixation-related evoked response. Interestingly, multivariate decoding analyses revealed much earlier preview effects than previously reported for words, and individual decoding performance correlated with participant reading scores. These results demonstrate that a parafoveal preview can influence relatively early aspects of post-saccadic word processing and help to resolve some discrepancies between the word and object literatures. |
Danilo Menicucci; Silvia Animali; Eleonora Malloggi; Angelo Gemignani; Enrica Bonanni; Francesco Fornai; Filippo Sean Giorgi; Paola Binda Correlated P300b and phasic pupil-dilation responses to motivationally significant stimuli Journal Article In: Psychophysiology, vol. 61, no. 6, pp. 1–14, 2024. @article{Menicucci2024, Motivationally significant events like oddball stimuli elicit both a characteristic event-related potential (ERPs) known as P300 and a set of autonomic responses including a phasic pupil dilation. Although co-occurring, P300 and pupil-dilation responses to oddball events have been repeatedly found to be uncorrelated, suggesting separate origins. We re-examined their relationship in the context of a three-stimulus version of the auditory oddball task, independently manipulating the frequency (rare vs. repeated) and motivational significance (relevance for the participant's task) of the stimuli. We used independent component analysis to derive a P300b component from EEG traces and linear modeling to separate a stimulus-related pupil-dilation response from a potentially confounding action-related response. These steps revealed that, once the complexity of ERP and pupil-dilation responses to oddball targets is accounted for, the amplitude of phasic pupil dilations and P300b are tightly and positively correlated (across participants: r =.69 p =.002), supporting their coordinated generation. |
Madaline Mocchi; Eleonora Bartoli; John Magnotti; Jan Willem Gee; Brian Metzger; Bailey Pascuzzi; Raissa Mathura; Suhruthaa Pulapaka; Wayne Goodman; Sameer Sheth; Matthew J. McGinley; Kelly Bijanki Aperiodic spectral slope tracks the effects of brain state on saliency responses in the human auditory cortex Journal Article In: Scientific Reports, vol. 14, no. 1, pp. 1–14, 2024. @article{Mocchi2024, Alteration of responses to salient stimuli occurs in a wide range of brain disorders and may be rooted in pathophysiological brain state dynamics. Specifically, tonic and phasic modes of activity in the reticular activating system (RAS) influence, and are influenced by, salient stimuli, respectively. The RAS influences the spectral characteristics of activity in the neocortex, shifting the balance between low- and high-frequency fluctuations. Aperiodic ‘1/f slope' has emerged as a promising composite measure of these brain state dynamics. However, the relationship of 1/f slope to state-dependent processes, such as saliency, is less explored, particularly intracranially in humans. Here, we record pupil diameter as a measure of brain state and intracranial local field potentials in auditory cortical regions of human patients during an auditory oddball stimulus paradigm. We find that phasic high-gamma band responses in auditory cortical regions exhibit an inverted-u shaped relationship to tonic state, as reflected in the 1/f slope. Furthermore, salient stimuli trigger state changes, as indicated by shifts in the 1/f slope. Taken together, these findings suggest that 1/f slope tracks tonic and phasic arousal state dynamics in the human brain, increasing the interpretability of this metric and supporting it as a potential biomarker in brain disorders. |
Kieran S. Mohr; Anna C. Geuzebroek; Simon P. Kelly Visual cortical area contributions to the transient, multifocal and steady- state VEP: A forward model- informed analysis Journal Article In: Imaging Neuroscience, vol. 2, pp. 1–26, 2024. @article{Mohr2024, Central to our understanding of how visual- evoked potentials (VEPs) contribute to visual processing is the question of where their anatomical sources are. Three well- established measures of low- level visual cortical activity are widely used: the first component (“C1”) of the transient and multifocal VEP, and the steady- state VEP (SSVEP). Although primary visual cortex (V1) activity has often been implicated in the generation of all three signals, their dominant sources remain uncertain due to the limited resolution and methodological heterogeneity of source modelling. Here, we provide the first characterisation of all three signals in one analytic framework centred on the “cruciform model”, which describes how scalp topographies of V1 activity vary with stimulus location due to the retinotopy and unique folding pattern of V1. We measured the transient C1, multifocal C1, and SSVEPs driven by an 18.75 Hz and 7.5 Hz flicker, and regressed them against forward models of areas V1, V2, and V3 generated from the Benson- 2014 retino- topy atlas. The topographic variations of all four VEP signals across the visual field were better captured by V1 mod- els, explaining between 2 and 6 times more variance than V2/V3. Models with all three visual areas improved fit further, but complementary analyses of temporal dynamics across all three signals indicated that the bulk of extrastriate contributions occur considerably later than V1. Overall, our data support the use of peak C1 amplitude and SSVEPs to probe V1 activity, although the SSVEP contains stronger extrastriate contributions. Moreover, we provide elabo- rated heuristics to distinguish visual areas in VEP data based on signal lateralisation as well as polarity inversion. |
Caoimhe Moran; Philippa A. Johnson; Ayelet N. Landau; Hinze Hogendoorn Decoding remapped spatial information in the peri-saccadic period Journal Article In: The Journal of Neuroscience, vol. 44, no. 30, pp. 1–12, 2024. @article{Moran2024, It has been suggested that, prior to a saccade, visual neurons predictively respond to stimuli that will fall in their receptive fields after completion of the saccade. This saccadic remapping process is thought to compensate for the shift of the visual world across the retina caused by eye movements. To map the timing of this predictive process in the brain, we recorded neural activity using electroencephalography during a saccade task. Human participants (male and female) made saccades between two fixation points while covertly attending to oriented gratings briefly presented at various locations on the screen. Data recorded during trials in which participants maintained fixation were used to train classifiers on stimuli in different positions. Subsequently, data collected during saccade trials were used to test for the presence of remapped stimulus information at the post-saccadic retinotopic location in the peri-saccadic period, providing unique insight into when remapped information becomes available. We found that the stimulus could be decoded at the remapped location ∼180 ms post-stimulus onset, but only when the stimulus was presented 100–200 ms before saccade onset. Within this range, we found that the timing of remapping was dictated by stimulus onset rather than saccade onset. We conclude that presenting the stimulus immediately before the saccade allows for optimal integration of the corollary discharge signal with the incoming peripheral visual information, resulting in a remapping of activation to the relevant post-saccadic retinotopic neurons. |
Kimberley Mulder; Sophie Brand; Lou Boves; Mirjam Ernestus Processing reduced speech in the L1 and L2: A combined eye-tracking and ERP study Journal Article In: Language, Cognition and Neuroscience, vol. 39, no. 5, pp. 527–551, 2024. @article{Mulder2024, We examined the cognitive processes underlying the comprehension of reduced word pronunciation variants in natives and advanced learners of French. In a passive listening visual world task, participants heard sentences containing either a reduced or a full form and saw pictures representing the target word, a phonological competitor and two neutral distractors. After each sentence they saw a picture and had to decide whether it matched the content of that sentence. Eye movements and EEG were recorded simultaneously. Because the two recordings offer complementary information about cognitive processes, we developed methods for analysing the signals in combination. We found a stronger effect of reduction on phonetic processing and semantic integration in learners than in natives, but the effects are different from the N100/N400 and P600 effects found in previous research. Time-locking EEG signals on fixation moments in the eye movements offers a window onto the time course of semantic integration. |
Marta Font-Alaminos; Nadia Paraskevoudi; Jordi Costa-Faidella; Iria SanMiguel Do actions structure auditory memory? Action-based event segmentation effects on sensory responses, pupil dilation and sequential memory Journal Article In: Psychophysiology, vol. 61, pp. 1–19, 2024. @article{FontAlaminos2024, Our actions shape our everyday experience: what we experience, how we perceive, and remember it are deeply affected by how we interact with the world. Performing an action to deliver a stimulus engages neurophysiological processes which are reflected in the modulation of sensory and pupil responses. We hypothesized that these processes shape memory encoding, parsing the experience by grouping self- and externally generated stimuli into differentiated events. Participants encoded sound sequences, in which either the first or last few sounds were self-generated and the rest externally generated. We tested recall of the sequential order of sounds that had originated from the same (within event) or different sources (across events). Memory performance was not higher for within-event sounds, suggesting that actions did not structure the memory representation. However, during encoding, we observed the expected electrophysiological response attenuation for self-generated sounds, together with increased pupil dilation triggered by actions. Moreover, at the boundary between events, physiological responses to the first sound from the new source were influenced by the direction of the source switch. Our results suggest that introducing actions creates a stronger contextual shift than removing them, even though actions do not directly contribute to memory performance. This study contributes to our understanding of how interacting with sensory input shapes experiences by exploring the relationships between action effects on sensory responses, pupil dilation, and memory encoding. Importantly, it challenges the notion of a meaningful contribution from low-level neurophysiological mechanisms associated with action execution in the modulation of the self-generation effect. |
Stefan L. Frank; Anna Aumeistere An eye-tracking-with-EEG coregistration corpus of narrative sentences Journal Article In: Language Resources and Evaluation, vol. 58, no. 2, pp. 641–657, 2024. @article{Frank2024, We present the Radboud Coregistration Corpus of Narrative Sentences (RaCCooNS), the first freely available corpus of eye-tracking-with-EEG data collected while participants read narrative sentences in Dutch. The corpus is intended for studying human sentence comprehension and for evaluating the cognitive validity of computational language models. RaCCooNS contains data from 37 participants (3 of which eye tracking only) reading 200 Dutch sentences each. Less predictable words resulted in significantly longer reading times and larger N400 sizes, replicating well-known surprisal effects in eye tracking and EEG simultaneously. We release the raw eye-tracking data, the preprocessed eye-tracking data at the fixation, word, and trial levels, the raw EEG after merger with eye-tracking data, and the preprocessed EEG data both before and after ICA-based ocular artifact correction. |
Francesco Grassia; Louisa Kulkeb; Alex Lepauvrec; Anne Schachta Relevance acquisition through motivational incentives: Modeling the time-course of associative learning and the role of visual features Journal Article In: Imaging Neuroscience, vol. 2, pp. 1–20, 2024. @article{Grassia2024, Motivational relevance associated with symbolic stimuli impacts both neural and behavioral responses, similar to visual stimuli with inherent emotional valence. However, the specific effects of associated relevance on early sensory stages and lexico-semantic processing of these stimuli remain unclear, part icularly considering the role of low- level visual features in relevance acquisition. To address these issues, we employed an associative learning paradigm in which we manipulated visual features, but not the stimuli themselves. The study (N = 48) included a learning phase, where pseudowords were associated with either gain, loss, or neutral outcomes. This was followed by a test phase the next day, involving an old/new decision task, in which stimuli were presented in either the same or a different font. During both phases, pupil responses and event- related brain potentials (P1, Early Posterior Negativity (EPN), Late Positive Complex (LPC), P3) were measured. Stronger pupil responses and increased neural activation in early visual encoding (P1) and lexico- semantic processing (EPN) were observed during relevance acquisition, particularly for loss associations. After relevance acquisition, the most substantial effect on modulating lexico-semantic processing was observed for gain associations, as evidenced by both behavioral responses and neural activity. During the test phase, exposure to incongruent visual features of the stimuli influenced the same processes that were observed during rele- vance acquisition. Notably, these effects of visual feature congruence were independent of those of associated motivational relevance. These results highlight the dynamic nature of motivational relevance effects, revealing differential effects observed during acquisition and the test phase, as well as between earlier perceptual processing and later neural and behavioral responses. |
John P. Grogan; Matthias Raemaekers; Maaike M. H. Swieten; Alexander L. Green; Martin J. Gillies; Sanjay G. Manohar Muscarinic receptors mediate motivation via preparatory neural activity in humans Journal Article In: eLife, vol. 13, pp. 1–28, 2024. @article{Grogan2024, Motivation depends on dopamine, but might be modulated by acetylcholine which influences 10 dopamine release in the striatum, and amplifies motivation in animal studies. A corresponding effect in humans would be important clinically, since anticholinergic drugs are frequently used in Parkinson's disease, a condition that can also disrupt motivation. Reward and dopamine make us more ready to respond, as indexed by reaction times (RT), and move faster, sometimes termed vigour. These effects may be controlled by preparatory processes that can be tracked using EEG. We 15 measured vigour in a placebo-controlled, double-blinded study of trihexyphenidyl (THP), a muscarinic antagonist, with an incentivised eye movement task and EEG. Participants responded faster and with greater vigour when incentives were high, but THP blunted these motivation effects, suggesting that muscarinic receptors facilitate invigoration by reward. Preparatory EEG build-up (contingent negative variation; CNV) was strengthened by high incentives and by muscarinic 20 blockade. The amplitude of preparatory activity predicted both vigour and RT, although over distinct scalp regions. Frontal activity predicted vigour, whereas a larger, earlier, central component predicted RT. Indeed the incentivisation of RT was partly mediated by the CNV, though vigour was not. Moreover, the CNV mediated the drug's effect on dampening incentives, suggesting that muscarinic receptors underlie the motivational influence on this preparatory activity. Taken 25 together, these findings show that a muscarinic blocker used to treat Parkinson's disease impairs motivated action in healthy people, and that medial frontal preparatory neural activity mediates this for RT. |
Courtney Guida; Minwoo J. B. Kim; Olivia A. Stibolt; Alyssa Lompado; James E. Hoffman The N400 component reflecting semantic and repetition priming of visual scenes is suppressed during the attentional blink Journal Article In: Attention, Perception, & Psychophysics, pp. 1–20, 2024. @article{Guida2024, In the attentional blink paradigm, participants attempt to identify two targets appearing in a rapidly presented stream of distractors. Report accuracy is typically high for the first target (T1) while identification of the second target (T2) is impaired when it follows within about 200–400 ms of T1. An important question is whether T2 is processed to a semantic level even when participants are unaware of its identity. We examined this issue in three studies that used natural scenes as stimuli and the N400 component of the event-related potential (ERP) as a measure of semantic priming. In the first experiment, the prime (e.g., a doghouse in a yard) was presented at the beginning of the trial and a test picture that was related (e.g., a dog standing in the kitchen) or unrelated (e.g., a coffee mug on a table) appeared as T2. In the second experiment, the prime was presented as T2 and the test picture appeared at the end of the picture sequence. In both experiments, we found robust semantic priming when participants were aware of the identity of the blinked picture and an absence of priming when they were unaware. In Experiment 3, we used identity priming to assess whether earlier representations preceding semantics were preserved, and again found that priming critically depended on awareness of the prime's identity. These results suggest that semantic priming in scenes, as measured with the N400, is a higher-level process that critically depends on attention and awareness. |
Divya Gulati; Supratim Ray Auditory and visual gratings elicit distinct gamma responses Journal Article In: eNeuro, vol. 11, no. 4, pp. 1–13, 2024. @article{Gulati2024, Sensory stimulation is often accompanied by fluctuations at high frequencies (>30 Hz) in brain signals. These could be “narrowband” oscillations in the gamma band (30–70 Hz) or nonoscillatory “broadband” high-gamma (70–150 Hz) activity. Narrowband gamma oscillations, which are induced by presenting some visual stimuli such as gratings and have been shown to weaken with healthy aging and the onset of Alzheimer's disease, hold promise as potential biomarkers. However, since delivering visual stimuli is cumbersome as it requires head stabilization for eye tracking, an equivalent auditory paradigm could be useful. Although simple auditory stimuli have been shown to produce high-gamma activity, whether specific auditory stimuli can also produce narrowband gamma oscillations is unknown. We tested whether auditory ripple stimuli, which are considered an analog to visual gratings, could elicit narrowband oscillations in auditory areas. We recorded 64-channel electroencephalogram from male and female (18 each) subjects while they either fixated on the monitor while passively viewing static visual gratings or listened to stationary and moving ripples, played using loudspeakers, with their eyes open or closed. We found that while visual gratings induced narrowband gamma oscillations with suppression in the alpha band (8–12 Hz), auditory ripples did not produce narrowband gamma but instead elicited very strong broadband high-gamma response and suppression in the beta band (14–26 Hz). Even though we used equivalent stimuli in both modalities, our findings indicate that the underlying neuronal circuitry may not share ubiquitous strategies for stimulus processing. |
Eren Günseli; Joshua J. Foster; David W. Sutterer; Lara Todorova; Edward K. Vogel; Edward Awh Encoded and updated spatial working memories share a common representational format in alpha activity Journal Article In: iScience, vol. 27, no. 2, pp. 1–15, 2024. @article{Guenseli2024, Working memory (WM) flexibly updates information to adapt to the dynamic environment. Here, we used alpha-band activity in the EEG to reconstruct the content of dynamic WM updates and compared this representational format to static WM content. An inverted encoding model using alpha activity precisely tracked both the initially encoded position and the updated position following an auditory cue signaling mental updating. The timing of the update, as tracked in the EEG, correlated with reaction times and saccade latency. Finally, cross-training analyses revealed a robust generalization of alpha-band reconstruction of WM contents before and after updating. These findings demonstrate that alpha activity tracks the dynamic updates to spatial WM and that the format of this activity is preserved across the encoded and updated representations. Thus, our results highlight a new approach for measuring updates to WM and show common representational formats during dynamic mental updating and static storage. |
Annika Hense; Antje Peters; Maximilian Bruchmann; Torge Dellert; Thomas Straube Electrophysiological correlates of sustained conscious perception Journal Article In: Scientific Reports, vol. 14, no. 1, pp. 1–11, 2024. @article{Hense2024, Previous research on the neural correlates of consciousness (NCC) in visual perception revealed an early event-related potential (ERP), the visual awareness negativity (VAN), to be associated with stimulus awareness. However, due to the use of brief stimulus presentations in previous studies, it remains unclear whether awareness-related negativities represent a transient onset-related response or correspond to the duration of a conscious percept. Studies are required that allow prolonged stimulus presentation under aware and unaware conditions. The present ERP study aimed to tackle this challenge by using a novel stimulation design. Male and female human participants (n = 62) performed a visual task while task-irrelevant line stimuli were presented in the background for either 500 or 1000 ms. The line stimuli sometimes contained a face, which needed so-called visual one-shot learning to be seen. Half of the participants were informed about the presence of the face, resulting in faces being perceived by the informed but not by the uninformed participants. Comparing ERPs between the informed and uninformed group revealed an enhanced negativity over occipitotemporal electrodes that persisted for the entire duration of stimulus presentation. Our results suggest that sustained visual awareness negativities (SVAN) are associated with the duration of stimulus presentation. |
Xin Huang; Brian W. L. Wong; Hezul Tin Yan Ng; Werner Sommer; Olaf Dimigen; Urs Maurer Neural mechanism underlying preview effects and masked priming effects in visual word processing Journal Article In: Attention, Perception, & Psychophysics, pp. 1–20, 2024. @article{Huang2024f, Two classic experimental paradigms – masked repetition priming and the boundary paradigm – have played a pivotal role in understanding the process of visual word recognition. Traditionally, these paradigms have been employed by different communities of researchers, with their own long-standing research traditions. Nevertheless, a review of the literature suggests that the brain-electric correlates of word processing established with both paradigms may show interesting similarities, in particular with regard to the location, timing, and direction of N1 and N250 effects. However, as of yet, no direct comparison has been undertaken between the two paradigms. In the current study, we used combined eye-tracking/EEG to perform such a within-subject comparison using the same materials (single Chinese characters) as stimuli. To facilitate direct comparisons, we used a simplified version of the boundary paradigm – the single word boundary paradigm. Our results show the typical early repetition effects of N1 and N250 for both paradigms. However, repetition effects in N250 (i.e., a reduced negativity following identical-word primes/previews as compared to different-word primes/previews) were larger with the single word boundary paradigm than with masked priming. For N1 effects, repetition effects were similar across the two paradigms, showing a larger N1 after repetitions as compared to alternations. Therefore, the results indicate that at the neural level, a briefly presented and masked foveal prime produces qualitatively similar facilitatory effects on visual word recognition as a parafoveal preview before a single saccade, although such effects appear to be stronger in the latter case. |
Caitlin M. Hudac; Kelsey Dommer; Monique Mahony; Trent D. DesChamps; Brianna Cairney; Rachel Earl; Evangeline C. Kurtz-Nelson; Jessica Bradshaw; Raphael A. Bernier; Evan E. Eichler; Emily Neuhaus; Sara Jane Webb; Frederick Shic Visual and auditory attention in individuals with DYRK1A and SCN2A disruptive variants Journal Article In: Autism Research, pp. 1–13, 2024. @article{Hudac2024, This preliminary study sought to assess biomarkers of attention using electroencephalography (EEG) and eye tracking in two ultra-rare monogenic populations associated with autism spectrum disorder (ASD). Relative to idiopathic ASD (n = 12) and neurotypical comparison (n = 49) groups, divergent attention profiles were observed for the monogenic groups, such that individuals with DYRK1A (n = 9) exhibited diminished auditory attention condition differences during an oddball EEG paradigm whereas individuals with SCN2A (n = 5) exhibited diminished visual attention condition differences noted by eye gaze tracking when viewing social interactions. Findings provide initial support for alignment of auditory and visual attention markers in idiopathic ASD and neurotypical development but not monogenic groups. These results support ongoing efforts to develop translational ASD biomarkers within the attention domain. |
Henry M. Jones; Gisella K. Diaz; William X. Q. Ngiam; Edward Awh Electroencephalogram decoding reveals distinct processes for directing spatial attention and encoding into working memory Journal Article In: Psychological Science, vol. 35, no. 10, pp. 1108 –1138, 2024. @article{Jones2024, Past work reveals a tight relationship between spatial attention and storage in visual working memory. But is spatially attending an item tantamount to working memory encoding? Here, we tracked electroencephalography (EEG) signatures of spatial attention and working memory encoding while independently manipulating the number of memory items and the spatial extent of attention in two studies of adults (N = 39; N = 33). Neural measures of spatial attention tracked the position and size of the attended area independent of the number of individuated items encoded into working memory. At the same time, multivariate decoding of the number of items stored in working memory was insensitive to variations in the breadth and position of spatial attention. Finally, representational similarity analyses provided converging evidence for a pure load signal that is insensitive to the spatial extent of the stored items. Thus, although spatial attention is a persistent partner of visual working memory, it is functionally dissociable from the selection and maintenance of individuated representations in working memory. |
Henry M. Jones; William S. Thyer; Darius Suplica; Edward Awh Cortically disparate visual features evoke content-independent load signals during storage in working memory Journal Article In: The Journal of Neuroscience, vol. 44, no. 44, pp. 1–18, 2024. @article{Jones2024a, It is well established that holding information in working memory (WM) elicits sustained stimulus-specific patterns of neural activity. Nevertheless, here we provide evidence for a distinct class of neural activity that tracks the number of individuated items in working memory, independent of the type of visual features stored. We present two EEG studies of young adults of both sexes that provide robust evidence for a signal tracking the number of individuated representations in working memory, regardless of the specific feature values stored. In Study 1, subjects maintained either colors or orientations across separate blocks in a single session. We found near-perfect generalization of the load signal between these two conditions, despite being able to simultaneously decode which feature had been voluntarily stored. In Study 2, participants attended to two features with very distinct cortical representations: color and motion coherence. We again found evidence for a neural load signal that robustly generalized across these distinct visual features, even though cortically disparate regions process color and motion coherence. Moreover, representational similarity analysis provided converging evidence for a content-independent load signal, while simultaneously showing that unique variance in EEG activity tracked the specific features that were stored. We posit that this load signal reflects a content-independent "pointer" operation that binds objects to the current context while parallel but distinct neural signals represent the features that are stored for each item in memory. |
Yuan Zhang; Matteo Valsecchi; Karl R. Gegenfurtner; Jing Chen The execution of saccadic eye movements suppresses visual processing of both color and luminance in the early visual cortex of humans Journal Article In: Journal of Neurophysiology, vol. 131, no. 6, pp. 1156–1167, 2024. @article{Zhang2024n, Our eyes execute rapid, directional movements known as saccades, occurring several times per second, to focus on objects of interest in our environment. During these movements, visual sensitivity is temporarily reduced. Despite numerous studies on this topic, the underlying mechanism remains elusive, including a lingering debate on whether saccadic suppression affects the parvocellular visual pathway. To address this issue, we conducted a study employing steady-state visual evoked potentials (SSVEPs) elicited by chromatic and luminance stimuli while observers performed saccadic eye movements. We also employed an innovative analysis pipeline to enhance the signal-to-noise ratio, yielding superior results compared to the previous method. Our findings revealed a clear suppression effect on SSVEP signals during saccades compared to fixation periods. Notably, this suppression effect was comparable for both chromatic and luminance stimuli. We went further to measure the suppression effect across various contrast levels, which enabled us to model SSVEP responses with contrast response functions. The results suggest that saccades primarily reduce response gain without significantly affecting contrast gain and that this reduction applies uniformly to both chromatic and luminance pathways. In summary, our study provides robust evidence that saccades similarly suppress visual processing in both the parvocellular and magnocellular pathways within the human early visual cortex, as indicated by SSVEP responses. The observation that saccadic eye movements impact response gain rather than contrast gain implies that they influence visual processing through a multiplicative mechanism. |
Carolin Zsigo; Ellen Greimel; Regine Primbs; Jürgen Bartling; Gerd Schulte-Körne; Lisa Feldmann Frontal alpha asymmetry during emotion regulation in adults with lifetime major depression Journal Article In: Cognitive, Affective, & Behavioral Neuroscience, vol. 24, no. 3, pp. 552–566, 2024. @article{Zsigo2024, Emotion regulation (ER) often is impaired in current or remitted major depression (MD), although the extent of the deficits is not fully understood. Recent studies suggest that frontal alpha asymmetry (FAA) could be a promising electrophysiological measure to investigate ER. The purpose of this study was to investigate ER differences between participants with lifetime major depression (lifetime MD) and healthy controls (HC) for the first time in an experimental task by using FAA. We compared lifetime MD (n = 34) and HC (n = 25) participants aged 18–24 years in (a) an active ER condition, in which participants were instructed to reappraise negative images and (b) a condition in which they attended to the images while an EEG was recorded. We also report FAA results from an independent sample of adolescents with current MD (n = 36) and HC adolescents (n = 38). In the main sample, both groups were able to decrease self-reported negative affect in response to negative images through ER, without significant group differences. We found no differences between groups or conditions in FAA, which was replicated within the independent adolescent sample. The lifetime MD group also reported less adaptive ER in daily life and higher difficulty of ER during the task. The lack of differences between in self-reported affect and FAA between lifetime MD and HC groups in the active ER task indicates that lifetime MD participants show no impairments when instructed to apply an adaptive ER strategy. Implications for interventional aspects are discussed. |
Artyom Zinchenko; Markus Conci; Hermann J. Müller; Thomas Geyer Environmental regularities mitigate attentional misguidance in contextual cueing of visual search Journal Article In: Journal of Experimental Psychology: Learning, Memory, and Cognition, vol. 50, no. 5, pp. 699–711, 2024. @article{Zinchenko2024, Visual search is faster when a fixed target location is paired with a spatially invariant (vs. randomly changing) distractor configuration, thus indicating that repeated contexts are learned, thereby guiding attention to the target (contextual cueing [CC]). Evidence for memory-guided attention has also been revealed with electrophysiological (electroencephalographic [EEG]) recordings, starting with an enhanced early posterior neg- ativity (N1pc), which signals a preattentive bias toward the target, and, subsequently, attentional and postselective components, such as the posterior contralateral negativity (PCN) and contralateral delay activ- ity (CDA), respectively. Despite effective learning, relearning of previously acquired contexts is inflexible: The CC benefits disappear when the target is relocated to a new position within an otherwise invariant context and corresponding EEG correlates are diminished. The present study tested whether global statistical properties that induce predictions going beyond the immediate invariant layout can facilitate contextual relearning. Global statistical regularities were implemented by presenting repeated and nonrepeated displays in separate streaks (mini blocks) of trials in the relocation phase, with individual displays being presented in a fixed and thus predictable order. Our results revealed a significant CC effect (and an associated modulation of the N1pc, PCN, and CDA components) during initial learning. Critically, the global statistical regularities in the relocation phase also resulted in a reliable CC effect, thus revealing effective relearning with predictive streaks. Moreover, this relearning was reflected in an enhanced PCN amplitude for repeated relative to non- repeated contexts. Temporally ordered contexts may thus adapt memory-based guidance of attention, par- ticularly the allocation of covert attention in the visual display. |
Annika Ziereis; Anne Schacht Additive effects of emotional expression and stimulus size on the perception of genuine and artificial facial expressions: An ERP study Journal Article In: Scientific Reports, vol. 14, no. 1, pp. 1–15, 2024. @article{Ziereis2024, Seeing an angry individual in close physical proximity can not only result in a larger retinal representation of that individual and an enhanced resolution of emotional cues, but may also increase motivation for rapid visual processing and action preparation. The present study investigated the effects of stimulus size and emotional expression on the perception of happy, angry, non-expressive, and scrambled faces. We analyzed event-related potentials (ERPs) and behavioral responses of N = 40 participants who performed a naturalness classification task on real and artificially created facial expressions. While the emotion-related effects on accuracy for recognizing authentic expressions were modulated by stimulus size, ERPs showed only additive effects of stimulus size and emotional expression, with no significant interaction with size. This contrasts with previous research on emotional scenes and words. Effects of size were present in all included ERPs, whereas emotional expressions affected the N170, EPN, and LPC, irrespective of size. These results imply that the decoding of emotional valence in faces can occur even for small stimuli. Supra-additive effects in faces may necessitate larger size ranges or dynamic stimuli that increase arousal. |
Zhihan Zhang; Chuhan Wu; Hongyi Chen; Hongyang Chen CogAware: Cognition-Aware framework for sentiment analysis with textual representations Journal Article In: Knowledge-Based Systems, vol. 299, pp. 1–9, 2024. @article{Zhang2024r, Sentiment analysis has become an important research area in artificial intelligence. Recently, the integration of sentiment analysis with cognitive neuroscience in natural language processing (NLP) tasks has attracted widespread attention. Cognitive signals and textual signals (i.e. word embeddings) both contain distinctive information for sentiment analysis tasks. However, most previous studies cannot effectively capture the specific features and cross-domain features while integrating cognitive signals acquired from brain activity and textual signals obtained from natural language processing (NLP). To address this issue, we propose CogAware, which learns to obtain a deep representation that combines purified specific features with cross-domain features from textual and cognitive signals. CogAware employs four private encoders to extract specific or cross-domain features from textual and cognitive signals alternately. It also employs feature reinforcement and orthogonality regularization to separate specific and cross-domain features from each modality. Moreover, a shared encoder and a modality discriminator are used to further capture cross-domain features from different modalities. Our designed architecture utilizes cognitive signals and word embeddings during model training, yet relies solely on word embeddings for model inference. Experiments on a public dataset show that CogAware achieves new state-of-the-art performance on the sentiment analysis task compared with other existing models. The source code of CogAware is available at: https://github.com/zhejiangzhuque/CogAware. |
Yuhong Zhang; Qin Li; Sujal Nahata; Tasnia Jamal; Shih Cheng; Gert Cauwenberghs; Tzyy Ping Jung Integrating large language model, eeg, and eye-tracking for word-level neural state classification in reading comprehension Journal Article In: IEEE Transactions on Neural Systems and Rehabilitation Engineering, vol. 32, pp. 3465–3475, 2024. @article{Zhang2024q, With the recent proliferation of large language models (LLMs), such as Generative Pre-trained Transformers (GPT), there has been a significant shift in exploring human and machine comprehension of semantic language meaning. This shift calls for interdisciplinary research that bridges cognitive science and natural language processing (NLP). This pilot study aims to provide insights into individuals’ neural states during a semantic inference reading-comprehension task. We propose jointly analyzing LLMs, eye-gaze, and electroencephalographic (EEG) data to study how the brain processes words with varying degrees of relevance to a keyword during reading. We also use feature engineering to improve the fixation-related EEG data classification while participants read words with high versus low relevance to the keyword. The best validation accuracy in this word-level classification is over 60% across 12 subjects. Words highly relevant to the inference keyword received significantly more eye fixations per word: 1.0584 compared to 0.6576, including words with no fixations. This study represents the first attempt to classify brain states at a word level using LLM-generated labels. It provides valuable insights into human cognitive abilities and Artificial General Intelligence (AGI), and offers guidance for developing potential reading-assisted technologies. |
Haojun Yang; Xiaojie Wei; Kailing Huang; Zhongling Wu; Qiong Zhang; Shirui Wen; Quan Wang; Li Feng Features of attention network impairment in patients with temporal lobe epilepsy: Evidence from eye-tracking and electroencephalogram Journal Article In: Epilepsy and Behavior, vol. 157, pp. 1–8, 2024. @article{Yang2024, Aim: To explore multiple features of attention impairments in patients with temporal lobe epilepsy (TLE). Methods: A total of 93 patients diagnosed with TLE at Xiangya Hospital during May 2022 and December 2022 and 85 healthy controls were included in this study. Participants were asked to complete neuropsychological scales and attention network test (ANT) with recording of eye-tracking and electroencephalogram. Results: All means of evaluation showed impaired attention functions in TLE patients. ANT results showed impaired orienting (p < 0.001) and executive control (p = 0.041) networks. Longer mean first saccade time (p = 0.046) and more total saccadic counts (p = 0.035) were found in eye-tracking results, indicating abnormal alerting and orienting networks. Both alerting, orienting and executive control networks were abnormal, manifesting as decreased amplitudes (N1 & P3, p < 0.001) and extended latency (P3 |
Hamideh Norouzi; Mohammad Reza Daliri Prediction of behavioral performance by alpha-band phase synchronization in working memory Journal Article In: Physiology and Behavior, vol. 284, pp. 1–11, 2024. @article{Norouzi2024, Working memory (WM) is a cognitive system with limited capacity that can temporarily store and process information. The purpose of this study was to investigate functional connectivity based on phase synchronization during WM and its relationship with the behavioral response. In this regard, we recorded EEG/Eye tracking data of seventeen healthy subjects while performing a memory-guided saccade (MGS) task with two different positions (near eccentricity and far eccentricity). We computed saccade error as memory performance and measured functional connectivity using Phase Locking Value (PLV) in the alpha frequency band (8–12 Hz). The results showed that PLV is negatively correlated with saccade error. Our finding indicated that during the maintenance period, PLV between the frontal and visual area in trials with low saccade error increased significantly compared to trials with high saccade error. Furthermore, we observed a significant difference between PLV for near and far conditions in the delay period. The results suggest that PLV in memory maintenance, in addition to predicting saccade error as behavioral performance, can be related to the coding of spatial information in WM. |
Amir Norouzpour; Tawna L. Roberts Fcirc statistic for steady-state evoked potentials; a generalized version of Tcirc2 statistic Journal Article In: Biomedical Signal Processing and Control, vol. 87, pp. 1–8, 2024. @article{Norouzpour2024, Introduction: Steady-state evoked potentials (ssEP) provide objective tools for studying brain function in different experimental conditions. Frequency components of brain response to repetitive stimuli have been analyzed using Tcirc2 statistic; however, Tcirc2 statistic is limited to comparisons between two means. Here, we present a generalized version of Tcirc2 statistic which enables us to compare multiple means of Fourier estimates corresponding to multiple conditions within participant(s) or multiple groups of participants. Methods: Frequency components of brain response are extracted from ssEP data using Fourier transform. Discrete Fourier measurements at frequency of interest are represented on the complex plane for statistical analyses. We present a new statistic called Fcirc statistic to compare three or more clusters of Fourier estimates whether they have equal or unequal variances or/and numbers of samples. Fcirc statistic derives from Welch's test but for multiple comparisons. Results: We demonstrate the validity of Fcirc statistic using simulated and empirical clusters of Fourier estimates with equal and unequal variances and numbers of samples. Type-I error remains 0.05 for all the conditions. Furthermore, we illustrate that the probability of achieving a significant difference among multiple means when the true means are unequal depends on the total length of ssEP data but is independent of the duration chosen for performing Fourier transform on a fixed length of ssEP data. Conclusion: Fcirc statistic is useful for multiple intra- and inter-participant and group comparisons of brain response at any frequency component extracted from ssEP data whether the group means have equal or unequal variances. |
Stijn A. Nuiten; Jan Willem Gee; Jasper B. Zantvoord; Johannes J. Fahrenfort; Simon Gaal Pharmacological elevation of catecholamine levels improves perceptual decisions, but not metacognitive insight Journal Article In: eNeuro, vol. 11, no. 7, pp. 1–20, 2024. @article{Nuiten2024, Perceptual decisions are often accompanied by a feeling of decision confidence. Where the parietal cortex is known for its crucial role in shaping such perceptual decisions, metacognitive evaluations are thought to additionally rely on the (pre)frontal cortex. Because of this supposed neural differentiation between these processes, perceptual and metacognitive decisions may be divergently affected by changes in internal (e.g., attention, arousal) and external (e.g., task and environmental demands) factors. Although intriguing, causal evidence for this hypothesis remains scarce. Here, we investigated the causal effect of two neuromodulatory systems on behavioral and neural measures of perceptual and metacognitive decision-making. Specifically, we pharmacologically elevated levels of catechol-amines (with atomoxetine) and acetylcholine (with donepezil) in healthy adult human participants performing a visual discrimination task in which we gauged decision confidence, while electroencephalography was measured. Where cholinergic effects were not robust, catecholaminergic enhancement improved perceptual sensitivity, while at the same time leaving metacognitive sensitivity unaffected. Neurally, catecholaminergic elevation did not affect sensory representations of task-relevant visual stimuli but instead enhanced well-known decision signals measured over the centroparietal cortex, reflecting the accumulation of sensory evidence over time. Crucially, catecholaminergic enhancement concurrently impoverished neural markers measured over the frontal cortex linked to the formation of metacognitive evaluations. Enhanced catecholaminergic neuromodulation thus improves perceptual but not metacognitive decision-making. |
Laura Pasqualette; Louisa Kulke Emotional expressions, but not social context, modulate attention during a discrimination task Journal Article In: Cognition and Emotion, pp. 1–19, 2024. @article{Pasqualette2024, Investigating social context effects and emotional modulation of attention in a laboratory setting is challenging. Electroencephalography (EEG) requires a controlled setting to avoid confounds, which goes against the nature of social interaction and emotional processing in real life. To bridge this gap, we developed a new paradigm to investigate the effects of social context and emotional expressions on attention in a laboratory setting. We co-registered eye-tracking and EEG to assess gaze behavior and brain activity while participants performed a discrimination task followed by feedback. Video clips of one second in which a confederate displayed either positive, neutral or negative expressions were presented as feedback to the discrimination task. Participants' belief was manipulated by telling them that the videos were selected either by the computer (non-social condition) or by the experimenter in the adjacent room that observed them via videochat (social condition). We found that emotional expressions modulated late attention processing in the brain (EPN and LPC), but neither early processing (P1) nor saccade latency. Social context did not influence any of the variables studied. We conclude this new paradigm serves as a stepping stone to the development of new paradigms to study social interaction within EEG experiments. |
Laura Pasqualette; Louisa Kulke Differences between overt, covert and natural attention shifts to emotional face Journal Article In: Neuroscience, vol. 559, pp. 283–292, 2024. @article{Pasqualette2024a, In daily life, individuals pay attention to emotional facial expressions and dynamically choose how to shift their attention, i.e. either overtly (with eye-movements) or covertly (without eye-movements). However, research on attention to emotional faces has mostly been conducted in controlled laboratory settings, in which people were instructed where to look. The current preregistered study co-registered EEG and eye-tracking to investigate differences in emotion-driven attention between instructed and uninstructed natural attention shifts in 48 adults. While a central stimulus was presented to the participant, a face appeared in the periphery, showing either a happy, neutral or an angry expression. In three counterbalanced blocks participants were instructed to either move their eyes overtly to the peripheral face, keep fixating the center and therefore covertly shift their attention, or freely look wherever they would like to look. We found that emotional content had stronger effects on the amplitude of the Early Posterior Negativity when participants shifted attention naturally, and that natural shifts of attention differed from instructed shifts in both saccade behavior and neural mechanisms. In summary, our results emphasize the importance of investigating modulation of attention using paradigms that allow par- ticipants to allocate their attention naturally. |
Galina Portnova; Guzal M. Khayrullina; Ivan V. Mikheev; Sofiya M. Byvsheva; Elena V. Proskurnina; Olga Martynova The dynamics of resting-state EEG and salivary trace elements in patients with obsessive-compulsive disorder Journal Article In: ACS Chemical Neuroscience, vol. 15, no. 7, pp. 1415–1423, 2024. @article{Portnova2024, The study of salivary microelements and their neurophysiological and behavioral correlates in patients with obsessive-compulsive disorder (OCD) is a pressing issue in modern psychiatry, which, however, lacks adequate research at this time. In this study, we tested the dynamics of behavioral parameters, resting-state electroencephalogram (EEG), and salivary iron, copper, manganese, magnesium, and zinc in 30 healthy volunteers and 30 individuals with OCD before and after an emotional antisaccade task. The eye-movement data served as a measure of behavioral performance. Our research revealed consistently higher manganese concentrations in the OCD group compared to healthy volunteers associated with a higher EEG ratio of amplitude transformation and symptom severity. The dynamics of salivary microelements and resting-state EEG, possibly influenced by cognitive and emotional load during the anticsaccade task, differed between groups. In healthy volunteers, there was a decrease in salivary iron level with an increase in high-frequency power spectral density of EEG. The OCD group showed a decrease in salivary copper with an increased Hjorth mobility of EEG. |
Jourdan J. Pouliot; Richard T. Ward; Caitlin M. Traiser; Payton Chiasson; Faith E. Gilbert; Andreas Keil Neurophysiological and autonomic dynamics of threat processing during sustained social fear generalization Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 2, pp. 482–497, 2024. @article{Pouliot2024, Survival in rapidly changing environments requires that organisms learn to predict noxious outcomes based on situational cues. One key facet of successful threat prediction is generalization from a specific predictive cue to similar cues, ensuring that a cue-outcome contingency is applied beyond the original learning environment. Generalization has also been observed in laboratory studies of human aversive conditioning: Most behavioral and physiological processes generalize responses from a stimulus paired with threat, (the CS+), to unpaired stimuli, with response magnitudes varying as a function of stimulus similarity. In contrast, work focusing on sensory responses in visual cortex has found a sharpening pattern, in which responses to stimuli closely resembling the CS+ are maximally suppressed, potentially reflecting lateral inhibitory interactions with the CS+ representation. Originally demonstrated with simple visual cues, changes in visuocortical tuning have also been observed in threat generalization learning across facial identity cues. It is however unclear to what extent these visuocortical changes represent transient or sustained effects and if generalization learning requires prior conditioning to the CS+. The present study addressed these questions using EEG and pupillometry in a paradigm involving several hundreds of trials of aversive generalization learning along a gradient of facial identities. Visuocortical ssVEP sharpening occurred after dozens of trials of generalization learning without prior differential conditioning, but diminished as learning progressed further. By contrast, generalization of EEG alpha power suppression, pupil dilation, and self-reported valence and arousal ratings was seen throughout the experimental session. Findings are consistent with models of threat processing emphasizing the role of changing visucocortical and attention dynamics in the formation, curation, and shaping of fear memories as observers continue learning about stimulus-outcome contingencies. |
Michal Pšurný; Stanislav Mokrý; Jana Stavkova Exploring consumers' perceptions of online purchase decision factors: Electroencephalography and eye-tracking evidence Journal Article In: Frontiers in Human Neuroscience, vol. 18, no. November, pp. 1–12, 2024. @article{Psurny2024, Introduction: Consumer behavior on the Internet is influenced by factors that can affect consumers' perceptions and attention to products. Understanding these processes at the neurobiological level can help to understand consumers' implicit responses to marketing stimuli. The objective of this study is to use electroencephalography (EEG) to investigate the differential effects of selected online purchase decision factors that are becoming increasingly important in online shopping. Methods: Using event-related potentials (ERPs) and simultaneous eye-tracking measurements, we identified differences in the perception of utilitarian and hedonic products when the products are exposed together with visual elements of the factors review, discount, and quantity discount. The ERP analysis focused on the P200 and late positive potential components (LPP). Results: By allowing free-viewing of stimuli during measurement, early automatic and later more complex attentional affective responses could be observed. The results suggest that the review and discount factors are processed faster than the product itself. However, the eye-tracking data indicate that the brain processes the factor without looking at it directly, i.e., from a peripheral view. Discussion: The study also demonstrates the possibilities of using new objective methods based on neurobiology and how they can be applied, especially in areas where the use of neuroscience is still rare, yet so much needed to objectify consumers' knowledge of their need satisfaction behavior. |
Zach V. Redding; Ian C. Fiebelkorn Separate cue- and alpha-related mechanisms for distractor suppression Journal Article In: Journal of Neuroscience, vol. 44, no. 25, pp. 1–13, 2024. @article{Redding2024, Research on selective attention has largely focused on the enhancement of behaviorally important information, with less focus on the suppression of distracting information. Enhancement and suppression can operate through a push−pull relationship attributable to competitive interactions among neural populations. There has been considerable debate, however, regarding (1) whether suppression can be voluntarily deployed, independent of enhancement, and (2) whether voluntary deployment of suppression is associated with neural processes occurring prior to the distractor onset. Here, we investigated the interplay between pre- and post-distractor neural processes, while male and female human subjects performed a visual search task with a cue that indicated the location of an upcoming distractor. We utilized two established EEG markers of suppression: the distractor positivity (PD) and alpha power (∼8–15 Hz). The PD—a component of event-related potentials—has been linked with successful distractor suppression, and increased alpha power has been linked with attenuated sensory processing. Cueing the location of an upcoming distractor speeded responses and led to an earlier PD, consistent with earlier suppression due to strategic use of a spatial cue. In comparison, higher predistractor alpha power contralateral to distractors led to a later PD, consistent with later suppression. Lower alpha power contralateral to distractors instead led to distractor-related attentional capture. Lateralization of alpha power was not linked to the spatial cue. This observation, combined with differences in the timing of suppression—as indexed by earlier and later PD components—demonstrates that cue-related, voluntary suppression can occur separate from alpha-related gating of sensory processing. |
Michele Bevilacqua; Sarah Feroldi; Fabienne Windel; Pauline Menoud; Roberto F. Salamanca-Giron; Sarah B. Zandvliet; Lisa Fleury; Friedhelm C. Hummel; Estelle Raffin Single session cross-frequency bifocal tACS modulates visual motion network activity in young healthy population and stroke patients Journal Article In: Brain Stimulation, vol. 17, no. 3, pp. 660–667, 2024. @article{Bevilacqua2024, Background: Phase synchronization over long distances underlies inter-areal communication and importantly, modulates the flow of information processing to adjust to cognitive demands. Objective: This study investigates the impact of single-session, cross-frequency (Alpha-Gamma) bifocal transcranial alternating current stimulation (cf-tACS) to the cortical visual motion network on inter-areal coupling between the primary visual cortex (V1) and the medio-temporal area (MT) and on motion direction discrimination. Methods: Based on the well-established phase-amplitude coupling (PAC) mechanism driving information processing in the visual system, we designed a novel directionally tuned cf-tACS protocol. Directionality of information flow was inferred from the area receiving low-frequency tACS (e.g., V1) projecting onto the area receiving high-frequency tACS (e.g., MT), in this case, promoting bottom-up information flow (Forward-tACS). The control condition promoted the opposite top-down connection (from MT to V1, called Backward-tACS), both compared to a Sham-tACS condition. Task performance and EEG activity were recorded from 45 young healthy subjects. An additional cohort of 16 stroke patients with occipital lesions and impairing visual processing was measured to assess the influence of a V1 lesion on the modulation of V1-MT coupling. Results: The results indicate that Forward cf-tACS successfully modulated bottom-up PAC (V1 α-phase-MT ɣ-amplitude) in both cohorts, while producing opposite effects on the reverse MT-to-V1 connection. Backward-tACS did not change V1-MT PAC in either direction in healthy participants but induced a slight decrease in bottom-up PAC in stroke patients. However, these changes in inter-areal coupling did not translate into cf-tACS-specific behavioural improvements. Conclusions: Single session cf-tACS can alter inter-areal coupling in intact and lesioned brains but is probably not enough to induce longer-lasting behavioural effects in these cohorts. This might suggest that a longer daily visual training protocol paired with tACS is needed to unveil the relationship between externally applied oscillatory activity and behaviourally relevant brain processing. |
Lénaïc Borot; Ruth Ogden; Simon J. Bennett Prefrontal cortex activity and functional organisation in dual-task ocular pursuit is affected by concurrent upper limb movement Journal Article In: Scientific Reports, vol. 14, no. 1, pp. 1–12, 2024. @article{Borot2024, Tracking a moving object with the eyes seems like a simple task but involves areas of prefrontal cortex (PFC) associated with attention, working memory and prediction. Increasing the demand on these processes with secondary tasks can affect eye movements and/or perceptual judgments. This is particularly evident in chronic or acute neurological conditions such as Alzheimer's disease or mild traumatic brain injury. Here, we combined near infrared spectroscopy and video-oculography to examine the effects of concurrent upper limb movement, which provides additional afference and efference that facilitates tracking of a moving object, in a novel dual-task pursuit protocol. We confirmed the expected effects on judgement accuracy in the primary and secondary tasks, as well as a reduction in eye velocity when the moving object was occluded. Although there was limited evidence of oculo-manual facilitation on behavioural measures, performing concurrent upper limb movement did result in lower activity in left medial PFC, as well as a change in PFC network organisation, which was shown by Graph analysis to be locally and globally more efficient. These findings extend upon previous work by showing how PFC is functionally organised to support eye-hand coordination when task demands more closely replicate daily activities. |
Ethan S. Bromberg-Martin; Yang-Yang Feng; Takaya Ogasawara; J. Kael White; Kaining Zhang; Ilya E. Monosov A neural mechanism for conserved value computations integrating information and rewards Journal Article In: Nature Neuroscience, vol. 27, pp. 1–17, 2024. @article{BrombergMartin2024, Behavioral and economic theory dictate that we decide between options based on their values. However, humans and animals eagerly seek information about uncertain future rewards, even when this does not provide any objective value. This implies that decisions are made by endowing information with subjective value and integrating it with the value of extrinsic rewards, but the mechanism is unknown. Here, we show that human and monkey value judgements obey strikingly conserved computational principles during multi-attribute decisions trading off information and extrinsic reward. We then identify a neural substrate in a highly conserved ancient structure, the lateral habenula (LHb). LHb neurons signal subjective value, integrating information's value with extrinsic rewards, and the LHb predicts and causally influences ongoing decisions. Neurons in key input areas to the LHb largely signal components of these computations, not integrated value signals. Thus, our data uncover neural mechanisms of conserved computations underlying decisions to seek information about the future. |
Jeroen Brus; Joseph A. Heng; Valeriia Beliaeva; Fabian Gonzalez Pinto; Antonino Mario Cassarà; Esra Neufeld; Marcus Grueschow; Lukas Imbach; Rafael Polanía Causal phase-dependent control of non-spatial attention in human prefrontal cortex Journal Article In: Nature Human Behaviour, vol. 8, no. 4, pp. 743–757, 2024. @article{Brus2024, Non-spatial attention is a fundamental cognitive mechanism that allows organisms to orient the focus of conscious awareness towards sensory information that is relevant to a behavioural goal while shifting it away from irrelevant stimuli. It has been suggested that attention is regulated by the ongoing phase of slow excitability fluctuations of neural activity in the prefrontal cortex, a hypothesis that has been challenged with no consensus. Here we developed a behavioural and non-invasive stimulation paradigm aiming at modulating slow excitability fluctuations of the inferior frontal junction. Using this approach, we show that non-spatial attention can be selectively modulated as a function of the ongoing phase of exogenously modulated excitability states of this brain structure. These results demonstrate that non-spatial attention relies on ongoing prefrontal excitability states, which are probably regulated by slow oscillatory dynamics, that orchestrate goal-oriented behaviour. |
Yi-Hsuan Hsuan Chang; He-Jun Jun Chen; Cesar Barquero; Hsu Jung Tsai; Wei-Kuang Kuang Liang; Chun-Hsien Hsien Hsu; Neil G. Muggleton; Chin-An An Wang Linking tonic and phasic pupil responses to P300 amplitude in an emotional face-word Stroop task Journal Article In: Psychophysiology, vol. 61, no. 4, pp. 1–26, 2024. @article{Chang2024b, The locus coeruleus-norepinephrine (LC-NE) system, which regulates arousal levels, is important for cognitive control, including emotional conflict resolution. Additionally, the LC-NE system is implicated in P300 generation. If the P300 is mediated by the LC-NE system, and considering the established correlations between LC activity and pupil dilation, P300 amplitude should correlate with task-evoked (phasic) pupil dilation on a trial-by-trial basis. However, prior studies, predominantly utilizing oddball-type paradigms, have not demonstrated correlations between concurrently recorded task-evoked pupil dilation and P300 responses. Using a recently developed emotional face-word Stroop task that links pupil dilation to the LC-NE system, here, we examined both intra- and inter-individual correlations between task-evoked pupil dilation and P300 amplitude. We found that lower accuracy, slower reaction times, and larger task-evoked pupil dilation were obtained in the incongruent compared to the congruent condition. Furthermore, we observed intra-individual correlations between task-evoked pupil dilation and P300 amplitude, with larger pupil dilation correlating with a greater P300 amplitude. In contrast, pupil dilation did not exhibit consistent correlations with N450 and N170 amplitudes. Baseline (tonic) pupil size also showed correlations with P300 and N170 amplitudes, with smaller pupil size corresponding to larger amplitude. Moreover, inter-individual differences in task-evoked pupil dilation between the congruent and incongruent conditions correlated with differences in reaction time and P300 amplitude, though these effects only approached significance. To summarize, our study provides evidence for a connection between task-evoked pupil dilation and P300 amplitude at the single-trial level, suggesting the involvement of the LC-NE system in P300 generation. |
Lixiang Chen; Radoslaw Martin Cichy; Daniel Kaiser Coherent categorical information triggers integration-related alpha dynamics Journal Article In: Journal of Neurophysiology, vol. 131, no. 4, pp. 619–625, 2024. @article{Chen2024c, To create coherent visual experiences, the brain spatially integrates the complex and dynamic information it receives from the environment. We previously demonstrated that feedback-related alpha activity carries stimulus-specific information when two spatially and temporally coherent naturalistic inputs can be integrated into a unified percept. In this study, we sought to determine whether such integration-related alpha dynamics are triggered by categorical coherence in visual inputs. In an EEG experiment, we manipulated the degree of coherence by presenting pairs of videos from the same or different categories through two apertures in the left and right visual hemifields. Critically, video pairs could be video-level coherent (i.e., stem from the same video), coherent in their basic-level category, coherent in their superordinate category, or incoherent (i.e., stem from videos from two entirely different categories). We conducted multivariate classification analyses on rhythmic EEG responses to decode between the video stimuli in each condition. As the key result, we significantly decoded the video-level coherent and basic-level coherent stimuli, but not the superordinate coherent and incoherent stimuli, from cortical alpha rhythms. This suggests that alpha dynamics play a critical role in integrating information across space, and that cortical integration processes are flexible enough to accommodate information from different exemplars of the same basic-level category. |
Shuyuan Chen; Erik D. Reichle; Yanping Liu Direct lexical control of eye movements in Chinese reading: Evidence from the co-registration of EEG and eye tracking Journal Article In: Cognitive Psychology, vol. 153, no. 135, pp. 1–21, 2024. @article{Chen2024f, The direct-lexical-control hypothesis stipulates that some aspect of a word's processing determines the duration of the fixation on that word and/or the next. Although the direct lexical control is incorporated into most current models of eye-movement control in reading, the precise implementation varies and the assumptions of the hypothesis may not be feasible given that lexical processing must occur rapidly enough to influence fixation durations. Conclusive empirical evidence supporting this hypothesis is therefore lacking. In this article, we report the results of an eye-tracking experiment using the boundary paradigm in which native speakers of Chinese read sentences in which target words were either high- or low-frequency and preceded by a valid or invalid preview. Eye movements were co-registered with electroencephalography, allowing standard analyses of eye-movement measures, divergence point analyses of fixation-duration distributions, and fixated-related potentials on the target words. These analyses collectively provide strong behavioral and neural evidence of early lexical processing and thus strong support for the direct-lexical-control hypothesis. We discuss the implications of the findings for our understanding of how the hypothesis might be implemented, the neural systems that support skilled reading, and the nature of eye-movement control in the reading of Chinese versus alphabetic scripts. |
Hsin-Hua Chin; Ying-Hsuan Tai; Rachel Yep; Yi-Hsuan Chang; Chun-Hsien Hsu; Chin-An Wang Investigating causal effects of pupil size on visual discrimination and visually evoked potentials in an optotype discrimination task Journal Article In: Frontiers in Neuroscience, vol. 18, pp. 1–10, 2024. @article{Chin2024, Pupil size primarily changes to regulate the amount of light entering the retina, optimizing the balance between visual acuity and sensitivity for effective visual processing. However, research directly examining the relationship between pupil size and visual processing has been limited. While a few studies have recorded pupil size and EEG signals to investigate the role of pupil size in visual processing, these studies have predominantly focused on the domain of visual sensitivity. Causal effects of pupil size on visual acuity, therefore, remain poorly understood. By manipulating peripheral background luminance levels and target stimulus contrast while simultaneously recording pupillometry and EEG signals, we examined how absolute pupil size affects visual discrimination and visually evoked potentials (VEP) in a task using optotype mimicking the Snellen eye chart, the most common assessment of visual acuity. Our findings indicate that both higher background luminance levels and higher target contrast were associated with improved target discrimination and faster correct reaction times. Moreover, while higher contrast visual stimuli evoked larger VEPs, the effects of pupil size on VEPs were not significant. Additionally, we did not observe inter-individual correlations between absolute pupil size and discrimination performance or VEP amplitude. Together, our results demonstrate that absolute pupil size, regulated by global luminance level, played a functional role in enhancing visual discrimination performance in an optotype discrimination task. The differential VEP effects of pupil size compared to those of stimulus contrast further suggested distinct neural mechanisms involved in facilitating visual acuity under small pupils. |
Samson Chota; Arnaud T. Bruat; Stefan Van der Stigchel; Christoph Strauch In: Journal of Cognitive Neuroscience, vol. 36, no. 5, pp. 800–814, 2024. @article{Chota2024, Visual working memory (VWM) allows storing goal-relevant information to guide future behavior. Prior work suggests that VWM is spatially organized and relies on spatial attention directed toward locations at which memory items were encoded, even if location is task-irrelevant. Importantly, attention often needs to be dynamically redistributed between locations, for example, in preparation for an upcoming probe. Very little is known about how attentional resources are distributed between multiple locations during a VWM task and even less about the dynamic changes govern-ing such attentional shifts over time. This is largely due to the inability to use behavioral outcomes to reveal fast dynamic changes within trials. We here demonstrated that EEG steady-state visual evoked potentials (SSVEPs) successfully track the dynamic allocation of spatial attention during a VWM task. Participants were presented with to-be-memorized gratings and distractors at two distinct locations, tagged with flickering discs. This allowed us to dynamically track attention allocated to memory and distractor items via their coupling with space by quantifying the amplitude and coherence of SSVEP responses in the EEG signal to flickering stimuli at the former memory and distractor locations. SSVEP responses did not differ between memory and distractor locations during early maintenance. However, shortly before probe comparison, we observed a decrease in SSVEP coherence over distractor locations indicative of a reallocation of spatial attentional resources. RTs were shorter when preceded by stronger decreases in SSVEP coherence at distractor locations, likely reflecting attentional shifts from the distractor to the probe or memory location. We demonstrate that SSVEPs can inform about dynamic processes in VWM, even if location does not have to be reported by participants. This finding not only supports the notion of a spatially organized VWM but also reveals that SSVEPs betray a dynamic prioritization process of working memory items and locations over time that is directly predictive of memory performance. |
Thérèse Collins; Emilie Zhu; Patrick Rateau The neural representation of stereotype content Journal Article In: Scientific Reports, vol. 14, no. 1, pp. 1–12, 2024. @article{Collins2024, Judgments about social groups are characterized by their position in a representational space defined by two axes, warmth and competence. We examined serial dependence (SD) in evaluations of warmth and competence while measuring participants' electroencephalographic (EEG) activity, as a means to address the independence between these two psychological axes. SD is the attraction of perceptual reports towards things seen in the recent past and has recently been intensely investigated in vision. SD occurs at multiple levels of visual processing, from basic features to meaningful objects. The current study aims to (1) measure whether SD occurs between non-visual objects, in particular social groups and (2) uncover the neural correlates of social group evaluation and SD using EEG. Participants' judgments about social groups such as “nurses” or “accountants” were serially dependent, but only when the two successive groups were close in representational space. The pattern of results argues in favor of a non-separability between the two axes, because groups nearby on one dimension but far on the other were not subject to SD, even though that other dimension was irrelevant to the task at hand. Using representational similarity analysis, we found a brain signature that differentiated social groups as a function of their position in the representational space. Our results thus argue that SD may be a ubiquitous cognitive phenomenon, that social evaluations are serially dependent, and that reproducible neural signatures of social evaluations can be uncovered. |
Aritra Das; Nilanjana Nandi; Supratim Ray Alpha and SSVEP power outperform gamma power in capturing attentional modulation in human EEG Journal Article In: Cerebral Cortex, vol. 34, no. 1, pp. 1–16, 2024. @article{Das2024, Attention typically reduces power in the alpha (8–12 Hz) band and increases power in gamma (>30 Hz) band in brain signals, as reported in macaque local field potential (LFP) and human electro/magneto-encephalogram (EEG/MEG) studies. In addition, EEG studies often use flickering stimuli that produce a specific measure called steady-state-visually-evoked-potential (SSVEP), whose power also increases with attention. However, effectiveness of these neural measures in capturing attentional modulation is unknown since stimuli and task paradigms vary widely across studies. In a recent macaque study, attentional modulation was more salient in the gamma band of the LFP, compared to alpha or SSVEP. To compare this with human EEG, we designed an orientation change detection task where we presented both static and counterphasing stimuli of matched difficulty levels to 26 subjects and compared attentional modulation of various measures under similar conditions. We report two main results. First, attentional modulation was comparable for SSVEP and alpha. Second, non-foveal stimuli produced weak gamma despite various stimulus optimizations and showed negligible attentional modulation although full-screen gratings showed robust gamma activity. Our results are useful for brain-machine-interfacing studies where suitable features are used for decoding attention, and also provide clues about spatial scales of neural mechanisms underlying attention. |
Yun Ding; Bradley R. Postle; Freek Ede Neural signatures of competition between voluntary and involuntary influences over the focus of attention in visual working memory Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 5, pp. 815–827, 2024. @article{Ding2024, Adaptive behavior relies on the selection and prioritization of relevant sensory inputs from the external environment as well as from among internal sensory representations held in working memory. Recent behavioral evidence suggests that the classic distinction between voluntary (goal-driven) and involuntary (stimulus-driven) influences over attentional allocation also applies to the selection of internal representations held in working memory. In the current EEG study, we set out to investigate the neural dynamics associated with the competition between voluntary and involuntary control over the focus of attention in visual working memory. We show that when voluntary and involuntary factors compete for the internal focus of attention, prioritization of the appropriate item is delayed—as reflected both in delayed gaze biases that track internal selection and in delayed neural beta (15–25 Hz) dynamics that track the planning for the upcoming memory-guided manual action. We further show how this competition is paralleled—possibly resolved—by an increase in frontal midline theta (4–8 Hz) activity that, moreover, predicts the speed of ensuing memory-guided behavior. Finally, because theta increased following retrocues that effectively reduced working-memory load, our data unveil how frontal theta activity during internal attentional focusing tracks demands on cognitive control over and above working-memory load. Together, these data yield new insight into the neural dynamics that govern the focus of attention in visual working memory, and disentangle the contributions of frontal midline theta activity to the processes of control versus retention in working memory. |
Carola Dolci; Einat Rashal; Elisa Santandrea; Suliann Ben; Leonardo Chelazzi; Emiliano Macaluso; C. Nico Boehler; Suliann Ben Hamed; Leonardo Chelazzi; Emiliano Macaluso; C. Nico Boehler The dynamics of statistical learning in visual search and its interaction with salience processing: An EEG study Journal Article In: NeuroImage, vol. 286, pp. 1–12, 2024. @article{Dolci2024, Visual attention can be guided by statistical regularities in the environment, that people implicitly learn from past experiences (statistical learning, SL). Moreover, a perceptually salient element can automatically capture attention, gaining processing priority through a bottom-up attentional control mechanism. The aim of our study was to investigate the dynamics of SL and if it shapes attentional target selection additively with salience processing, or whether these mechanisms interact, e.g. one gates the other. In a visual search task, we therefore manipulated target frequency (high vs. low) across locations while, in some trials, the target was salient in terms of colour. Additionally, halfway through the experiment, the high-frequency location changed to the opposite hemifield. EEG activity was simultaneously recorded, with a specific interest in two markers related to target selection and post-selection processing, respectively: N2pc and SPCN. Our results revealed that both SL and saliency significantly enhanced behavioural performance, but also interacted with each other, with an attenuated saliency effect at the high-frequency target location, and a smaller SL effect for salient targets. Concerning processing dynamics, the benefit of salience processing was more evident during the early stage of target selection and processing, as indexed by a larger N2pc and early-SPCN, whereas SL modulated the underlying neural activity particularly later on, as revealed by larger late-SPCN. Furthermore, we showed that SL was rapidly acquired and adjusted when the spatial imbalance changed. Overall, our findings suggest that SL is flexible to changes and, combined with salience processing, jointly contributes to establishing attentional priority. |
Laura Doll; Andrew R. Dykstra; Alexander Gutschalk Perceptual awareness of near-threshold tones scales gradually with auditory cortex activity and pupil dilation Journal Article In: iScience, vol. 27, no. 8, pp. 1–20, 2024. @article{Doll2024, Negative-going responses in sensory cortex co-vary with perceptual awareness of sensory stimuli. Given that this awareness negativity has also been observed for undetected stimuli, some have challenged its role for perception. To address this question, we combined magnetoencephalography, electroencephalography, and pupillometry to study how sustained attention and response criterion affect the auditory awareness negativity. Participants first detected distractor sounds and denied hearing task-irrelevant near-threshold tones, which evoked neither awareness negativity nor pupil dilation. These same tones evoked both responses when task-relevant, stronger for hit but also present for miss trials. Participants then rated their perception on a six-point scale to test whether response criterion explains the presence of these responses for miss trials. Decreasing perception ratings were associated with gradually reduced evoked responses, consistent with signal detection theory. These results support the concept of an awareness negativity that is modulated by attention but does not require a non-linear threshold mechanism. |
Y B Eisma; S T Vliet; A J Nederveen; J C F Winter Assessing the influence of visual stimulus properties on steady-state visually evoked potentials and pupil diameter Journal Article In: Biomedical Physics & Engineering Express, vol. 1o, pp. 1–12, 2024. @article{Eisma2024, Steady-State Visual Evoked Potentials (SSVEPs) are brain responses measurable via electroencephalo- graphy (EEG) in response to continuous visual stimulation at a constant frequency. SSVEPs have been instrumental in advancing our understanding ofhuman vision and attention, as well as in the development ofbrain-computer interfaces (BCIs). Ongoing questions remain about which type of visual stimulus causes the most potent SSVEP response. The current study investigated the effects of color, size, and flicker frequencyon the signal-to-noise ratio ofSSVEPs, complemented bypupillary light reflex measurements obtained through an eye-tracker. Six participants were presented with visual stimuli that differed in terms ofcolor (white, red, green), shape (circles, squares, triangles), size (10,000 to 30,000 pixels), flicker frequency (8to25Hz), and grouping (one stimulus at a time versus four stimuli presented in a 2×2 matrix to simulate a BCI). The results indicated that larger stimuli elicited stronger SSVEP responses and more pronounced pupil constriction. Additionally, the results revealed an interaction between stimulus color and flicker frequency, with red being more effective at lower frequencies and white at higher frequencies. Future SSVEP research could focus on the recommended waveform, interactions between SSVEP and power grid frequency, a wider range of flicker frequencies, a larger sample ofparticipants, and a systematic comparison ofthe information transfer obtained through SSVEPs, pupil diameter, and eye movements. |
Hadeel Ershaid; Mikel Lizarazu; D. J. Drew McLaughlin; Martin Cooke; Olympia Simantiraki; Maria Koutsogiannaki; Marie Lallier Contributions of listening effort and intelligibility to cortical tracking of speech in adverse listening conditions Journal Article In: Cortex, vol. 172, pp. 54–71, 2024. @article{Ershaid2024, Cortical tracking of speech is vital for speech segmentation and is linked to speech intelligibility. However, there is no clear consensus as to whether reduced intelligibility leads to a decrease or an increase in cortical speech tracking, warranting further investigation of the factors influencing this relationship. One such factor is listening effort, defined as the cognitive resources necessary for speech comprehension, and reported to have a strong negative correlation with speech intelligibility. Yet, no studies have examined the relationship between speech intelligibility, listening effort, and cortical tracking of speech. The aim of the present study was thus to examine these factors in quiet and distinct adverse listening conditions. Forty-nine normal hearing adults listened to sentences produced casually, presented in quiet and two adverse listening conditions: cafeteria noise and re- verberant speech. Electrophysiological responses were registered with electroencephalogram, and listening effort was estimated subjectively using self-reported scores and objectively using pupillometry. Results indicated varying impacts of adverse conditions on intelligibility, listening effort, and cortical tracking of speech, depending on the preservation of the speech temporal envelope. The more distorted envelope in the reverberant condition led to higher listening effort, as reflected in higher subjective scores, increased pupil diameter, and stronger cortical tracking of speech in the delta band. These findings suggest that using measures of listening effort in addition to those of intelligibility is useful for interpreting cortical tracking of speech results. Moreover, reading and phonological skills of participants were positively correlated with listening effort in the cafeteria condition, suggesting a special role of expert language skills in processing speech in this noisy condition. Implications for future research and theories linking atypical cortical tracking of speech and reading disorders are further discussed. |
Camille Fakche; Laura Dugué Perceptual cycles travel across retinotopic space Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 1, pp. 200–216, 2024. @article{Fakche2024, Visual perception waxes and wanes periodically over time at low frequencies (theta: 4–7 Hz; alpha: 8–13 Hz), creating “per-ceptual cycles.” These perceptual cycles can be induced when stimulating the brain with a flickering visual stimulus at the theta or alpha frequency. Here, we took advantage of the well-known organization of the visual system into retinotopic maps (topographic correspondence between visual and cortical spaces) to assess the spatial organization of induced perceptual cycles. Specifically, we tested the hypothesis that they can propagate across the retinotopic space. A disk oscillating in luminance (inducer) at 4, 6, 8, or 10 Hz was presented in the periphery of the visual field to induce perceptual cycles at specific frequencies. EEG recordings verified that the brain responded at the corresponding inducer frequencies and their first harmonics. Perceptual cycles were assessed with a concurrent detection task—target stimuli were displayed at threshold contrast (50% detection) at random times during the inducer. Behavioral results confirmed that perceptual performance was modulated periodically by the inducer at each frequency. We additionally manipulated the distance between the target and the inducer (three possible positions) and showed that the optimal phase, that is, moment of highest target detection, shifted across target distance to the inducer, specifically when its flicker frequency was in the alpha range (8 and 10 Hz). These results demonstrate that induced alpha perceptual cycles travel across the retinotopic space in humans at a propagation speed of 0.3–0.5 m/sec, consistent with the speed of unmyelinated horizontal connections in the visual cortex. |
Zepeng Fang; Yuanyuan Dang; Zhipei Ling; Yongzheng Han; Hulin Zhao; Xin Xu; Mingsha Zhang The involvement of the human prefrontal cortex in the emergence of visual awareness Journal Article In: eLife, vol. 13, pp. 1–25, 2024. @article{Fang2024, Exploring the neural mechanisms of awareness is a fundamental task of cognitive neuroscience. There is an ongoing dispute regarding the role of the prefrontal cortex (PFC) in the emergence of awareness, which is partially raised by the confound between report-and awareness-related activity. To address this problem, we designed a visual awareness task that can minimize report-related motor confounding. Our results show that saccadic latency is significantly shorter in the aware trials than in the unaware trials. Local field potential (LFP) data from six patients consis-tently show early (200–300ms) awareness-related activity in the PFC, including event-related potential and high-gamma activity. Moreover, the awareness state can be reliably decoded by the neural activity in the PFC since the early stage, and the neural pattern is dynamically changed rather than being stable during the representation of awareness. Furthermore, the enhancement of dynamic functional connectivity, through the phase modulation at low frequency, between the PFC and other brain regions in the early stage of the awareness trials may explain the mechanism of conscious access. These results indicate that the PFC is critically involved in the emergence of awareness. |
Andrew H. Farkas; Richard T. Ward; Faith E. Gilbert; Jourdan Pouliot; Payton Chiasson; Skylar Mcilvanie; Caitlin Traiser; Kierstin Riels; Ryan Mears; Andreas Keil Auditory aversive generalization learning prompts threat-specific changes in alpha-band activity Journal Article In: Cerebral Cortex, vol. 34, no. 3, pp. 1–11, 2024. @article{Farkas2024, Pairing a neutral stimulus with aversive outcomes prompts neurophysiological and autonomic changes in response to the conditioned stimulus (CS+), compared to cues that signal safety (CS-). One of these changes - selective amplitude reduction of parietal alpha-band oscillations - has been reliably linked to processing of visual CS+. It is, however, unclear to what extent auditory conditioned cues prompt similar changes, how these changes evolve as learning progresses, and how alpha reduction in the auditory domain generalizes to similar stimuli. To address these questions, 55 participants listened to three sine wave tones, with either the highest or lowest pitch (CS+) being associated with a noxious white noise burst. A threat-specific (CS+) reduction in occipital-parietal alpha-band power was observed similar to changes expected for visual stimuli. No evidence for aversive generalization to the tone most similar to the CS+ was observed in terms of alpha-band power changes, aversiveness ratings, or pupil dilation. By-trial analyses found that selective alpha-band changes continued to increase as aversive conditioning continued, beyond when participants reported awareness of the contingencies. The results support a theoretical model in which selective alpha power represents a cross-modal index of continuous aversive learning, accompanied by sustained sensory discrimination of conditioned threat from safety cues. |
Hsing-Hao Lee; Antonio Fernandez; Marisa Carrasco Adaptation and exogenous attention interact in the early visual cortex : A TMS study Journal Article In: iScience, vol. 27, pp. 1–12, 2024. @article{Lee2024c, Transcranial magnetic stimulation (TMS) to early visual cortex modulates the effect of adaptation and eliminates the effect of exogenous (involuntary) attention on contrast sensitivity. Here, we investigated whether adaptation modulates exogenous attention under TMS to V1/V2. Observers performed an orientation discrimination task while attending to one of two stimuli, with or without adaptation. Following an attentional cue, two stimuli were presented in the stimulated region and its contralateral symmetric re- gion. A response cue indicated the stimulus whose orientation observers had to discriminate. Without adaptation, in the distractor-stimulated condition, contrast sensitivity increased at the attended location and decreased at the unattended location via response gain—but these effects were eliminated in the target-stimulated condition. Critically, after adaptation, exogenous attention altered performance similarly in both distractor-stimulated and target-stimulated conditions. These results reveal that (1) adaptation and attention interact in the early visual cortex, and (2) adaptation shields exogenous attention from TMS effects. |
Shira Baror; Thomas J. Baumgarten; Biyu J. He Neural mechanisms determining the duration of task-free, self-paced visual perception Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 5, pp. 756–775, 2024. @article{Baror2024, Humans spend hours each day spontaneously engaging with visual content, free from specific tasks and at their own pace. Currently, the brain mechanisms determining the duration of self-paced perceptual behavior remain largely unknown. Here, participants viewed naturalistic images under task-free settings and self-paced each image's viewing duration while undergoing EEG and pupillometry recordings. Across two independent data sets, we observed large inter-and intra-individual variability in viewing duration. However, beyond an image's presentation order and category, specific image content had no consistent effects on spontaneous viewing duration across participants. Overall, longer viewing durations were associated with sustained enhanced posterior positivity and anterior negativity in the ERPs. Individual-specific variations in the spontaneous viewing duration were consistently correlated with evoked EEG activity amplitudes and pupil size changes. By contrast, presentation order was selectively correlated with baseline alpha power and baseline pupil size. Critically, spontaneous viewing duration was strongly predicted by the temporal stability in neural activity patterns starting as early as 350 msec after image onset, suggesting that early neural stability is a key predictor for sustained perceptual engagement. Interestingly, neither bottom–up nor top–down predictions about image category influenced spontaneous viewing duration. Overall, these results suggest that individual-specific factors can influence perceptual processing at a surprisingly early time point and influence the multifaceted ebb and flow of spontaneous human perceptual behavior in naturalistic settings. |
Katarina Begus; Elizabeth Bonawitz Infants evaluate informativeness of evidence and predict causal events as revealed in theta oscillations and predictive looking Journal Article In: Communications Psychology, vol. 2, no. 1, pp. 1–9, 2024. @article{Begus2024, This study investigates 16-month-old infants' sensitivity to the informativeness of evidence and its potential link to infants' ability to draw accurate causal inferences and predict unfolding events. Employing concurrent EEG and eye tracking, data from 66 infants revealed significantly increased theta oscillatory activity when infants expected to see causally unconfounded evidence compared to confounded evidence, suggesting heightened cognitive engagement in anticipation of informative evidence. Crucially, this difference was more pronounced in the subset of infants who later made correct predictions, suggesting that they had correctly inferred the causal structure based on the evidence presented. This research sheds light on infants' motivation to seek explanatory causal information, suggesting that even at 16 months, infants can strategically direct attention to situations conducive to acquiring informative evidence, potentially laying the groundwork for the impressive abilities of humans to rapidly acquire knowledge and develop causal theories of the world. Living |
Eser Sendesen; Didem Turkyilmaz Investigation of the behavior of tinnitus patients under varying listening conditions with simultaneous electroencephalography and pupillometry Journal Article In: Brain and Behavior, vol. 14, no. 6, pp. 1–11, 2024. @article{Sendesen2024a, Objective: This study aims to control all hearing thresholds, including extended high frequencies (EHFs), presents stimuli of varying difficulty levels, and measures electroencephalography (EEG) and pupillometry responses to determine whether listening difficulty in tinnitus patients is effort or fatigue-related. Methods: Twenty-one chronic tinnitus patients and 26 matched healthy controls having normal pure-tone averages with symmetrical hearing thresholds were included. Subjects were evaluated with 0.125−20 kHz pure-tone audiometry, Montreal Cognitive Assessment Test (MoCA), Tinnitus Handicap Inventory (THI), EEG, and pupillometry. Results: Pupil dilatation and EEG alpha power during the “encoding” phase of the presented sentence in tinnitus patients were less in all listening conditions (p <.05). Also, there was no statistically significant relationship between EEG and pupillometry components for all listening conditions and THI or MoCA (p >.05). Conclusion: EEG and pupillometry results under various listening conditions indicate potential listening effort in tinnitus patients even if all frequencies, including EHFs, are controlled. Also, we suggest that pupillometry should be interpreted with caution in autonomic nervous system-related conditions such as tinnitus. |
Dongping Shi; Qing Yu Distinct neural signatures underlying information maintenance and manipulation in working memory Journal Article In: Cerebral Cortex, vol. 34, no. 3, pp. 1–14, 2024. @article{Shi2024a, Previous working memory research has demonstrated robust stimulus representations during memory maintenance in both voltage and alpha-band activity in electroencephalography. However, the exact functions of these 2 neural signatures have remained controversial. Here we systematically investigated their respective contributions to memory manipulation. Human participants either maintained a previously seen spatial location, or manipulated the location following a mental rotation cue over a delay. Using multivariate decoding, we observed robust location representations in low-frequency voltage and alpha-band oscillatory activity with distinct spatiotemporal dynamics: location representations were most evident in posterior channels in alpha-band activity, but were most prominent in the more anterior, central channels in voltage signals. Moreover, the temporal emergence of manipulated representation in central voltage preceded that in posterior alpha-band activity, suggesting that voltage might carry stimulus-specific source signals originated internally from anterior cortex, whereas alpha-band activity might ref lect feedback signals in posterior cortex received from higher-order cortex. Lastly, while location representations in both signals were coded in a low-dimensional neural subspace, location representation in central voltage was higher-dimensional and underwent a representational transformation that exclusively predicted memory behavior. Together, these results highlight the crucial role of central voltage in working memory, and support functional distinctions between voltage and alpha-band activity. |
James Siklos-Whillans; Roxane J. Itier Effects of inversion and fixation location on the processing of face and house stimuli – a mass univariate analysis Journal Article In: Brain Topography, vol. 37, pp. 972–992, 2024. @article{SiklosWhillans2024, Most Event Related Potential studies investigating the time course of visual processing have focused mainly on the N170 component. Stimulus orientation affects the N170 amplitude for faces but not for objects, a finding interpreted as reflecting holistic/configural processing for faces and featural processing for objects. Furthermore, while recent studies suggest where on the face people fixate impacts the N170, fixation location effects have not been investigated in objects. A data-driven mass univariate analysis (all time points and electrodes) was used to investigate the time course of inversion and fixation location effects on the neural processing of faces and houses. Strong and widespread orientation effects were found for both faces and houses, from 100-350ms post-stimulus onset, including P1 and N170 components, and later, a finding arguing against a lack of holistic processing for houses. While no clear fixation effect was found for houses, fixation location strongly impacted face processing early, reflecting retinotopic mapping around the C2 and P1 components, and during the N170-P2 interval. Face inversion effects were also largest for nasion fixation around 120ms. The results support the view that facial feature integration (1) depends on which feature is being fixated and where the other features are situated in the visual field, (2) occurs maximally during the P1-N170 interval when fixation is on the nasion and (3) continues past 200ms, suggesting the N170 peak, where weak effects were found, might be an inflexion point between processes rather than the end of a feature integration into a whole process. |
Jack W. Silcox; Karen Bennett; Allyson Copeland; Sarah Hargus Ferguson; Brennan R. Payne The costs (and benefits?) of effortful listening for older adults: Insights from simultaneous electrophysiology, pupillometry, and memory Journal Article In: Journal of Cognitive Neuroscience, vol. 36, no. 6, pp. 997–1020, 2024. @article{Silcox2024, Although the impact of acoustic challenge on speech processing and memory increases as a person ages, older adults may engage in strategies that help them compensate for these demands. In the current preregistered study, older adults (n = 48) listened to sentences—presented in quiet or in noise—that were high constraint with either expected or unexpected endings or were low constraint with unexpected endings. Pupillometry and EEG were simultaneously recorded, and subsequent sentence recognition and word recall were measured. Like young adults in prior work, we found that noise led to increases in pupil size, delayed and reduced ERP responses, and decreased recall for unexpected words. However, in contrast to prior work in young adults where a larger pupillary response predicted a recovery of the N400 at the cost of poorer memory performance in noise, older adults did not show an associated recovery of the N400 despite decreased memory performance. Instead, we found that in quiet, increases in pupil size were associated with delays in N400 onset latencies and increased recognition memory performance. In conclusion, we found that transient variation in pupil-linked arousal predicted trade-offs between real-time lexical processing and memory that emerged at lower levels of task demand in aging. Moreover, with increased acoustic challenge, older adults still exhibited costs associated with transient increases in arousal without the corresponding benefits. |
Connor Spiech; Anne Danielsen; Bruno Laeng; Tor Endestad Oscillatory attention in groove Journal Article In: Cortex, vol. 174, pp. 137–148, 2024. @article{Spiech2024, Attention is not constant but rather fluctuates over time and these attentional fluctuations may prioritize the processing of certain events over others. In music listening, the pleasurable urge to move to music (termed ‘groove' by music psychologists) offers a particularly convenient case study of oscillatory attention because it engenders synchronous and oscillatory movements which also vary predictably with stimulus complexity. In this study, we simultaneously recorded pupillometry and scalp electroencephalography (EEG) from participants while they listened to drumbeats of varying complexity that they rated in terms of groove afterwards. Using the intertrial phase coherence of the beat frequency, we found that while subjects were listening, their pupil activity became entrained to the beat of the drumbeats and this entrained attention persisted in the EEG even as subjects imagined the drumbeats continuing through subsequent silent periods. This entrainment in both the pupillometry and EEG worsened with increasing rhythmic complexity, indicating poorer sensory precision as the beat became more obscured. Additionally, sustained pupil dilations revealed the expected, inverted U-shaped relationship between rhythmic complexity and groove ratings. Taken together, this work bridges oscillatory attention to rhythmic complexity in relation to musical groove. |
Yannik Stegmann; Janna Teigeler; Arash Mirifar; Andreas Keil; Matthias Gamer Electrocortical responses in anticipation of avoidable and inevitable threats: A multisite study Journal Article In: The Journal of Neuroscience, vol. 44, no. 42, pp. 1–12, 2024. @article{Stegmann2024, When faced with danger, human beings respond with a repertoire of defensive behaviors, including freezing and active avoidance. Previous research has revealed a pattern of physiological responses, characterized by heart rate bradycardia, reduced visual exploration, and heightened sympathetic arousal in reaction to avoidable threats, suggesting a state of attentive immobility in humans. However, the electrocortical underpinnings of these behaviors remain largely unexplored. To investigate the visuocortical components of attentive immobility, we recorded parieto-occipital alpha activity, along with eye-movements and autonomic responses, while participants awaited either an avoidable, inevitable, or no threat. To test the robustness and generalizability of our findings, we collected data from a total of 101 participants (76 females, 35 males) at two laboratories. Across sites, we observed an enhanced suppression of parieto-occipital alpha activity during avoidable threats, in contrast to inevitable or no threat trials, particularly towards the end of the trial that prompted avoidance responses. This response pattern coincided with heart rate bradycardia, centralization of gaze and increased sympathetic arousal. Furthermore, our findings expand on previous research by revealing that the amount of alpha suppression, along with centralization of gaze, and heart rate changes, predict the speed of motor responses. Collectively, these findings indicate that when individuals encounter avoidable threats, they enter a state of attentive immobility, which enhances perceptual processing and facilitates action preparation. This state appears to reflect freezing-like behavior in humans. Significance Statement In response to avoidable danger, organisms often exhibit freezing-like behavior. Recent research suggests that freezing is not merely a passive response but involves a state of attentive immobility aimed at enhancing threat avoidance and perception. However, the attentional mechanisms involved in response to avoidable threats at the level of the brain remain poorly understood. To address this gap, we employed EEG, eye-tracking, and measurements of autonomic activity. Our findings revealed a suppression of EEG alpha power, along with cardiac deceleration, reduced eye-movements, and heightened sympathetic activity during the anticipation of avoidable threats. Moreover, this response pattern was predictive of motor response times. These results underscore the significance of heightened perceptual processing during freezing-like states in humans. |
Caleb Stone; Jason B. Mattingley; Stefan Bode; Dragan Rangelov Distinct neural markers of evidence accumulation index metacognitive processing before and after simple visual decisions Journal Article In: Cerebral Cortex, vol. 34, no. 5, pp. 1–11, 2024. @article{Stone2024, Perceptual decision-making is affected by uncertainty arising from the reliability of incoming sensory evidence (perceptual uncertainty) and the categorization of that evidence relative to a choice boundary (categorical uncertainty). Here, we investigated how these factors impact the temporal dynamics of evidence processing during decision-making and subsequent metacognitive judgments. Participants performed a motion discrimination task while electroencephalography was recorded. We manipulated perceptual uncertainty by varying motion coherence, and categorical uncertainty by varying the angular offset of motion signals relative to a criterion. After each trial, participants rated their desire to change their mind. High uncertainty impaired perceptual and metacognitive judgments and reduced the amplitude of the centro-parietal positivity, a neural marker of evidence accumulation. Coherence and offset affected the centro-parietal positivity at different time points, suggesting that perceptual and categorical uncertainty affect decision-making in sequential stages. Moreover, the centro-parietal positivity predicted participants' metacognitive judgments: larger predecisional centro-parietal positivity amplitude was associated with less desire to change one's mind, whereas larger postdecisional centro-parietal positivity amplitude was associated with greater desire to change one's mind, but only following errors. These findings reveal a dissociation between predecisional and postdecisional evidence processing, suggesting that the CPP tracks potentially distinct cognitive processes before and after a decision. |
Dawid Strzelczyk; Nicolas Langer Pre-stimulus activity mediates event-related theta synchronization and alpha desynchronization during memory formation in healthy aging Journal Article In: Imaging Neuroscience, vol. 2, pp. 1–22, 2024. @article{Strzelczyk2024, The capacity to learn is a key determinant for the quality of life, but is known to decline to varying degrees with age. However, despite mounting evidence of memory deficits in older age, the neural mechanisms contributing to successful or impeded memory remain unclear. Previous research has primarily focused on memory formation through remembered versus forgotten comparisons, lacking the ability to capture the incremental nature of learning. Moreover, previous electroencephalography (EEG) studies have primarily examined oscillatory brain activity during the encoding phase, such as event- related synchronization (ERS) of mid-frontal theta and desynchronization (ERD) of parietal alpha, while neglecting the potential influence of pre-stimulus activity. To address these limitations, we employed a sequence learning paradigm, where 113 young and 117 older participants learned a fixed sequence of visual locations through repeated observations (6,423 sequence repetitions, 55 '944 stimuli). This paradigm enabled us to investigate mid- frontal theta ERS, parietal alpha ERD, and how they are affected by pre-stimulus activity during the incremental learning process. Behavioral results revealed that young subjects learned significantly faster than older subjects, in line with expected age-related cognitive decline. Successful incremental learning was directly linked to decreases of mid-frontal theta ERS and increases of parietal alpha ERD. Notably, these neurophysiological changes were less pronounced in older individuals, reflecting a slower rate of learning. Importantly, the mediation analysis revealed that in both age groups, mid-frontal pre-stimulus theta partially mediated the relationship between learning and mid- frontal theta ERS. Furthermore, the overall impact of learning on parietal alpha ERD was primarily driven by its positive influence on pre-stimulus alpha activity. Our findings offer new insights into the age- related differences in memory formation and highlight the importance of pre-stimulus activity in explaining post- stimulus responses during learning. |
Binbin Sun; Elombe Issa Calvert; Alyssa Ye; Heng Mao; Kevin Liu; Raymond Kong Wang; Xin Yuan Wang; Zhi Liu Wu; Zhen Wei; Xue Jun Kong Interest paradigm for early identification of autism spectrum disorder: An analysis from electroencephalography combined with eye tracking Journal Article In: Frontiers in Neuroscience, vol. 18, pp. 1–13, 2024. @article{Sun2024, Introduction: Early identification of Autism Spectrum Disorder (ASD) is critical for effective intervention. Restricted interests (RIs), a subset of repetitive behaviors, are a prominent but underutilized domain for early ASD diagnosis. This study aimed to identify objective biomarkers for ASD by integrating electroencephalography (EEG) and eye-tracking (ET) to analyze toddlers' visual attention and cortical responses to RI versus neutral interest (NI) objects. Methods: The study involved 59 toddlers aged 2-4 years, including 32 with ASD and 27 non-ASD controls. Participants underwent a 24-object passive viewing paradigm, featuring RI (e.g., transportation items) and NI objects (e.g., balloons). ET metrics (fixation time and pupil size) and EEG time-frequency (TF) power in theta (4-8 Hz) and alpha (8-13 Hz) bands were analyzed. Statistical methods included logistic regression models to assess the predictive potential of combined EEG and ET biomarkers. Results: Toddlers with ASD exhibited significantly increased fixation times and pupil sizes for RI objects compared to NI objects, alongside distinct EEG patterns with elevated theta and reduced alpha power in occipital regions during RI stimuli. The multimodal logistic regression model, incorporating EEG and ET metrics, achieved an area under the curve (AUC) of 0.75, demonstrating robust predictive capability for ASD. Discussion: This novel integration of ET and EEG metrics highlights the potential of RIs as diagnostic markers for ASD. The observed neural and attentional distinctions underscore the utility of multimodal biomarkers for early diagnosis and personalized intervention strategies. Future work should validate findings across broader age ranges and diverse populations. |
Alessandro Toso; Annika P. Wermuth; Ayelet Arazi; Anke Braun; Tineke Grent-‘t Jong; Peter J. Uhlhaas; Tobias H. Donner 40 Hz steady-state response in human auditory cortex is shaped by gabaergic neuronal inhibition Journal Article In: The Journal of Neuroscience, vol. 44, no. 24, pp. 1–10, 2024. @article{Toso2024, The 40 Hz auditory steady-state response (ASSR), an oscillatory brain response to periodically modulated auditory stimuli, is a promising, noninvasive physiological biomarker for schizophrenia and related neuropsychiatric disorders. The 40 Hz ASSR might be amplified by synaptic interactions in cortical circuits, which are, in turn, disturbed in neuropsychiatric disorders. Here, we tested whether the 40 Hz ASSR in the human auditory cortex depends on two key synaptic components of neuronal interactions within cortical circuits: excitation via N-methyl-aspartate glutamate (NMDA) receptors and inhibition via gamma-amino-butyric acid (GABA) receptors. We combined magnetoencephalography (MEG) recordings with placebo-controlled, low-dose pharmacological interventions in the same healthy human participants (13 males, 7 females). All participants exhibited a robust 40 Hz ASSR in auditory cortices, especially in the right hemisphere, under a placebo. The GABAA receptor–agonist lorazepam increased the amplitude of the 40 Hz ASSR, while no effect was detectable under the NMDA blocker memantine. Our findings indicate that the 40 Hz ASSR in the auditory cortex involves synaptic (and likely intracortical) inhibition via the GABAA receptor, thus highlighting its utility as a mechanistic signature of cortical circuit dysfunctions involving GABAergic inhibition. |
Marius Tröndle; Nicolas Langer Decomposing neurophysiological underpinnings of age-related decline in visual working memory Journal Article In: Neurobiology of Aging, vol. 139, pp. 30–43, 2024. @article{Troendle2024, Exploring the neural basis of age-related decline in working memory is vital in our aging society. Previous electroencephalographic studies suggested that the contralateral delay activity (CDA) may be insensitive to age-related decline in lateralized visual working memory (VWM) performance. Instead, recent evidence indicated that task-induced alpha power lateralization decreases in older age. However, the relationship between alpha power lateralization and age-related decline of VWM performance remains unknown, and recent studies have questioned the validity of these findings due to confounding factors of the aperiodic signal. Using a sample of 134 participants, we replicated the age-related decrease of alpha power lateralization after adjusting for the aperiodic signal. Critically, the link between task performance and alpha power lateralization was found only when correcting for aperiodic signal biases. Functionally, these findings suggest that age-related declines in VWM performance may be related to the decreased ability to prioritize relevant over irrelevant information. Conversely, CDA amplitudes were stable across age groups, suggesting a distinct neural mechanism possibly related to preserved VWM encoding or early maintenance. |
A. Van Den Kerchove; H. Si-Mohammed; M. M. Van Hulle; F. Cabestaing Correcting for ERP latency jitter improves gaze-independent BCI decoding Journal Article In: Journal of Neural Engineering, vol. 21, no. 4, pp. 1–15, 2024. @article{VanDenKerchove2024, Objective. Patients suffering from heavy paralysis or Locked-in-Syndrome can regain communication using a Brain-Computer Interface (BCI). Visual event-related potential (ERP) based BCI paradigms exploit visuospatial attention (VSA) to targets laid out on a screen. However, performance drops if the user does not direct their eye gaze at the intended target, harming the utility of this class of BCIs for patients suffering from eye motor deficits. We aim to create an ERP decoder that is less dependent on eye gaze. Approach. ERP component latency jitter plays a role in covert visuospatial attention (VSA) decoding. We introduce a novel decoder which compensates for these latency effects, termed Woody Classifier-based Latency Estimation (WCBLE). We carried out a BCI experiment recording ERP data in overt and covert visuospatial attention (VSA), and introduce a novel special case of covert VSA termed split VSA, simulating the experience of patients with severely impaired eye motor control. We evaluate WCBLE on this dataset and the BNCI2014-009 dataset, within and across VSA conditions to study the dependency on eye gaze and the variation thereof during the experiment. Main results. WCBLE outperforms state-of-the-art methods in the VSA conditions of interest in gaze-independent decoding, without reducing overt VSA performance. Results from across-condition evaluation show that WCBLE is more robust to varying VSA conditions throughout a BCI operation session. Significance. Together, these results point towards a pathway to achieving gaze independence through suited ERP decoding. Our proposed gaze-independent solution enhances decoding performance in those cases where performing overt VSA is not possible. |
Daniel Walper; Alexandra Bendixen; Sabine Grimm; Anna Schubö; Wolfgang Einhäuser Attention deployment in natural scenes: Higher-order scene statistics rather than semantics modulate the N2pc component Journal Article In: Journal of vision, vol. 24, no. 6, pp. 1–28, 2024. @article{Walper2024, Which properties of a natural scene affect visual search? We consider the alternative hypotheses that low-level statistics, higher-level statistics, semantics, or layout affect search difficulty in natural scenes. Across three experiments (n = 20 each), we used four different backgrounds that preserve distinct scene properties: (a) natural scenes (all experiments); (b) 1/f noise (pink noise, which preserves only low-level statistics and was used in Experiments 1 and 2); (c) textures that preserve low-level and higher-level statistics but not semantics or layout (Experiments 2 and 3); and (d) inverted (upside-down) scenes that preserve statistics and semantics but not layout (Experiment 2). We included "split scenes" that contained different backgrounds left and right of the midline (Experiment 1, natural/noise; Experiment 3, natural/texture). Participants searched for a Gabor patch that occurred at one of six locations (all experiments). Reaction times were faster for targets on noise and slower on inverted images, compared to natural scenes and textures. The N2pc component of the event-related potential, a marker of attentional selection, had a shorter latency and a higher amplitude for targets in noise than for all other backgrounds. The background contralateral to the target had an effect similar to that on the target side: noise led to faster reactions and shorter N2pc latencies than natural scenes, although we observed no difference in N2pc amplitude. There were no interactions between the target side and the non-target side. Together, this shows that-at least when searching simple targets without own semantic content-natural scenes are more effective distractors than noise and that this results from higher-order statistics rather than from semantics or layout. |
Kangning Wang; Wei Wei; Weibo Yi; Shuang Qiu; Huiguang He; Minpeng Xu; Dong Ming Contrastive fine-grained domain adaptation network for EEG-based vigilance estimation Journal Article In: Neural Networks, vol. 179, pp. 1–18, 2024. @article{Wang2024d, Vigilance state is crucial for the effective performance of users in brain-computer interface (BCI) systems. Most vigilance estimation methods rely on a large amount of labeled data to train a satisfactory model for the specific subject, which limits the practical application of the methods. This study aimed to build a reliable vigilance estimation method using a small amount of unlabeled calibration data. We conducted a vigilance experiment in the designed BCI-based cursor-control task. Electroencephalogram (EEG) signals of eighteen participants were recorded in two sessions on two different days. And, we proposed a contrastive fine-grained domain adaptation network (CFGDAN) for vigilance estimation. Here, an adaptive graph convolution network (GCN) was built to project the EEG data of different domains into a common space. The fine-grained feature alignment mechanism was designed to weight and align the feature distributions across domains at the EEG channel level, and the contrastive information preservation module was developed to preserve the useful target-specific information during the feature alignment. The experimental results show that the proposed CFGDAN outperforms the compared methods in our BCI vigilance dataset and SEED-VIG dataset. Moreover, the visualization results demonstrate the efficacy of the designed feature alignment mechanisms. These results indicate the effectiveness of our method for vigilance estimation. Our study is helpful for reducing calibration efforts and promoting the practical application potential of vigilance estimation methods. |
Wei Wei; Kangning Wang; Shuang Qiu; Huiguang He A MultiModal Vigilance (MMV) dataset during RSVP and SSVEP brain-computer interface tasks Journal Article In: Scientific Data, vol. 11, no. 1, pp. 1–14, 2024. @article{Wei2024, Vigilance represents an ability to sustain prolonged attention and plays a crucial role in ensuring the reliability and optimal performance of various tasks. In this report, we describe a MultiModal Vigilance (MMV) dataset comprising seven physiological signals acquired during two Brain-Computer Interface (BCI) tasks. The BCI tasks encompass a rapid serial visual presentation (RSVP)-based target image retrieval task and a steady-state visual evoked potential (SSVEP)-based cursor-control task. The MMV dataset includes four sessions of seven physiological signals for 18 subjects, which encompasses electroencephalogram(EEG), electrooculogram (EOG), electrocardiogram (ECG), photoplethysmogram (PPG), electrodermal activity (EDA), electromyogram (EMG), and eye movement. The MMV dataset provides data from four stages: 1) raw data, 2) pre-processed data, 3) trial data, and 4) feature data that can be directly used for vigilance estimation. We believe this dataset will achieve flexible reuse and meet the various needs of researchers. And this dataset will greatly contribute to advancing research on physiological signal-based vigilance research and estimation. |
Jaeger Wongtrakun; Shou-Han Zhou; Mark A. Bellgrove; Trevor T. J. Chong; James P. Coxon The effect of congruent versus incongruent distractor positioning on electrophysiological signals during perceptual decision-making Journal Article In: The Journal of Neuroscience, vol. 44, no. 45, pp. 1–9, 2024. @article{Wongtrakun2024, Key event-related potentials (ERPs) of perceptual decision-making such as centroparietal positivity (CPP) elucidate how evidence is accumulated toward a given choice. Furthermore, this accumulation can be impacted by visual target selection signals such as the N2 contralateral (N2c). How these underlying neural mechanisms of perceptual decision-making are influenced by the spatial congruence of distractors relative to target stimuli remains unclear. Here, we used electroencephalography (EEG) in humans of both sexes to investigate the effect of distractor spatial congruency (same vs different hemifield relative to targets) on perceptual decision-making. We confirmed that responses for perceptual decisions were slower for spatially incongruent versus congruent distractors of high salience. Similarly, markers of target selection (N2c peak amplitude) and evidence accumulation (CPP slope) were found to be lower when distractors were spatially incongruent versus congruent. To evaluate the effects of congruency further, we applied drift diffusion modeling to participant responses, which showed that larger amplitudes of both ERPs were correlated with shorter nondecision times when considering the effect of congruency. The modeling also suggested that congruency's effect on behavior occurred prior to and during evidence accumulation when considering the effects of the N2c peak and CPP slope. These findings point to spatially incongruent distractors, relative to congruent distractors, influencing decisions as early as the initial sensory processing phase and then continuing to exert an effect as evidence is accumulated throughout the decision-making process. Overall, our findings highlight how key electrophysiological signals of perceptual decision-making are influenced by the spatial congruence of target and distractor. |
2023 |
Elizabeth R. Schotter; Sara Milligan; Victoria M. Estevez Event-related potentials show that parafoveal vision is insufficient for semantic integration Journal Article In: Psychophysiology, vol. 60, no. 7, pp. 1–25, 2023. @article{Schotter2023, Readers extract information from a word from parafoveal vision prior to looking at it. It has been argued that parafoveal perception allows readers to initiate linguistic processes, but it is unclear which stages of word processing are engaged: the process of extracting letter information to recognize words, or the process of extracting meaning to comprehend them. This study used the event-related brain potential (ERP) technique to investigate how word recognition (indexed by the N400 effect for unexpected or anomalous compared to expected words) and semantic integration (indexed by the Late-positive component; LPC effect for anomalous compared to expected words) are or are not elicited when the word is perceived only in parafoveal vision. Participants read a target word following a sentence that made it expected, unexpected, or anomalous, and read the sentences presented three words at a time in the Rapid Serial Visual Presentation (RSVP) with flankers paradigm so that words were perceived in parafoveal and foveal vision. We orthogonally manipulated whether the target word was masked in parafoveal and/or foveal vision to dissociate the processing associated with perception of the target word from either location. We found that the N400 effect was generated from parafoveally perceived words, and was reduced for foveally perceived words if they were previously perceived parafoveally. In contrast, the LPC effect was only elicited if the word was perceived foveally, suggesting that readers must attend to a word directly in foveal vision in order to attempt to integrate its meaning into the sentence context. |
Svea C. Y. Schroeder; David Aagten-Murphy; Niko A. Busch Contralateral delay activity, but not alpha lateralization, indexes prioritization of information for working memory storage Journal Article In: Attention, Perception, & Psychophysics, vol. 85, no. 3, pp. 718–733, 2023. @article{Schroeder2023, Working memory is inherently limited, which makes it important to select and maintain only task-relevant information and to protect it from distraction. Previous research has suggested the contralateral delay activity (CDA) and lateralized alpha oscillations as neural candidates for such a prioritization process. While most of this work focused on distraction during encoding, we examined the effect of external distraction presented during memory maintenance. Participants memorized the orientations of three lateralized objects. After an initial distraction-free maintenance interval, distractors appeared in the same location as the targets or in the opposite hemifield. This distraction was followed by another distraction-free interval. Our results show that CDA amplitudes were stronger in the interval before compared with the interval after the distraction (i.e., CDA amplitudes were stronger in response to targets compared with distractors). This amplitude reduction in response to distractors was more pronounced in participants with higher memory accuracy, indicating prioritization and maintenance of relevant over irrelevant information. In contrast, alpha lateralization did not change from the interval before distraction compared with the interval after distraction, and we found no correlation between alpha lateralization and memory accuracy. These results suggest that alpha lateralization plays no direct role in either selective maintenance of task-relevant information or inhibition of distractors. Instead, alpha lateralization reflects the current allocation of spatial attention to the most salient information regardless of task-relevance. In contrast, CDA indicates flexible allocation of working memory resources depending on task-relevance. |
Joshua Snell; Jeremy Yeaton; Jonathan Mirault; Jonathan Grainger Parallel word reading revealed by fixation-related brain potentials Journal Article In: Cortex, vol. 162, pp. 1–11, 2023. @article{Snell2023, During reading, the brain is confronted with many relevant objects at once. But does lexical processing occur for multiple words simultaneously? Cognitive science has yet to answer this prominent question. Recently it has been argued that the issue warrants supplementing the field's traditional toolbox (response times, eye-tracking) with neuroscientific techniques (EEG, fMRI). Indeed, according to the OB1-reader model, upcoming words need not impact oculomotor behavior per se, but parallel processing of these words must nonetheless be reflected in neural activity. Here we combined eye-tracking with EEG, time-locking the neural window of interest to the fixation on target words in sentence reading. During these fixations, we manipulated the identity of the subsequent word so that it posed either a syntactically legal or illegal continuation of the sentence. In line with previous research, oculomotor measures were unaffected. Yet, syntax impacted brain potentials as early as 100 ms after the target fixation onset. Given the EEG literature on syntax processing, the presently observed timings suggest parallel word reading. We reckon that parallel word processing typifies reading, and that OB1-reader offers a good platform for theorizing about the reading brain. |