All EyeLink eye tracker neuroscience peer-reviewed journal publications (with concurrent eye tracking) up until 2025 (with some early 2026s) are listed below by year. These publications include fMRI, MEG, EEG, fNIRS, and TMS articles. You can search the eye-tracking publications using keywords such as NIRS, Visual Cortex, Neural Plasticity, MEG, P300, etc. You can also search for individual author names. If we missed any EyeLink neuroscience articles, please email us!
2025 |
Ankan Biswas; Wupadrasta Santosh Kumar; Kanishka Sharma; Supratim Ray Stimulus-induced gamma sources reduce in power but not in spatial extent with healthy aging in human EEG Journal Article In: European Journal of Neuroscience, vol. 61, no. 10, pp. 1–12, 2025. @article{Biswas2025,Aging alters brain structure and function, and studying such changes may help understand the neural basis underlying aging and devise interventions to detect deviations from healthy progression. Electroencephalogram (EEG) offers an effective way to study healthy aging owing to its high temporal resolution and affordability. Recent studies have shown that narrow-band stimulus-induced gamma oscillations (20–70 Hz) in EEG, induced with Cartesian gratings in a fixation task paradigm, weaken with healthy aging and onset of Alzheimer's disease (AD) while remaining highly reproducible for a given subject and thus hold promise as potential biomarkers. However, functional connectivity (FC) sometimes changes in a different way compared with sensor power with aging. This difference could be potentially addressed by studying how underlying gamma sources change with aging, since either a reduction in source power or a shrinkage of the sources (or both) could reduce the power in the sensors but may have different effects on other measures such as FC. We therefore reconstructed EEG gamma sources through a linear inverse method called exact low-resolution tomography analysis (eLORETA) on a large (N = 217) cohort of healthy elderly subjects (> 50 years). We further characterized gamma distribution in cortical space as an exponential fall-off from a seed voxel with maximal gamma source power to delineate a reduction in magnitude versus shrinkage. We found a significant reduction in magnitude but not shrinkage with healthy aging. Overall, our results shed light on changes in EEG gamma source distribution with healthy aging, which could provide clues about underlying neural mechanisms. |
Dock H. Duncan; Norman Forschack; Dirk Moorselaar; Matthias M. Müller; Jan Theeuwes Learning modulates early encephalographic responses to distracting stimuli: A combined SSVEP and ERP study Journal Article In: Journal of Neuroscience, vol. 45, no. 21, pp. 1–13, 2025. @article{Duncan2025,Through experience, humans can learn to suppress locations that frequently contain distracting stimuli. However, the neural mechanism underlying learned suppression remains largely unknown. In this study, we combined steady-state visually evoked potentials (SSVEPs) with event-related potentials (ERPs) to investigate the mechanism behind statistically learned spatial suppression. Twenty-four male and female human participants performed a version of the additional singleton search task in which one location contained a distractor stimulus frequently. The search stimuli constantly flickered on-and-off the screen, resulting in steady-state entrainment. Prior to search onset, no differences in the SSVEP response were found, though a post hoc analysis did reveal proactive alpha lateralization. Following search onset, clear evoked differences in both the SSVEP and ERP signals emerged at the suppressed location relative to all other locations. Crucially, the early timing of these evoked modulations suggests that learned distractor suppression occurs at the initial stages of visual processing. |
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, vol. 87, no. 4, pp. 1199–1218, 2025. @article{Guida2025,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. |
Francesco Mantegna; Emanuele Olivetti; Philipp Schwedhelm; Daniel Baldauf Covariance-based decoding reveals a category-specific functional connectivity network for imagined visual objects Journal Article In: NeuroImage, vol. 311, pp. 1–15, 2025. @article{Mantegna2025,The coordination of different brain regions is required for the visual imagery of complex objects (e.g., faces and places). Short-range connectivity within sensory areas is necessary to construct the mental image. Long-range connectivity between control and sensory areas is necessary to re-instantiate and maintain the mental image. While dynamic changes in functional connectivity are expected during visual imagery, it is unclear whether a category-specific network exists in which the strength and the spatial destination of the connections vary depending on the imagery target. In this magnetoencephalography study, we used a minimally constrained experimental paradigm wherein imagery categories were prompted using visual word cues only, and we decoded face versus place imagery based on their underlying functional connectivity patterns as estimated from the spatial covariance across brain regions. A subnetwork analysis further disentangled the contribution of different connections. The results show that face and place imagery can be decoded from both short-range and long-range connections. Overall, the results show that imagined object categories can be distinguished based on functional connectivity patterns observed in a category-specific network. Notably, functional connectivity estimates rely on purely endogenous brain signals suggesting that an external reference is not necessary to elicit such category-specific network dynamics. |
Brian R. Cornwell; Paige R. Didier; Shannon E. Grogans; Allegra S. Anderson; Samiha Islam; Hyung Cho Kim; Manuel Kuhn; Rachael M. Tillman; Juyoen Hur; Zachary S. Scott; Andrew S. Fox; Kathryn A. DeYoung; Jason F. Smith; Alexander J. Shackman A shared threat-anticipation circuit is dynamically engaged at different moments by certain and uncertain threat Journal Article In: The Journal of Neuroscience, vol. 45, no. 16, pp. 1–19, 2025. @article{Cornwell2025,Temporal dynamics play a central role in models of emotion: “fear” is widely conceptualized as a phasic response to certain-and-imminent danger, whereas “anxiety” is a sustained response to uncertain-or-distal harm. Yet the underlying neurobiology remains contentious. Leveraging a translationally relevant fMRI paradigm and theory-driven modeling approach in 220 adult humans, we demonstrate that certain- and uncertain-threat anticipation recruit a shared circuit that encompasses the central extended amygdala (EAc), periaqueductal gray, midcingulate, and anterior insula. This circuit exhibits persistently elevated activation when threat is uncertain and distal and transient bursts of activation just before certain encounters with threat. Although there is agreement that the EAc plays a critical role in orchestrating responses to threat, confusion persists about the respective contributions of its major subdivisions, the bed nucleus of the stria terminalis (BST) and central nucleus of the amygdala (Ce). Here we used anatomical regions of interest to demonstrate that the BST and Ce exhibit statistically indistinguishable threat dynamics. Both regions exhibited activation dynamics that run counter to popular models, with the Ce (and BST) showing sustained responses to uncertain-and-distal threat and the BST (and Ce) showing phasic responses to certain-and-imminent threat. For many scientists, feelings are the hallmark of fear and anxiety. Here we used an independently validated multivoxel brain “signature” to covertly probe the moment-by-moment dynamics of anticipatory distress for the first time. Results mirrored the dynamics of neural activation. These observations provide fresh insights into the neurobiology of threat-elicited emotions and set the stage for more ambitious clinical and mechanistic research. |
Ziyi Duan; Marissa H. Evans; Bonnie Lawrence; Clayton E. Curtis Effector general representation of movement goals in human frontal and parietal cortex Journal Article In: NeuroImage, vol. 310, pp. 1–9, 2025. @article{Duan2025,In the nonhuman primate, discrete parts of premotor frontal and parietal cortex appear to code for movements of different effectors. However, the evidence regarding homologous effector selectivity within the human brain remains inconclusive. Here, we measured neural activity in the human brain using functional magnetic resonance imaging while participants remembered a target location and planned either saccades or reaches that matched the rich kinematics used in seminal monkey studies. We compared activity patterns during the planning period and used assumption-free multivariate searchlight analysis to identify brain regions that could decode the spatial goals of planned movements. Critically, we performed two types of decoding analyses to determine if the spatial information embedded in activation patterns was effector-specific or effector-general. For effector-specific spatial coding, we compared brain regions that could decode target locations within each effector. However, we did not identify areas that coded spatial information in one effector but not the other. For effector-general spatial coding, we performed spatial decoding using trials across effectors and conducted cross-effector decoding. Both analyses identified several areas in the frontal and parietal regions that encoded spatial information for both effectors, including precentral sulcus, superior parietal lobe, and intraparietal sulcus. Our results indicate that premotor frontal and parietal cortex encode the spatial metrics of movement goals that can be read out and converted into effector-specific motor metrics for saccades and reaches. |
Robert Woodry; Clayton E. Curtis; Jonathan Winawer Feedback scales the spatial tuning of cortical responses during both visual working memory and long-term memory Journal Article In: Journal of Neuroscience, vol. 45, no. 17, pp. 1–15, 2025. @article{Woodry2025,Perception, working memory, and long-term memory each evoke neural responses in the visual cortex. While previous neuroimaging research on the role of the visual cortex in memory has largely emphasized similarities between perception and memory, we hypothesized that responses in the visual cortex would differ depending on the origins of the inputs. Using fMRI, we quantified spatial tuning in the visual cortex while participants (both sexes) viewed, maintained in working memory, or retrieved from long-term memory a peripheral target. In each condition, BOLD responses were spatially tuned and aligned with the target's polar angle in all measured visual field maps including V1. As expected given the increasing sizes of receptive fields, polar angle tuning during perception increased in width up the visual hierarchy from V1 to V2, V3, hV4, and beyond. In stark contrast, the tuned responses were broad across the visual hierarchy during long-term memory (replicating a prior result) and during working memory. This pattern is consistent with the idea that mnemonic responses in V1 stem from top-down sources, even when the stimulus was recently viewed and is held in working memory. Moreover, in long-term memory, trial-to-trial biases in these tuned responses (clockwise or counterclockwise of target) predicted matched biases in memory, suggesting that the reinstated cortical responses influence memory-guided behavior. We conclude that feedback widens spatial tuning in the visual cortex during memory, where earlier visual maps inherit broader tuning from later maps thereby impacting the precision of memory. |
Mana Biabani; Kevin Walsh; Shou Han Zhou; Joseph Wagner; Alexandra Johnstone; Julia Paterson; Beth P. Johnson; Natasha Matthews; Gerard M. Loughnane; Redmond G. O'Connell; Mark A. Bellgrove Neurophysiology of perceptual decision-making and its alterations in attention-deficit hyperactivity disorder Journal Article In: The Journal of Neuroscience, vol. 45, no. 14, pp. 1–12, 2025. @article{Biabani2025,Despite the prevalence of attention-deficit hyperactivity disorder (ADHD), efforts to develop a detailed understanding of the neuropsychology of this neurodevelopmental condition are complicated by the diversity of interindividual presentations and the inability of current clinical tests to distinguish between its sensory, attentional, arousal, or motoric contributions. Identifying objective methods that can explain the diverse performance profiles across individuals diagnosed with ADHD has been a long-held goal. Achieving this could significantly advance our understanding of etiological processes and potentially inform the development of personalized treatment approaches. Here, we examine key neuropsychological components of ADHD within an electrophysiological (EEG) perceptual decision-making paradigm that is capable of isolating distinct neural signals of several key information processing stages necessary for sensory-guided actions from attentional selection to motor responses. Using a perceptual decision-making task (random dot motion), we evaluated the performance of 79 children (aged 8–17 years) and found slower and less accurate responses, along with a reduced rate of evidence accumulation (drift rate parameter of drift diffusion model), in children with ADHD (n = 37; 13 female) compared with typically developing peers (n = 42; 18 female). This was driven by the atypical dynamics of discrete electrophysiological signatures of attentional selection, the accumulation of sensory evidence, and strategic adjustments reflecting urgency of response. These findings offer an integrated account of decision-making in ADHD and establish discrete neural signals that might be used to understand the wide range of neuropsychological performance variations in individuals with ADHD. |
Steven P. Errington; Jeffrey D. Schall A preparatory cranial potential for saccadic eye movements in macaque monkeys Journal Article In: eNeuro, vol. 12, no. 4, pp. 1–10, 2025. @article{Errington2025,Response preparation is accomplished by gradual accumulation in neural activity until a threshold is reached. In humans, such a preparatory signal, referred to as the lateralized readiness potential (LRP), can be observed in the EEG over sensorimotor cortical areas before execution of a voluntary movement. Although well described for manual movements, less is known about preparatory EEG potentials for saccadic eye movements in humans and nonhuman primates. Hence, we describe a LRP over the frontolateral cortex in macaque monkeys. Homologous to humans, we observed lateralized electrical potentials ramping before the execution of both rewarded and nonrewarded contralateral saccades. This potential parallels the neural spiking of saccadic movement neurons in the frontal eye field (FEF), suggesting that it may offer a noninvasive correlate of intracortical spiking activity. However, unlike neural spiking in the FEF, polarization in frontolateral channels did not distinguish between saccade generation and inhibition. These findings provide new insights into noninvasive electrophysiological signatures of saccadic preparation in nonhuman primates, highlighting the potential of EEG measures to bridge invasive neural recordings and noninvasive studies of eye movement control in humans. |
Daniela Gresch; Larissa Behnke; Freek Ede; Anna C. Nobre; Sage E. P. Boettcher Neural dynamics of reselecting visual and motor contents in working memory after external interference Journal Article In: The Journal of Neuroscience, vol. 45, no. 18, pp. 1–13, 2025. @article{Gresch2025,In everyday tasks, we must often shift our focus away from internal representations held in working memory to engage with perceptual events in the external world. Here, we investigated how our internal focus is reestablished following an interrupting task by tracking the reselection of visual representations and their associated action plans in working memory. Specifically, we asked whether reselection occurs for both visual and motor memory attributes and when this reselection occurs. We developed a visual-motor working-memory task in which participants were retrospectively cued to select one of two memory items before being interrupted by a perceptual discrimination task. To determine what information was reselected, the memory items had distinct visual and motor attributes. To determine when internal representations were reselected, the interrupting task was presented at one of three distinct time points following the retro-cue. We employed electroencephalography time–frequency analyses to track the initial selection and later reselection of visual and motor representations, as operationalized through modulations of posterior alpha (8–12 Hz) activity relative to the memorized item location (visual) and of central beta (13–30 Hz) activity relative to the required response hand (motor). Our results show that internal visual and motor contents were concurrently reselected immediately after completing the interrupting task, rather than only when internal information was required for memory-guided behavior. Thus, following interruption, we swiftly resume our internal focus in working memory through the simultaneous reselection of memorized visual representations and their associated action plans, thereby restoring internal contents to a ready-to-use state. |
Catherine N. Moran; David P. McGovern; Mike Melnychuk; Alan F. Smeaton; Paul M. Dockree Oscillations of the wandering mind: Neural evidence for distinct exploration/exploitation strategies in younger and older adults Journal Article In: Human Brain Mapping, vol. 46, no. 6, pp. 1–23, 2025. @article{Moran2025a,This study traced the neurophysiological signals of fluctuating attention and task-related processing to ascertain the mechanistic basis of transient strategic shifts between competing task focus and mind-wandering, as expressed by the ‘exploitation/exploration' framework, and explored how they are differentially affected with age. Thirty-four younger (16 female, mean age 22 years) and 34 healthy older (20 female, mean age 71 years) adults performed the Gradual Contrast Change Detection task; monitoring a continuously presented flickering annulus for intermittent gradual contrast reductions and responding to experience sampling probes to discriminate the nature of their thoughts at discrete moments. Electroencephalography and pupillometry were concurrently recorded during target- and probe-related intervals. Older adults tracked the downward stimulus trajectory with greater sensory integrity (reduced target SSVEP amplitude) and demonstrated earlier initiation of evidence accumulation (earlier onset CPP), attenuated variability in the attentional signal (posterior alpha) and more robust phasic pupillary responses to the target, suggesting steadier attentional engagement with age. Younger adults only exhibited intermittent sensory encoding, indexed by greater variability in the sensory (SSVEP) and attentional (alpha) signals before mind-wandering relative to focused states. Attentional variability was accompanied by disrupted behavioural performance and reduced task-related neural processing, independent of age group. Together, this elucidates distinct performance strategies employed by both groups. Older adults suspended mind-wandering and implemented an exploitative oscillation strategy to circumvent their reduced cognitive resources and allay potential behavioural costs. Conversely, younger adults exhibited greater exploration through mind-wandering, utilising their greater cognitive resources to flexibly alternate between competing goal-directed and mind-wandering strategies, with limited costs. |
Nir Ofir; Ayelet N. Landau Motor preparation tracks decision boundary crossing rather than accumulated evidence in temporal decision-making Journal Article In: The Journal of Neuroscience, vol. 45, no. 17, pp. 1–11, 2025. @article{Ofir2025,Interval timing, the ability of animals to estimate the passage of time, is thought to involve diverse neural processes rather than a single central “clock” (Paton and Buonomano, 2018). Each of the different processes engaged in interval timing follows a different dynamic path, according to its specific function. For example, attention tracks anticipated events, such as offsets of intervals (Rohenkohl and Nobre, 2011), while motor processes control the timing of the behavioral output (De Lafuente et al., 2024). However, which processes are involved and how they are orchestrated over time to produce a temporal decision remains unknown. Here, we study motor preparation in the temporal bisection task, in which human (female and male) participants categorized intervals as “long” or “short.” In contrast to typical perceptual decisions, where motor plans for all response alternatives are prepared simultaneously (Shadlen and Kiani, 2013), we find that temporal bisection decisions develop sequentially. While preparation for “long” responses was already underway before interval offset, no preparation was found for “short” responses. Furthermore, within intervals categorized as “long,” motor preparation was stronger at interval offset for faster responses. Our findings support the two-stage model of temporal decisions, where “long” decisions are considered during the interval itself, while “short” decisions are only considered after the interval is over. Viewed from a wider perspective, our study offers methods to study the neural mechanisms of temporal decisions, by studying the multiple processes that produce them. |
Ying Que; Xiao Hu Enhancing learners' reading comprehension with preferred background music: An eye-tracking, EEG, and heart rate study Journal Article In: Reading Research Quarterly, vol. 60, no. 2, pp. 1–17, 2025. @article{Que2025,It is often observed that many learners prefer reading texts with background music (BGM), which may enhance engagement and mood in reading. However, the impact of BGM on reading comprehension performance or process remains heterogeneous. This study examined how learners' self-provided preferred BGM affected reading comprehension accuracy and cognitive processes as measured by self-reports and multimodal psychophysiological signals (e.g., eye movements, EEG signals, heart rates) during reading, while also considering the role of personal traits and music characteristics. Data were collected from a within-subject experiment with 52 nonnative English speakers, who read half of the English text passages with their self-provided BGM and the other half in silence. The results indicated that overall BGM did not significantly affect reading task performance, self-reports, or multimodal psychophysiological responses. Hierarchical linear modeling revealed that learners' personal traits (e.g., working memory capacity, multitasking ability, language proficiency, BGM listening habits, prior knowledge) moderated the effects of BGM on their cognitive processes during reading as indicated by the psychophysiological responses. Stepwise linear regression showed that the presence of lyrics predicted decreased reading comprehension accuracy and disrupted word-level lexical processing, while a fast tempo was related to more efficient text processing. These findings offer empirical evidence on how BGM influences learners' reading task performance and cognitive processes and provide practical guidance for stakeholders (e.g., learners, instructors, learning environment designers) on the personalized and appropriate use of BGM for enhancing reading comprehension. |
David Richter; Dirk Moorselaar; Jan Theeuwes Proactive distractor suppression in early visual cortex Journal Article In: eLife, vol. 13, pp. 1–22, 2025. @article{Richter2025,Avoiding distraction by salient yet irrelevant stimuli is critical when accomplishing daily tasks. One possible mechanism to accomplish this is by suppressing stimuli that may be distracting such that they no longer compete for attention. While the behavioral benefits of distractor suppression are well established, its neural underpinnings are not yet fully understood. In a functional MRI (fMRI) study, we examined whether and how sensory responses in early visual areas show signs of distractor suppression after incidental learning of spatial statistical regularities. Participants were exposed to an additional singleton task where, unbeknownst to them, one location more frequently contained a salient distractor. We analyzed whether visual responses in terms of fMRI BOLD were modulated by this distractor predictability. Our findings indicate that implicit spatial priors shape sensory processing even at the earliest stages of cortical visual processing, evident in early visual cortex as a suppression of stimuli at locations which frequently contained distracting information. Notably, while this suppression was spatially (receptive field) specific, it did extend to nearby neutral locations and occurred regardless of whether distractors, nontarget items, or targets were presented at this location, suggesting that suppression arises before stimulus identification. Crucially, we observed similar spatially specific neural suppression even if search was only anticipated, but no search display was presented. Our results highlight proactive modulations in early visual cortex, where potential distractions are suppressed preemptively, before stimulus onset, based on learned expectations. Combined, our study underscores how the brain leverages implicitly learned prior knowledge to optimize sensory processing and attention allocation. |
Grace E. Hallenbeck; Nathan Tardiff; Thomas C. Sprague; Clayton E. Curtis Prioritizing working memory resources depends on the prefrontal cortex Journal Article In: The Journal of Neuroscience, vol. 45, no. 11, pp. 1–10, 2025. @article{Hallenbeck2025,How the prefrontal cortex contributes to working memory remains controversial, as theories differ in their emphasis on its role in storing memories versus controlling their content. To adjudicate between these competing ideas, we tested how perturbations to the human (both sexes) lateral prefrontal cortex impact the storage and control aspects of working memory during a task that requires human subjects to allocate resources to memory items based on their behavioral priority. Our computational model made a strong prediction that disruption of this control process would counterintuitively improve memory for low-priority items. Remarkably, transcranial magnetic stimulation of retinotopically-defined superior precentral sulcus, but not intraparietal sulcus, unbalanced the prioritization of resources, improving memory for low-priority items as predicted by the model. Therefore, these results provide direct causal support for models in which the prefrontal cortex controls the allocation of resources that support working memory, rather than simply storing the features of memoranda. |
Amie J. Durston; Roxane J. Itier Event-related potentials to facial expressions are related to stimulus-level perceived arousal and valence Journal Article In: Psychophysiology, vol. 62, no. 3, 2025. @article{Durston2025,Facial expressions provide critical details about social partners' inner states. We investigated whether event-related potentials (ERP) related to the visual processing of facial expressions are modulated by participants' perceived arousal and valence at the stimulus level. ERPs were recorded while participants (N = 80) categorized the gender of faces expressing fear, anger, happiness, and no emotion. Participants then viewed each face again and rated them on arousal and valence using 1–9 Likert scales. For each participant, ratings of each unique face were linked back to corresponding ERP trials. ERPs were analyzed at all time points and electrodes using hierarchical mass univariate statistics. Three different ANOVA models were employed: the original emotion model, and models with valence or arousal ratings as trial-level regressors. Results from models with ratings highly overlapped with the original model, although they were more temporally restricted. The N170 component was the most impacted by arousal and valence ratings, with four out of six emotion contrasts revealing significant valence or arousal interactions. Emotion effects on the P2 component were mostly unrelated to ratings. On the EPN component, only two contrasts related to both arousal and valence ratings. Thus, ERP emotion effects are related to participants' perceived arousal and valence of the stimuli, although this association depends on the contrast analyzed. These findings, their limitations, and generalizability are discussed in reference to existing theories and literature. |
Caoimhe Moran; Philippa A. Johnson; Hinze Hogendoorn; Ayelet N. Landau The representation of stimulus features during stable fixation and active vision Journal Article In: The Journal of Neuroscience, vol. 45, no. 12, pp. 1–11, 2025. @article{Moran2025,Predictive updating of an object's spatial coordinates from presaccade to postsaccade contributes to stable visual perception. Whether object features are predictively remapped remains contested. We set out to characterize the spatiotemporal dynamics of feature processing during stable fixation and active vision. To do so, we applied multivariate decoding methods to EEG data collected while human participants (male and female) viewed brief visual stimuli. Stimuli appeared at different locations across the visual field at either high or low spatial frequency (SF). During fixation, classifiers were trained to decode SF presented at one parafoveal location and cross-tested on SF from either the same, adjacent, or more peripheral locations. When training and testing on the same location, SF was classified shortly after stimulus onset (∼79 ms). Decoding of SF at locations farther from the trained location emerged later (∼144–295 ms), with decoding latency modulated by eccentricity. This analysis provides a detailed time course for the spread of feature information across the visual field. Next, we investigated how active vision impacts the emergence of SF information. In the presence of a saccade, the decoding time of peripheral SF at parafoveal locations was earlier, indicating predictive anticipation of SF due to the saccade. Crucially, however, this predictive effect was not limited to the specific remapped location. Rather, peripheral SF was correctly classified, at an accelerated time course, at all parafoveal positions. This indicates spatially coarse, predictive anticipation of stimulus features during active vision, likely enabling a smooth transition on saccade landing. |
Veera Ruuskanen; C. Nico Boehler; Sebastiaan Mathôt The interplay of spontaneous pupil-size fluctuations and EEG power in near-threshold detection Journal Article In: Psychophysiology, vol. 62, no. 3, pp. 1–11, 2025. @article{Ruuskanen2025,Detection of near-threshold stimuli depends on the properties of the stimulus and the state of the observer. In visual detection tasks, improved accuracy is associated with larger prestimulus pupil size. However, it is still unclear whether this association is due to optical effects (more light entering the eye), correlations with arousal, correlations with cortical excitability (as reflected in alpha power), or a mix of these. To better understand this, we investigated the relative contributions of pupil size and power in the alpha, beta, and theta frequency bands on near-threshold detection. We found that larger prestimulus pupil size is associated with improved accuracy and more stimulus-present responses, and these effects were not mediated by spectral power in the EEG. Pupil size was also positively correlated with power in the beta and alpha bands. Taken together, our results show an independent effect of pupil size on detection performance that is not driven by cortical excitability but may be driven by optical effects, physiological arousal, or a mix of both. |
Markus R Tünte; Stefanie Hoehl; Moritz Wunderwald; Johannes Bullinger; Asena Boyadziheva; Lara Maister; Birgit Elsner; Manos Tsakiris; Ezgi Kayhan Respiratory and cardiac interoceptive sensitivity in the first two years of life Journal Article In: eLife, vol. 12, pp. 1–40, 2025. @article{Tuente2025,Several recent theoretical accounts have posited that interoception, the perception of internal bodily signals, plays a vital role in early human development. Yet, empirical evidence of cardiac interoceptive sensitivity in infants to date has been mixed. Furthermore, existing evidence does not go beyond the perception of cardiac signals and focuses only on the age of 5–7 mo, limiting the generalizability of the results. Here, we used a modified version of the cardiac interoceptive sensitivity paradigm introduced by Maister et al., 2017 in 3-, 9-, and 18-mo-old infants using cross-sectional and longitudinal approaches. Going beyond, we introduce a novel experimental paradigm, namely the iBREATH, to investigate respiratory interoceptive sensitivity in infants. Overall, for cardiac interoceptive sensitivity ( total n =135) we find rather stable evidence across ages with infants on average preferring stimuli presented synchronously to their heartbeat. For respiratory interoceptive sensitivity ( total n =120) our results show a similar pattern in the first year of life, but not at 18 mo. We did not observe a strong relationship between cardiac and respiratory interoceptive sensitivity at 3 and 9 mo but found some evidence for a relationship at 18 mo. We validated our results using specification curve- and mega-analytic approaches. By examining early cardiac and respiratory interoceptive processing, we provide evidence that infants are sensitive to their interoceptive signals. |
Dirk Moorselaar; Ningkai Wang; Jan Theeuwes Differential neural mechanisms underlying inhibition of color and dynamic motion distractors Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 3, pp. 543–554, 2025. @article{Moorselaar2025a,Navigating visually complex environments requires focusing on relevant information while filtering out (salient) distractions. The signal suppression hypothesis posits that salient stimuli generate an automatic saliency signal that captures attention unless overridden by learned suppression mechanisms. In support of this, ERP studies have demonstrated that salient stimuli that do not capture attention elicit a distractor positivity (PD), a putative neural index of suppression. Yet, to date, this hypothesis has been primarily tested with color singletons, leaving it unclear if the PD reflects general suppression or is specific to color singletons. This study compared lateralized ERPs elicited by color singleton and dynamic motion distractors using a variant of the additional singleton paradigm that has been shown to result in proactive suppression of colored distractors. Behavioral results showed a singleton presence benefit for both distractor types, indicating distractor suppression. However, ERP data revealed clear differences in the underlying neural mechanisms: Color singletons elicited a PD component indicative of proactive suppression, whereas motion singletons elicited a later positivity preceded by an N2pc, suggesting reactive suppression. Our findings suggest that motion singletons, unlike color singletons, are suppressed reactively after initial capture. This study highlights the importance of considering distractor feature dimensions in understanding attentional suppression mechanisms and underscores the need for caution in establishing proactive suppression based on a single metric. Further research is needed to clarify the conditions under which the early PD reliably indicates proactive suppression and to explore the neural processes underlying the suppression of various salient distractors. |
Monica Vanoncini; Ezgi Kayhan; Birgit Elsner; Moritz Wunderwald; Sebastian Wallot; Stefanie Hoehl; Natalie Boll-Avetisyan Individual differences in infants' speech segmentation performance: The role of mother-infant cardiac synchrony Journal Article In: Infancy, vol. 30, no. 2, pp. 1–13, 2025. @article{Vanoncini2025,Caregiver-infant coregulation is an early form of communication. This study investigated whether mother-infant biological coregulation is associated with 9-month-olds' word segmentation performance, a crucial milestone predicting language development. We hypothesized that coregulation would relate with infants' word segmentation performance. Additionally, we examined whether this relationship is influenced by the caregiving environment (i.e., parental reflective functioning) and the infant's emotional state (i.e., positive affect). Coregulation was investigated via cardiac synchrony in 28 nine-month-old infants (16 females) during a 5-min free-play with their German-speaking mothers. Cardiac synchrony was measured through Respiratory Sinus Arrhythmia (RSA), employing Recurrence Quantification Analysis to evaluate dyadic coupling (i.e., Recurrence Rate) and dyadic predictability (i.e., Entropy). Infants' word segmentation was measured with an eye-tracking central-fixation procedure. A stepwise regression revealed that higher dyadic coupling, but not predictability, of the dyads' RSA was associated with infants looking longer toward the screen when listening to novel as compared to familiar test words, indicating advanced word segmentation performance (Cohen's d = 0.25). Moreover, cardiac synchrony correlated positively with maternal sensitivity to their infant's mental states, but not with the infant's positive affect. These results suggest that caregiver-infant biological coregulation may play a foundational role in language acquisition. |
Britta U. Westner; Ella Bosch; Christian Utzerath; Jan Buitelaar; Floris P. Lange Typical neural adaptation for familiar images in autistic adolescents Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–15, 2025. @article{Westner2025,It has been proposed that autistic perception may be marked by a reduced influence of temporal context. Following this theory, prior exposure to a stimulus should lead to a weaker or absent alteration of the behavioral and neural response to the stimulus in autism, compared with a typical population. To examine these hypotheses, we recruited two samples of human volunteers: a student sample (N = 26), which we used to establish our analysis pipeline, and an adolescent sample (N = 36), which consisted of a group of autistic (N = 18) and a group of non-autistic (N = 18) participants. All participants were presented with visual stimulus streams consisting of novel and familiar image pairs, while they attentively monitored each stream. We recorded task performance and used magnetoencephalography (MEG) to measure neural responses, and to compare the responses with familiar and novel images. We found behavioral facilitation as well as a reduction of event-related field (ERF) amplitude for familiar, compared with novel, images in both samples. Crucially, we found statistical evidence against between-group effects of familiarity on both behavioral and neural responses in the adolescent sample, suggesting that the influence of familiarity is comparable between autistic and non-autistic adolescents. These findings challenge the notion that perception in autism is marked by a reduced influence of prior exposure. |
Yuting Xu; Ayumu Yamashita; Kyuto Uno; Tomoya Kawashima; Kaoru Amano Prediction of alpha power using multiple subjective measures and autonomic responses Journal Article In: Psychophysiology, vol. 62, no. 3, pp. 1–13, 2025. @article{Xu2025d,Alpha oscillations are associated with various cognitive functions. However, the determinants of alpha power variation remain ambiguous, primarily due to its inconsistent associations with autonomic responses and subjective states under different experimental conditions. To thoroughly examine the correlations between alpha power variation and these factors, we implemented a range of experimental conditions, encompassing attentional and emotional tasks, as well as a resting-state. In addition to the electroencephalogram data, we gathered a suite of autonomic response measurements and subjective ratings. We employed multiple linear regression analysis, utilizing autonomic responses and subjective reports as predictors of alpha power. We also subtracted the aperiodic components for better estimation of the power of periodic alpha oscillations. Our results from two separately conducted experiments robustly demonstrated that the combined use of autonomic response measurements and subjective ratings effectively predicted the parietal-occipital periodic alpha power variation across a range of conditions. These predictions were supported by leave-one-participant-out cross-validation and cross-experiment validation, confirming that multiple linear relationships can be generalized to new participants. This study demonstrates the links of alpha power variations with autonomic responses and subjective states, suggesting that during investigations of the cognitive functions of alpha oscillations, it is important to consider the potential influences of autonomic responses and subjective states on alpha oscillations. |
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, vol. 44, 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 |
Xuan Li; Simon B. Eickhoff; Susanne Weis Stimulus selection influences prediction of individual phenotypes in naturalistic conditions Journal Article In: Human Brain Mapping, vol. 46, no. 3, pp. 1–22, 2025. @article{Li2025p,While the use of naturalistic stimuli such as movie clips for understanding individual differences and brain–behaviour relationships attracts increasing interest, the influence of stimulus selection remains largely unclear. By using machine learning to predict individual traits (phenotypes) from brain activity evoked during various movie clips, we show that different movie stimuli can result in distinct prediction performances. In brain regions related to lower-level processing of the stimulus, prediction to a certain degree benefits from stronger synchronisation of brain activity across subjects. By contrast, better predictions in frontoparietal brain regions are mainly associated with larger inter-subject variability. Furthermore, we demonstrate that while movie clips with rich social content in general achieve better predictions, the importance of specific movie features for prediction highly depends on the phenotype under investigation. Overall, our findings underscore the importance of careful stimulus selection and provide novel insights into stimulus selection for phenotype prediction in naturalistic conditions, opening new avenues for future research. |
Timo Kerkoerle; Louise Pape; Milad Ekramnia; Xiaoxia Feng; Jordy Tasserie; Morgan Dupont; Xiaolian Li; Bechir Béchir Jarraya; Wim Vanduffel; Stanislas Dehaene; Ghislaine Dehaene-Lambertz Brain mechanisms of reversible symbolic reference: A potential singularity of the human brain Journal Article In: eLife, vol. 12, pp. 1–28, 2025. @article{Kerkoerle2025,The emergence of symbolic thinking has been proposed as a dominant cognitive criterion to distinguish humans from other primates during hominization. Although the proper definition of a symbol has been the subject of much debate, one of its simplest features is bidirectional attachment: the content is accessible from the symbol, and vice versa. Behavioral observations scattered over the past four decades suggest that this criterion might not be met in non-human primates, as they fail to generalize an association learned in one temporal order (A to B) to the reverse order (B to A). Here, we designed an implicit fMRI test to investigate the neural mechanisms of arbitrary audio-visual and visual-visual pairing in monkeys and humans and probe their spontaneous reversibility. After learning a unidirectional association, humans showed surprise signals when this learned association was violated. Crucially, this effect occurred spontaneously in both learned and reversed directions, within an extended network of high-level brain areas, including, but also going beyond the language network. In monkeys, by contrast, violations of association effects occurred solely in the learned direction and were largely confined to sensory areas. We propose that a human-specific brain network may have evolved the capacity for reversible symbolic reference. ### Competing Interest Statement The authors have declared no competing interest. |
Xizi Gong; Tao He; Qian Wang; Junshi Lu; Fang Fang Time course of orientation ensemble representation in the human brain Journal Article In: The Journal of Neuroscience, vol. 45, no. 7, pp. 1–13, 2025. @article{Gong2025,Natural scenes are filled with groups of similar items. Humans employ ensemble coding to extract the summary statistical information of the environment, thereby enhancing the efficiency of information processing, something particularly useful when observing natural scenes. However, the neural mechanisms underlying the representation ofensemble information in the brain remain elusive. In particular, whether ensemble representation results from the mere summation of individual item representations or it engages other specific processes remains unclear. In this study, we utilized a set of orientation ensembles wherein none ofthe individual item orientations were the same as the ensemble orientation. We recorded magnetoencephalography (MEG) signals from human participants (both sexes) when they performed an ensemble orientation discrimination task. Time-resolved multivariate pattern analysis (MVPA) and the inverted encoding model (IEM) were employed to unravel the neural mechanisms of the ensemble orientation representation and track its time course. First, we achieved successful decoding of the ensemble orientation, with a high correlation between the decoding and behavioral accuracies. Second, the IEM analysis demonstrated that the representation of the ensemble orientation differed from the sum of the representations of individual item orientations, suggesting that ensemble coding could fur- ther modulate orientation representation in the brain. Moreover, using source reconstruction, we showed that the representation of ensemble orientation manifested in early visual areas. Taken together, our findings reveal the emergence of the ensemble representation in the human visual cortex and advance the understanding of how the brain captures and represents ensemble information. |
Changrun Huang; Dirk Moorselaar; Joshua Foster; Mieke Donk; Jan Theeuwes Neural mechanisms of learned suppression uncovered by probing the hidden attentional priority map Journal Article In: eLife, vol. 13, pp. 1–21, 2025. @article{Huang2025b,Attentional capture by an irrelevant salient distractor is attenuated when the distractor appears more frequently in one location, suggesting learned suppression of that location. However, it remains unclear whether suppression is proactive (before attention is directed) or reactive (after attention is allocated). Here, we investigated this using a ‘pinging' technique to probe the attentional distribution before search onset. In an EEG experiment, participants searched for a shape singleton while ignoring a color singleton distractor at a high-probability location. To reveal the hidden attentional priority map, participants also performed a continuous recall spatial memory task, with a neutral placeholder display presented before search onset. Behaviorally, search was more efficient when the distractor appeared at the high-probability location. Inverted encoding analysis of EEG data showed tuning profiles that decayed during memory maintenance but were revived by the placeholder display. Notably, tuning was most pronounced at the to-be-suppressed location, suggesting initial spatial selection followed by suppression. These findings suggest that learned distractor suppression is a reactive process, providing new insights into learned spatial distractor suppression mechanisms. |
Rose Nasrawi; Mika Mautner-Rohde; Freek Ede Memory load influences our preparedness to act on visual representations in working memory without affecting their accessibility Journal Article In: Progress in Neurobiology, vol. 245, pp. 1–11, 2025. @article{Nasrawi2025,It is well established that when we hold more content in working memory, we are slower to act upon part of that content when it becomes relevant for behavior. Here, we asked whether this load-related slowing is due to slower access to the sensory representations held in working memory (as predicted by serial working-memory search), or by a reduced preparedness to act upon those sensory representations once accessed. To address this, we designed a visual-motor working-memory task in which participants memorized the orientation of two or four colored bars, of which one was cued for reproduction. We independently tracked EEG markers associated with the selection of visual (cued item location) and motor (relevant manual action) information from the EEG time-frequency signal, and compared their latencies as a function of memory load. We confirm slower memory-guided behavior with higher working-memory load and show that this is associated with delayed motor selection. In contrast, we find no evidence for a concomitant delay in the latency of visual selection. Moreover, we show that variability in decision times within each memory-load condition is associated with corresponding changes in the latency of motor, but not visual selection. These results reveal how memory load affects our preparedness to act on sensory representations in working memory, while leaving sensory access itself unaffected. This posits action readiness as a key factor that shapes the speed of memory-guided behavior and that underlies delayed responding with higher working-memory load. |
Camila Dias; Teresa Sousa; Susana Mouga; Miguel Castelo-Branco A link between error monitoring and basal ganglia-dependent learning in autism Journal Article In: Brain Communications, vol. 7, no. 6, pp. 1–18, 2025. @article{Dias2025,Altered error monitoring mechanisms may contribute to the neurobehavioral manifestations of autism, particularly those affecting trial-and-error learning. In this study, we examined how error-related neural responses relate to learning processes in autistic individuals. To address this, we employed a paradigm designed to assess both error monitoring and the learning of socioemotional orienting cues. Fifteen autistic and fifteen non-autistic adult males performed a challenging functional MRI task that required the integration of facial expression and gaze cues to extract the correct social-orienting direction. We evaluated how error-related neural responses evolved with learning and compared these dynamics across groups. We hypothesized differences in key error monitoring regions—the anterior cingulate cortex and anterior insula—and in the putamen, which is implicated in trial-and-error learning. Behaviourally, both groups exhibited similar learning rates, but the autistic group showed significantly lower accuracy (P = 9.60 × 10−5). At the neural level, although the progression of error-related activity throughout learning was similar across both groups, autistic individuals demonstrated attenuated differentiation between correct responses and errors in the dorsal anterior cingulate cortex (F(2, 53.23) = 9.68 |
Goi Khia Eng; Arielle Tambini; Molly S. Hermiller; Nicolette Recchia; Jeanmarie R. Harvey; Dan V. Iosifescu; Russell H. Tobe; Emily R. Stern Personalized non-invasive neuromodulation for sensory-based urge suppression in individuals with OCD: A proof-of-concept investigation Journal Article In: Frontiers in Human Neuroscience, vol. 19, pp. 1–9, 2025. @article{Eng2025,Obsessive-compulsive disorder (OCD) is chronic and impairing. While OCD often involves fear of harm or bad events, many patients experience “sensory phenomena,” which are aversive sensory experiences that drive repetitive behaviors regardless of specific fears. Standard treatments do not effectively address sensory phenomena, and novel approaches are needed. Transcranial magnetic stimulation (TMS) is a safe and non-invasive neuromodulation technique increasingly used in psychiatric disorders, including OCD. This work presents a data-driven approach to identifying TMS brain targets for modulating sensory urges in OCD incorporating both behavioral and clinical criteria (Study 1) for a proof-of-concept investigation (Study 2). Study 1 included 69 individuals with OCD and 23 controls who completed an urges-for-action fMRI task involving instructed eyeblink suppression as an experimental model for sensory-based urges. Data-driven conjunction analysis revealed several brain regions, including the right postcentral gyrus, that were associated with more blink suppression failure (behavioral), more severe sensory phenomena (clinical), and were hyperactivated in OCD patients compared to controls. Study 2 administered single-session inhibitory TMS on 4 returning OCD patients using individualized targets within the postcentral gyrus identified from Study 1. Compared to sham, inhibitory TMS delivered to individualized postcentral gyrus targets resulted in fewer blink suppression failures, reduced activation in the target (postcentral gyrus) and key urge-related areas (insula, mid-cingulate), and greater reduction in self-reported urge to engage in OCD-related compulsions, with medium to large effect sizes. These findings demonstrate the potential of utilizing data-driven approaches incorporating behavioral and clinical criteria to target hard-to-treat sensory phenomena in OCD. |
Juyoen Hur; Rachael M. Tillman; Hyung Cho Kim3; Paige Didier; Allegra S. Anderson; Samiha Islam; Melissa D. Stockbridge; Andres De Los Reyes; Kathryn A. DeYoung; Jason F. Smith; Alexander J. Shackman In: Journal of Psychopathology and Clinical Science, vol. 134, no. 1, pp. 41–56, 2025. @article{Hur2025,Social anxiety-which typically emerges in adolescence-lies on a continuum and, when extreme, can be devastating. Socially anxious individuals are prone to heightened fear, anxiety, and the avoidance of contexts associated with potential social scrutiny. Yet most neuroimaging research has focused on acute social threat. Much less attention has been devoted to understanding the neural systems recruited during the uncertain anticipation of potential encounters with social threat. Here we used a novel fMRI paradigm to probe the neural circuitry engaged during the anticipation and acute presentation of threatening faces and voices in a racially diverse sample of 66 adolescents selectively recruited to encompass a range of social anxiety and enriched for clinically significant levels of distress and impairment. Results demonstrated that adolescents with more severe social anxiety symptoms experience heightened distress when anticipating encounters with social threat, and reduced discrimination of uncertain social threat and safety in the bed nucleus of the stria terminalis (BST), a key division of the central extended amygdala (EAc). Although the EAc-including the BST and central nucleus of the amygdala-was robustly engaged by the acute presentation of threatening faces and voices, the degree of EAc engagement was unrelated to the severity of social anxiety. Together, these observations provide a neurobiologically grounded framework for conceptualizing adolescent social anxiety and set the stage for the kinds of prospective-longitudinal and mechanistic research that will be necessary to determine causation and, ultimately, to develop improved interventions for this often-debilitating illness. |
Michaela Klímová; Ilona M. Bloem; Sam Ling How does orientation-tuned normalization spread across the visual field? Journal Article In: Journal of Neurophysiology, vol. 133, no. 2, pp. 539–546, 2025. @article{Klimova2025,Visuocortical responses are regulated by gain control mechanisms, giving rise to fundamental neural and perceptual phenomena such as surround suppression. Suppression strength, determined by the composition and relative properties of stimuli, controls the strength of neural responses in early visual cortex, and in turn, the subjective salience of the visual stimulus. Notably, suppression strength is modulated by feature similarity; for instance, responses to a center-surround stimulus in which the components are collinear to each other are weaker than when they are orthogonal. However, this feature-tuned aspect of normalization, and how it may affect the gain of responses, has been understudied. Here, we examine the contribution of the tuned component of suppression to contrast response modulations across the visual field. To do so, we used functional magnetic resonance imaging (fMRI) to measure contrast response functions (CRFs) in early visual cortex (areas V1–V3) in 10 observers while they viewed full-field center-surround gratings. The center stimulus varied in contrast between 2.67% and 96% and was surrounded by a collinear or orthogonal surround at full contrast. We found substantially stronger suppression of responses when the surround was parallel to the center, manifesting as shifts in the population CRF. The magnitude of the CRF shift was strongly dependent on voxel spatial preference and seen primarily in voxels whose receptive field spatial preference corresponds to the area straddling the center-surround boundary in our display, with little-to-no modulation elsewhere. |
Heather L. Kosakowski; Jingnan Du; Vaibhav Tripathi; Mark C. Eldaief; Randy L. Buckner Ventral striatum is preferentially correlated with the salience network including regions in dorsolateral prefrontal cortex Journal Article In: Journal of Neurophysiology, vol. 134, no. 1, pp. 193–215, 2025. @article{Kosakowski2025,The ventral striatum (VS) receives input from the cerebral cortex and is modulated by midbrain dopaminergic projections in support of processing reward and motivation. Here, we explored the organization of cortical regions linked to the human VS using within-individual functional connectivity MRI (fcMRI) in intensively scanned participants. In two initial participants (scanned 31 sessions each), seed regions in the VS were preferentially correlated with distributed cortical regions that are part of the salience network. The VS seed regions recapitulated salience network topography and replicated in each individual, including anterior and posterior midline regions, anterior insula, and dorsolateral prefrontal cortex (DLPFC). The topography was distinct from adjacent striatal seed regions and from cortical networks associated with domain-flexible cognitive control. Unbiased comprehensive analyses of the full striatum confirmed that the VS is coupled to the salience network while also revealing the established, spatially separated cognitive zones of the caudate and motor zones of the putamen. VS correlation with the salience network, including DLPFC, was observed in 15 additional participants (scanned 8 or more times each), indicating it is a robust and generalizable finding. These results suggest that the VS contributes to a cortico-basal ganglia loop that is part of the salience network and raise the possibility that the DLPFC may be an effective neuromodulatory target for neuropsychiatric disorders of reward and motivation because of its preferential coupling to the VS. |
Mrinmayi Kulkarni; Lydia Jiang; Jessica Robin; Jung Won Choi; Bradley R. Buchsbaum; Rosanna K. Olsen Scene-sensitive medial temporal lobe subregions are recruited for the integration of non-scene stimuli Journal Article In: Journal of Cognitive Neuroscience, vol. 38, no. 1, pp. 100–125, 2025. @article{Kulkarni2025,A hallmark feature of episodic memory is the ability to flexibly recombine information across episodes to form new associations and guide behavior. This process, termed associative inference, relies on the hippocampus and surrounding medial temporal lobe (MTL) subregions. We previously found that cross-episode binding was improved when episodes were linked by scenes rather than by faces or objects. Here, we tested whether differential recruitment of category-sensitive MTL subregions underlies these behavioral differences. Participants completed study-test phases of the Associative Inference in Memory task while undergoing fMRI scanning. During the study phase, they encoded overlapping AB and BC pairs. A and C items were always objects. The linking B item was either a face or a scene. At test, memory for the direct (AB, BC) and indirect associations (inferred AC) was tested. Category sensitivity in MTL subregions was tested using an independent functional localizer and the low integration (AB) trials from the study phase of the Associative Inference in Memory task. Within the MTL, no subregions exhibited face sensitivity. The anterior hippocampal head, anterolateral and posteromedial entorhinal cortices, and parahippocampal cortex were identified as scene sensitive. Although accuracy of the indirect inferences did not differ between pairs linked by faces and scenes, MTL subregion recruitment differed across categories. Scene-sensitive subregions in MTL cortex (anterolateral entorhinal cortex, posteromedial entorhinal cortex, and parahippocampal cortex), but not the hippocampus (anterior hippocampal head), were recruited to support associative inference for faces during encoding. These findings suggest that regions in MTL cortex identified as scene sensitive here may be involved in integrating disparate elements of episodes into coherent representations, and may be recruited for non-scene stimuli when integration demands during encoding are high (e.g., during associative inference). |
Hsin Hung Li; Thomas C. Sprague; Aspen H. Yoo; Wei Ji Ma; Clayton E. Curtis Neural mechanisms of resource allocation in working memory Journal Article In: Science Advances, vol. 11, no. 15, pp. 1–14, 2025. @article{Li2025d,To mitigate capacity limits of working memory, people allocate resources according to an item's relevance. However, the neural mechanisms supporting such a critical operation remain unknown. Here, we developed computational neuroimaging methods to decode and demix neural responses associated with multiple items in working memory with different priorities. In striate and extrastriate cortex, the gain of neural responses tracked the priority of memoranda. We decoded higher-priority memoranda with smaller error and lower uncertainty. Moreover, these neural differences predicted behavioral differences in memory prioritization between and within participants. Trial-wise variability in the magnitude of delay activity in the frontal cortex predicted differences in decoded precision between low- and high-priority items in visual cortex. These results support a model in which feedback signals broadcast from frontal cortex sculpt the gain of memory representations in the visual cortex according to behavioral relevance, thus identifying a neural mechanism for resource allocation. |
Milena I. Mihovilovic; Thomas Stephan; Andreas Straube; Marianne Dieterich; Thomas Eggert Brain activity during acquisition of long visuospatial sequences Journal Article In: Frontiers in Cognition, vol. 4, pp. 1–15, 2025. @article{Mihovilovic2025,Explicitly acquiring a visuospatial sequence involves various fundamental attentional and processing mechanisms that can be difficult to disentangle. To this end, we performed an fMRI study (n = 34) on the acquisition of visuospatial targets in a delayed imitation paradigm. Task phases alternated between presentation and recall of a 20-target-long sequence. Behavioral data from the recall phase was used to determine encoding progress as a function of time during presentation, with this progress taken as a continuous predictor of BOLD activity. A separate, attention-only task was devised in order to isolate activity related to spatial attention shifts specifically. General linear model analysis using the constructed learning and attention predictors revealed heightened activation for both tasks in bilateral superior parietal lobules (SPL), bilateral V5, and bilateral middle frontal gyri (MFG). Increased response during learning was seen in the SPL and V5, but not MFG. Repeated measures ANOVA indicated significant interactions between region and task, as well as a right-biased tendency in the hemisphere*task interaction. This suggests a role for the SPL and V5 during sequence acquisition that cannot be explained by attention alone. |
William Narhi-Martinez; Yong Min Choi; Blaire Dube; Julie D. Golomb Allocation of spatial attention in human visual cortex as a function of endogenous cue validity Journal Article In: Cortex, vol. 185, pp. 4–19, 2025. @article{NarhiMartinez2025,Several areas of visual cortex contain retinotopic maps of the visual field, and neuroimaging studies have shown that covert attentional guidance will result in increases of activity within the regions representing attended locations. However, little research has been done to directly compare neural activity for different types of attentional cues. Here, we used fMRI to investigate how retinotopically-specific cortical activity would be modulated depending on whether we provided deterministic or probabilistic spatial information. On each trial, a four-item memory array was presented and participants' memory for one of the items would later be probed. Critically, trials began with a foveally-presented endogenous cue that was either 100% valid (deterministic runs), 70% valid (probabilistic runs), or neutral. By dividing visual cortex into quadrant-specific regions of interest (qROIs), we could examine how attention was spatially distributed across the visual field within each trial, depending on cue type and delay. During the anticipatory period prior to the memory array, we found increased activation at the cued location compared to noncued locations, with surprisingly comparable levels of facilitation for both deterministic and probabilistic cues. However, we found significantly greater facilitation on deterministic relative to probabilistic trials following the onset of the memory array, with only deterministic cue-related facilitation persisting through the presentation of the probe. These findings reveal how cue validity can drive differential allocations of neural resources over time across cued and noncued locations, and that the allocation of attention should not be assumed to invariably scale alongside the validity of a cue. |
Vincent Plikat; Pablo R. Grassi; Julius Frack; Andreas Bartels; Vincent Plikata; Pablo R. Grassia; Julius Frackd; Andreas Bartels Hierarchical surprise signals in naturalistic violation of expectations Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–23, 2025. @article{Plikat2025,Surprise responses signal both high-level cognitive alerts that information is missing, and increasingly specific back-propagating error signals that allow updates in processing nodes. Studying surprise is, hence, central for cognitive neuroscience to understand internal world representations and learning. Yet, only few prior studies used naturalistic stimuli targeting our high-level understanding of the world. Here, we use magic tricks in an fMRI experiment to investigate neural responses to violations of core assumptions held by humans about the world. We showed participants naturalistic videos of three types of magic tricks, involving objects appearing, changing color, or disappearing, along with control videos without any violation of expectation. Importantly, the same videos were presented with and without prior knowledge about the tricks' explanation. Results revealed generic responses in frontal and parietal areas, together with responses specific to each of the three trick types in posterior sensory areas. A subset of these regions, the midline areas of the default mode network (DMN), showed surprise activity that depended on prior knowledge. Equally, sensory regions showed sensitivity to prior knowledge, reflected in differing decoding accuracies. These results suggest a hierarchy of surprise signals involving generic processing of violation of expectations in frontal and parietal areas with concurrent surprise signals in sensory regions that are specific to the processed features. |
Luke Priestley; Mark Chiew; Mo Shahdloo; Ali Mahmoodi; Xinghao Cheng; Robin Cleveland; Matthew Rushworth; Nima Khalighinejad Dorsal raphe nucleus controls motivation-state transitions in monkeys Journal Article In: Science Advances, vol. 11, no. 26, pp. 1–20, 2025. @article{Priestley2025,The dorsal raphe nucleus (DRN) is an important source of serotonin in the brain, but fundamental aspects of its function remain elusive. Here, we present a combination of minimally invasive recording and disruption studies to show that DRN brings about changes in motivation states. We use recently developed methods for identifying temporal patterns in behavior to show that monkeys change their motivation depending on the availability of rewards in the environment. Distinctive patterns of DRN activity occur when monkeys transition between a high-motivation state occupied when rewards are abundant, to a low-motivation state engendered by reward scarcity. Disrupting DRN diminishes sensitivity to the reward environment and perturbs transitions in motivational states. |
Ayushi Sangoi; Farzin Hajebrahimi; Suril Gohel; Mitchell Scheiman; Tara L. Alvarez Efferent compared to afferent neural substrates of the vergence eye movement system evoked via fMRI Journal Article In: Frontiers in Neuroscience, vol. 18, pp. 1–13, 2025. @article{Sangoi2025,Introduction: The vergence neural system was stimulated to dissect the afferent and efferent components of symmetrical vergence eye movement step responses. The hypothesis tested was whether the afferent regions of interest would differ from the efferent regions to serve as comparative data for future clinical patient population studies. Methods: Thirty binocularly normal participants participated in an oculomotor symmetrical vergence step block task within a functional MRI experiment compared to a similar sensory task where the participants did not elicit vergence eye movements. Results: For the oculomotor vergence task, functional activation was observed within the parietal eye field, supplemental eye field, frontal eye field, and cerebellar vermis, and activation in these regions was significantly diminished during the sensory task. Differences between the afferent sensory and efferent oculomotor experiments were also observed within the visual cortex. Discussion: Differences between the vergence oculomotor and sensory tasks provide a protocol to delineate the afferent and efferent portion of the vergence neural circuit. Implications with clinical populations and future therapeutic intervention studies are discussed. |
Ayushi Sangoi; Farzin Hajebrahimi; Suril Gohel; Mitchell Scheiman; Arlene Goodman; Melissa Noble; Tara L. Alvarez In: Frontiers in Neuroscience, vol. 19, pp. 1–13, 2025. @article{Sangoi2025a,Introduction: The CONCUSS clinical trial examined the neural changes associated with office-based vergence/accommodative therapy with movement (OBVAM) in concussion-related convergence insufficiency (CONC-CI). Methods: The following assessments were collected at baseline and post-OBVAM therapy: activity evoked from a functional MRI vergence oculomotor task, near point of convergence (NPC), positive fusional vergence (PFV), vergence facility (VF), and visual symptoms from a sensorimotor vision exam. Fifty-four CONC-CI participants, diagnosed with persisting concussion symptoms between one- and six-months post-injury, were analyzed in group-level results. Results: Functional activity in the vergence oculomotor network, specifically the frontal eye fields, supplemental eye fields, parietal eye fields, cerebellar vermis (CV), and visual cortex, increased post-OBVAM compared to baseline assessments. Significant increases in post-OBVAM compared to baseline assessments were observed in the visual cortex (bilateral V3 and right area PH) and the CV, via a paired t-test with family-wise error corrected for multiple comparisons (p < 0.05). The pooled baseline and post-OBVAM measures revealed that the bilateral functional activities of V3 and CV were significantly correlated with the NPC, PFV, and VF clinical signs, and the right hemisphere area PH within the visual cortex was significantly correlated with VF (Bonferroni-corrected; p < 0.001). To determine whether the CONC-CI post-OBVAM functional brain activity differed from that of the binocularly normal control (BNC) data, an unpaired t-test was performed comparing 46 age-matched BNC datasets with 54 CONC-CI datasets. Significant differences in functional activity between BNC and CONC-CI at post-OBVAM datasets were not observed (p > 0.05). Discussion: Results support that OBVAM improves functional brain activity in CONC-CI correlated with NPC, PFV, and VF. |
Sarah Schuster; Kim Lara Weiss; Florian Hutzler; Martin Kronbichler; Stefan Hawelka Interactive and additive effects of word frequency and predictability: A fixation-related fMRI study Journal Article In: Brain and Language, vol. 260, pp. 1–7, 2025. @article{Schuster2025,The effects of word frequency and predictability are informative with respect to bottom-up and top-down mechanisms during reading. Word frequency is assumed to index bottom-up, whereas word predictability top-down information. Findings regarding potential interactive effects, however, are inconclusive. An interactive effect would suggest an early lexical impact of contextual top-down mechanisms where both variables are processed concurrently in early stages of word recognition. An additive effect, to the contrary, would suggest that contextual top-down processing only occurs post-lexically. We evaluated potential interactions between word frequency and predictability during silent reading by means of functional magnetic resonance imaging and simultaneous eye-tracking (i.e., fixation-related fMRI). Our data revealed exclusively additive effects. Specifically, we observed effects of word frequency and word predictability in left inferior frontal regions, whereas word frequency additionally exhibited an effect in the left occipito-temporal cortex. We interpret our findings in terms of contextual top-down processing facilitation. |
Leslie Tricoche; Marion Royer D'Halluin; Martine Meunier; Denis Pélisson Neural bases of social facilitation and inhibition: How peer presence affects elementary eye movements Journal Article In: Social Cognitive and Affective Neuroscience, vol. 19, no. 1, pp. 1–16, 2025. @article{Tricoche2025,Social facilitation/inhibition (SFI) refers to how others' presence influences task performance positively or negatively. Our previous study revealed that peer presence modulated saccadic eye movements, a fundamental sensorimotor activity. Pro- and antisaccades were either facilitated or inhibited depending on trial block complexity Tricoche L, Ferrand-Verdejo J, Pélisson D et al. (Peer Presence Effects on Eye Movements and Attentional Performance. Front Behav Neurosci 2020;13:1–13. https://doi.org/10.3389/fnbeh.2019.00280). In the present fMRI study, we adapted our paradigm to investigate the neural basis of SFI on saccades. Considering inter- and intra-individual variabilities, we evaluated the shared and distinct neural patterns between social facilitation and inhibition. We predicted an involvement of the saccade-related and attention networks, alongside the Theory-of-Mind (ToM) network, with opposite activity changes between facilitation and inhibition. Results confirmed peer presence modulation in frontoparietal areas related to saccades and attention, in opposite directions for facilitation and inhibition. Additionally, the ventral attention network was modulated during inhibition. Default mode regions, including ToM areas, were also modulated. Finally, pupil size, often linked to arousal, increased with peers and correlated with dorsal attention regions and anterior insula activities. These results suggest that SFI engages task-specific and domain-general networks, modulated differently based on observed social effect. Attention network seemed to play a central role at both basic (linked to arousal or vigilance) and cognitive control levels. |
Janina Von Der Gablentz; Andreas Sprenger; Nina Overbeeke; Dagmar Timmann; Norbert Brüggemann; Christoph Helmchen Dissociable effects of medication on visual-vestibular brain excitability by visual motion stimuli in episodic ataxia type 2 Journal Article In: Brain Communications, vol. 7, no. 5, pp. 1–13, 2025. @article{VonDerGablentz2025,The clinical hallmark of episodic ataxia type 2 (EA2) consists of episodes of recurrent severe vestibulo-cerebellar dysfunction, characterized by marked postural unsteadiness and oscillopsia. Triggering factors of EA2 attacks, such as physical exertion and sensory stimulation, the high comorbidity with migraine, and the increased risk of epilepsy in EA2 suggest abnormal brain excitability. To investigate this, we assessed brain excitability in response to visual (checkerboard) and visual motion (optic flow) stimuli using interictal functional magnetic resonance imaging. Visual stimulation elicited strong bilateral neural activity in the primary visual cortex (V1-V3) and in motion-sensitive visual areas (V5) in 21 EA2 patients and 21 age-matched healthy participants (HP). Compared to HP, EA2 patients revealed decreased activity in the primary visual cortex (V1), cerebellar Crus I and II and caudal vermis but increased activation of multisensory vestibular processing areas (posterior insula, superior temporal and supramarginal gyrus, inferior parietal lobe). Interestingly, the abnormal excitability in the vestibular processing cortex areas was primarily found in patients without medication (4-aminopyridine, acetazolamide) but hardly seen in patients on medication. Our findings in treatment-naïve patients reflect disease-inherent changes in visual cortical excitability in EA2, which may be reversible through anti-episodic medication. As excitability by visual motion stimuli in multi-sensory vestibular processing cortical areas was largely found in patients on medication it may also indicate an inhibitory effect on the physiological reciprocal inhibitory visual-vestibular interaction as a multisensory mechanism for self-motion perception: the annoying oscillopsia of EA2 patients is counterbalanced by decreased visual cortex activity and hence smaller inhibition of the vestibular cortex. |
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. |
Ayelet Arazi; Alessandro Toso; Tineke Grent-‘t-Jong; Peter J. Uhlhaas; Tobias H. Donner Large-scale maps of altered cortical dynamics in early-stage psychosis are related to GABAergic and glutamatergic neurotransmission Journal Article In: Science Advances, vol. 11, no. 33, pp. 1–19, 2025. @article{Arazi2025,Psychotic disorders affect GABAergic inhibition and glutamatergic excitation via NMDA receptors across the cerebral cortex. The mechanisms by which these distributed synaptic alterations produce the heterogenous symptoms of psychosis remain poorly understood. Using magnetoencephalographical source imaging, we mapped psychosis-related alterations of various features of intrinsic neural population dynamics across the human cortex. The cortex-wide patterns of these features were highly reproducible and related to the anatomical hierarchy of cortical areas. We found similar changes in these patterns for individuals with a first episode of psychosis and those at clinical high risk for psychosis. Maps of psychosis-induced changes in dynamics resembled the maps of GABA-A receptor densities and of pharmacological GABA-A or NMDA manipulation effects on cortical dynamics in healthy participants. The level of pattern similarity to GABA-A manipulation effects in individual patients correlated with positive symptoms, while the pattern similarity to NMDA effects correlated with negative symptoms. Our results open up a window on the distributed mechanisms of psychotic symptoms. |
Cemre Baykan; Alexander C. Schütz Electroencephalographic responses to the number of objects in partially occluded and uncovered scenes Journal Article In: Journal of Cognitive 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. |
Merit Bruckmaier; Artyom Zinchenko; Hermann J. Müller; Thomas Geyer Increasing signal, reducing noise: Contrasting neural mechanisms of attention in visual search Journal Article In: Journal of Cognitive Neuroscience, vol. 38, no. 1, pp. 89–99, 2025. @article{Bruckmaier2025,When invariant target–distractor arrays are presented repeatedly during visual search, participants respond faster on repeated versus novel configuration trials. This effect reflects attentional guidance through long-term memory (LTM) templates—a phenomenon termed contextual cueing. Subsequently, relocating the target within the same distractor layout abolishes any contextual cueing effects, and relearning the new target position is much harder than the initial learning—likely due to consistent attentional misguidance toward the initial (learnt) target position. Here, we studied how the different processes involved in contextual cueing and relearning affect the variability of neural responses in human participants as measured with EEG. Attention has previously been shown to reduce trial-by-trial variability in EEG [Arazi, A., Yeshurun, Y., & Dinstein, I. Neural variability is quenched by attention. Journal of Neuroscience, 39, 5975–5985, 2019], indicating that, in addition to increasing the neural response to an attended stimulus, attention may reduce the noise within the neural response itself. While repeated versus novel contexts did not modulate the trial-by-trial variability during initial learning, significant lateralized variability reductions were observed for repeated but not novel context trials in the relocations phase. This contrasts with how contextual cueing affected lateralized ERPs in past research. Zinchenko and colleagues [Zinchenko, A., Conci, M., Töllner, T., Müller, H. J., & Geyer, T. Automatic guidance (and misguidance) of visuospatial attention by acquired scene memory: Evidence from an N1pc polarity reversal. Psychological Science, 31, 1531–1543, 2020] found that lateralized ERPs signal correct and incorrect (i.e., misguided) attentional selection of target positions learned earlier. This phenomenon was observed during both the learning and relocation phases. Thus, variability and lateralized ERPs may represent different facets of attention, where variability becomes evident specifically under high attentional demand conditions, such as when participants must override the misguidance caused by LTM templates. |
Yi-Hsuan Chang; Chia-Shiang Lin; Cesar Barquero; Chin-An Wang Emotional conflict affects microsaccade dynamics in the emotional face–word Stroop task Journal Article In: Annals of the New York Academy of Sciences, vol. 1547, no. 1, pp. 204–219, 2025. @article{Chang2025f,Achieving optimal performance requires effectively resolving emotional conflict arising from the interference of task-irrelevant, emotionally salient stimuli. While microsaccade behavior has been linked to various cognitive and emotional processes, whether emotional conflict affects microsaccade responses remains to be determined. Additionally, pupil dilation is known to be modulated by emotional conflict signals, and both microsaccades and pupil dilation are arguably mediated by the superior colliculus (SC). However, the relationship between microsaccades and pupil dilation remains poorly understood. In this study, we investigated the effects of emotional conflict on microsaccade rates and metrics by presenting an emotional face–word stimulus in the face–word Stroop task. Larger microsaccade amplitudes (or higher peak velocities) were observed in the incongruent condition compared to the congruent condition, while microsaccade rates were similar between the two conditions. Additionally, microsaccade amplitudes were larger in incongruent trials following congruent trials than in those following incongruent trials. Furthermore, interindividual correlations between differences in microsaccade responses and State-Trait Anxiety Inventory scores were observed. Finally, trials with lower microsaccade rates were associated with larger pupil dilation. These results demonstrate the modulation of microsaccade metrics by emotional conflict, implicating the SC in integrating signals from the locus coeruleus network to coordinate these responses. |
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. |
Michael C. B. David; Emma Jane Mallas; Lucia M. Li; Magdalena A. Kolanko; Ramin Nilforooshan; Man Lai Tsoi; Hanim Karakoc; Karen Hoang; Johanna Brandt; Charikleia Triantafyllou; Dragos C. Gruia; Darije Custovic; Peter J. Lally; Karl A. Zimmerman; David Sharp; Care Research Institute; Technology Group UK Dementia Research; Paresh A. Malhotra; Gregory Scott1; David J. Sharp Pupil-linked arousal, cortical activity, and cognition in Alzheimer's disease Journal Article In: Brain Communications, vol. 7, no. 4, pp. 1–17, 2025. @article{David2025,Arousal dysfunction contributes to impairments seen in Alzheimer's disease. However, the nature and degree of this dysfunction have not been studied in detail. We investigated changes in tonic and phasic arousal using simultaneous pupillometry-EEG, relating these changes to locus coeruleus integrity, a key arousal nucleus. Forty Alzheimer's disease participants and 30 controls underwent neuropsychological testing using the Alzheimer's Disease Assessment Scale-Cognitive Subscale (ADAS-Cog), MRI designed to show contrast in the locus coeruleus as a measure of integrity and simultaneous pupillometry-EEG during 5 min of eyes-open resting-state. Pupillometry-EEG was then also applied during an oddball task which included a passive session and sessions in which responses to target stimuli were required, to test the effect of salience. Alzheimer's disease had lower locus coeruleus integrity (b = -0.26 |
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 Fischer; Danilo Postin; Lina Johanna Meiners; Louisa Kulke; Pascal Vrtička; Anne Schacht Challenging the negativity bias in affective scene viewing: The role of social content Journal Article In: Social Cognitive and Affective Neuroscience, vol. 20, no. 1, pp. 1–11, 2025. @article{Fischer2025,How does the brain prioritize information when visual scenes contain multiple sources of relevance? While emotionally evocative content has long been considered central to attentional capture, social content constitutes another key dimension of relevance. However, the neural mechanisms underlying the integration of these relevance dimensions remain unclear. We co-registered event-related potentials (ERPs) and eye movements while participants viewed complex scenes varying in social content (social, non-social) and emotional valence (positive, negative, neutral). Early ERP responses (P1) showed enhanced amplitudes for positive social images, suggesting that social relevance can mitigate the early-stage negativity bias. Social content also modulated the EPN, while emotional valence shaped later components (P300, LPC), with larger amplitudes for negative scenes. Eye-tracking measures mirrored these ERP effects: initial saccades were faster for social images, and fixation patterns indicated increased visual exploration for both positive and negative social scenes relative to neutral ones. Together, these results support a sequential appraisal process in which social content is prioritized at early perceptual stages, while emotional valence influences later evaluative processing. This pattern challenges the notion of a general negativity bias and underscores the interactive and stage-specific contributions of social and emotional relevance in affective scene perception. |
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. |
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, and Psychophysics, vol. 87, no. 1, pp. 5–24, 2025. @article{Huang2025f,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. |
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 |
Dennis Y. Jung; Bikash C. Sahoo; Adam C. Snyder Distractor anticipation during working memory is associated with theta and beta oscillations across spatial scales Journal Article In: Frontiers in Integrative Neuroscience, vol. 19, pp. 1–4, 2025. @article{Jung2025,Introduction: Anticipating distractors during working memory maintenance is critical to reduce their disruptive effects. In this study, we aimed to identify the oscillatory correlates of this process across different spatial scales of neural activity. Methods: We simultaneously recorded local field potentials (LFP) from the lateral prefrontal cortex (LPFC) and electroencephalograms (EEG) from the scalp of monkeys performing a modified memory-guided saccade (MGS) task. The monkeys were required to remember the location of a target visual stimulus while anticipating distracting visual stimulus, flashed at 50% probability during the delay period. Results: We found significant theta-band activity across spatial scales during anticipation of a distractor, closely linked with underlying working memory dynamics, through decoding and cross-temporal generalization analyses. EEG particularly reflected reactivation of memory around the anticipated time of a distractor, even in the absence of stimuli. During this anticipated time, beta-band activity exhibited transiently enhanced intrahemispheric communication between the LPFC and occipitoparietal brain areas. These oscillatory phenomena were observed only when the monkeys successfully performed the task, implicating their possible functional role in mitigating anticipated distractors. Discussion: Our results demonstrate that distractor anticipation recruits multiple oscillatory processes across the brain during working memory maintenance, with a key activity observed predominantly in the theta and beta bands. |
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. |
Lua Koenig; Biyu J. He Spontaneous slow cortical potentials and brain oscillations independently influence conscious visual perception Journal Article In: PLoS Biology, vol. 23, no. 1, pp. 1–33, 2025. @article{Koenig2025,Perceptual awareness results from an intricate interaction between external sensory input and the brain's spontaneous activity. Pre-stimulus ongoing activity influencing conscious perception includes both brain oscillations in the alpha (7 to 14 Hz) and beta (14 to 30 Hz) frequency ranges and aperiodic activity in the slow cortical potential (SCP, <5 Hz) range. However, whether brain oscillations and SCPs independently influence conscious perception or do so through shared mechanisms remains unknown. Here, we addressed this question in 2 independent magnetoencephalography (MEG) data sets involving near-threshold visual perception tasks in humans using low-level (Gabor patches) and high-level (objects, faces, houses, animals) stimuli, respectively. We found that oscillatory power and large-scale SCP activity influence conscious perception through independent mechanisms that do not have shared variance. In addition, through mediation analysis, we show that pre-stimulus oscillatory power and SCP activity have different relations to pupil size—an index of arousal—in their influences on conscious perception. Together, these findings suggest that oscillatory power and SCPs independently contribute to perceptual awareness, with distinct relations to pupil-linked arousal. |
Changhyun Lim; Hyun Jun Jeon; Woojae Jung; Youngshin Kwak; Myong Young Lee; Jung Hyun Ham; Hyun Bin Joo; Oh Sang Kwon; Dongil Chung Quantifying objective and perceived image quality through eeg and eye-tracking Journal Article In: IEEE Access, vol. 13, pp. 61250–61260, 2025. @article{Lim2025,Image quality evaluation of electronic displays is inherently subjective. To date, user studies in focus groups and viewers' self-reports have been the primary sources of feedback used as judgment criteria for the display quality. However, little has been known whether the measured responses to visual stimuli, other than self-reports, reflect the objective and subjective levels of image quality on displays. Here, we used electroencephalogram (EEG) and eye-tracking to investigate whether individuals' neural and physiological responses to visual stimuli track three objectively discerned levels of image quality, as well as their self-ratings on vividness. Our findings reveal that event-related potentials (ERPs) at 200-300 msec after stimulus onset in the frontocentral brain region, as well as saccade and blink frequencies, significantly tracked (cluster-based permutation p <0.05) the objective perspective of image quality. In contrast, ERPs at 600 msec in the frontolateral region and saccade peak velocity tracked (cluster-based permutation p <0.05) the subjective evaluation of image quality. Individuals' pupil diameter successfully tracked (cluster-based permutation p <0.05) both objective and subjective perspectives of image quality. These patterns highlight both the shared characteristics and measurable distinctions between objective image quality and subjective vividness ratings. This study demonstrates the effectiveness of EEG and eye-tracking as quantitative tools for assessing objective image quality disparities and the subjective affective nuances of users' visual display experiences, potentially reducing reliance on self-reports. |
Baiwei Liu; Freek Ede Concurrent selection of internal goals and external sensations during visual search Journal Article In: Science Advances, vol. 11, no. 45, pp. 1–10, 2025. @article{Liu2025d,Flexible goal-directed behavior relies on selecting relevant internal goal representations and external sensations. Yet, these selection processes have classically been studied in isolation, leaving unclear how they are coordinated in time to support behavior. To address this, we developed a visual search task to simultaneously track selection among internal search goals held in working memory and external search targets in the environment. Capitalizing on sensitive gaze and neural markers, we provide proof-of-principle evidence in humans that internal and external selection processes do not necessarily take turns in a strictly serial manner but can develop concurrently. These concurrent processes are supported by largely nonoverlapping neural activity patterns in the human brain and can be performed effectively even when engaging opposite spatial locations in working memory and perception. Our findings challenge views portraying brain states as being either internally or externally focused and bring insight into how internal and external selection processes work together to yield efficient search behavior. |
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. |
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, vol. 87, no. 1, pp. 120–132, 2025. @article{Mei2025,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, and Psychophysics, vol. 87, no. 1, pp. 94–119, 2025. @article{Melcher2025,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. |
Sara Milligan; Ayah Elaboudi; Brian Nestor; Elizabeth R. Schotter In: Cognitive, Affective & Behavioral Neuroscience, vol. 25, no. 4, pp. 1063–1082, 2025. @article{Milligan2025,The N400 ERP component has been characterized as a response reflecting binding of semantic memory states to create a “multimodal conceptual representation” (Kutas & Federmeier, 2011). An assumption of this characterization is that the measured response itself reflects the synchronization of various neural processes required for language comprehension. Less is known, however, about how these processes vary across individuals and how specific language skills may modulate particular underlying subcomponents of the N400. In the current study, we measured the N400 response to words that were (1) unexpected (i.e., low cloze in low constraint) or (2) orthographically related (OR) anomalies compared with expected words (i.e., high cloze in high constraint), and we investigated how these effects were related to participants' reading comprehension ability, vocabulary, and spelling ability. We found that contextual support N400 effects were larger for individuals with superior reading comprehension skills, whereas OR anomaly N400 effects were larger for individuals with superior spelling ability. These findings support the characterization of the N400 as a composite of various comprehension-related neural processes. The current study demonstrates that individual differentiation in the strength of these skills is differentially associated with the N400 subcomponents related to contextual facilitation and orthographic processing. Reading comprehension ability is associated with stronger contextual support effects, which may reflect more effective use of contextual support in facilitating semantic retrieval. Spelling skill is more strongly associated with OR anomaly effects (in contexts supporting an orthographic neighbor of the presented anomalous word), which may reflect more precise word identification. |
Sara Milligan; Milca Jaime Brunet; Neslihan Caliskan; Elizabeth R. Schotter Parafoveal N400 effects reveal that word skipping is associated with deeper lexical processing in the presence of context-driven expectations Journal Article In: Attention, Perception, & Psychophysics, vol. 87, no. 1, pp. 76–93, 2025. @article{Milligan2025a,Readers are able to begin processing upcoming words before directly fixating them, and in some cases skip words altogether (i.e., never fixated). However, the exact mechanisms and recognition thresholds underlying skipping decisions are not entirely clear. In the current study, we test whether skipping decisions reflect instances of more extensive lexical processing by recording neural language processing (via electroencephalography; EEG) and eye movements simultaneously, and we split trials based on target word-skipping behavior. To test lexical processing of the words, we manipulated the orthographic and phonological relationship between upcoming preview words and a semantically correct (and in some cases, expected) target word using the gaze-contingent display change paradigm. We also manipulated the constraint of the sentences to investigate the extent to which the identification of sublexical features of words depends on a reader's expectations. We extracted fixation-related brain potentials (FRPs) during the fixation on the preceding word (i.e., in response to parafoveal viewing of the manipulated previews). We found that word skipping is associated with larger neural responses (i.e., N400 amplitudes) to semantically incongruous words that did not share a phonological representation with the correct word, and this effect was only observed in high-constraint sentences. These findings suggest that word skipping can be reflective of more extensive linguistic processing, but in the absence of expectations, word skipping may occur based on less fine-grained linguistic processing and be more reflective of identification of plausible or expected sublexical features rather than higher-level lexical processing (e.g., semantic access). |
Kieran S. Mohr; Anna C. Geuzebroek; Simon P. Kelly Pattern-pulses and pattern-reversals evoke different cascades of cortical sources in the multifocal visual evoked potential Journal Article In: Journal of Vision, vol. 25, no. 7, pp. 1–19, 2025. @article{Mohr2025,The multifocal visual evoked potential (mVEP), elicited by either pattern-pulses or pattern-reversals, provides an effective means to study visual processing and to identify retinal damage and visual field defects. It is often assumed that the first components of these VEPs, the C1 and N75, respectively, are generated in V1, based on source modeling and their polarity reversal between upper- and lower-field stimulus presentations. However, limitations in the spatial resolution of source modeling and the non-uniqueness of the polarity reversal heuristic leave this assumed V1 source uncertain. We recently demonstrated the utility of a novel method to resolve visual sources by correlating retinotopically varying VEP topographies with predictions from the Benson-2014 retinotopy atlas. Here, we apply this method to study the sources of both the pulse and reversal mVEP, presented at the same stimulus event rates of between 3–8 Hz per location (approximately 35 Hz overall event rate). This analysis suggested that although V1 dominated the generation of the pulse mVEP throughout its time course, the initial component of the reversal mVEP was instead dominated by extrastriate areas, with V1 dominance emerging later from approximately 110 ms onwards. Although the initial component of the reversal mVEP did exhibit the classic sign of a V1 source—polarity reversal across the horizontal meridian—this basic feature is also produced by extrastriate areas such as V2 and V3, and the strong lateralization of topographies near the vertical meridian predicted by a V1 source was not observed. These results suggest that the pulse and reversal mVEP evoke different cascades of generative visual areas when evoked at the event rate tested here. |
Ascensión Pagán; Federica Degno; Sara V. Milledge; Richard D. Kirkden; Sarah J. White; Simon P. Liversedge; Kevin B. Paterson Aging and word predictability during reading: Evidence from eye movements and fixation-related potentials Journal Article In: Attention, Perception, & Psychophysics, vol. 87, no. 1, pp. 50–75, 2025. @article{Pagan2025,The use of context to facilitate the processing of words is recognized as a hallmark of skilled reading. This capability is also hypothesized to change with older age because of cognitive changes across the lifespan. However, research investigating this issue using eye movements or event-related potentials (ERPs) has produced conflicting findings. Specifically, whereas eye-movement studies report larger context effects for older than younger adults, ERP findings suggest that context effects are diminished or delayed for older readers. Crucially, these contrary findings may reflect methodological differences, including use of unnatural sentence displays in ERP research. To address these limitations, we used a coregistration technique to record eye movements (EMs) and fixation-related potentials (FRPs) simultaneously while 44 young adults (18–30 years) and 30 older adults (65+ years) read sentences containing a target word that was strongly or weakly predicted by prior context. Eye-movement analyses were conducted over all data (full EM dataset) and only data matching FRPs. FRPs were analysed to capture early and later components 70–900 ms following fixation-onset on target words. Both eye-movement datasets and early FRPs showed main effects of age group and context, while the full EM dataset and later FRPs revealed larger context effects for older adults. We argue that, by using coregistration methods to address limitations of earlier ERP research, our experiment provides compelling complementary evidence from eye movements and FRPs that older adults rely more on context to integrate words during reading. |
Laura Pasqualette; Louisa Kulke Emotional expressions, but not social context, modulate attention during a discrimination task Journal Article In: Cognition and Emotion, vol. 39, no. 5, pp. 1108–1126, 2025. @article{Pasqualette2025,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. |
Lisa-Marie G. Pohle; Moritz M. Nickel; Birgit Nierula; Markus Ploner; Ulrike Horn; Falk Eippert Expectation effects on repetition suppression in nociception Journal Article In: Journal of Neurophysiology, vol. 134, no. 4, pp. 1244–1256, 2025. @article{Pohle2025,Repetition suppression, the reduced neural response upon repeated presentation of a stimulus, can be explained by models focusing on bottom-up (i.e., adaptation) or top-down (i.e., expectation) mechanisms. Predictive coding models fall into the latter category and propose that repetitions are expected and therefore elicit smaller prediction error responses. Although studies in the visual and auditory domain provide some support for such models, in nociception evidence remains inconclusive, despite the substantial influence expectations exert on pain perception. To assess expectation effects on repetition suppression in nociception, we developed a paradigm in which healthy volunteers received brief CO2 laser stimuli, while we acquired electroencephalographic (EEG) and peripheral physiological data. Importantly, laser stimuli could be either repeated after one second or not be repeated, with the probability of repetitions manipulated in a block-wise fashion, such that repetitions were either expected or unexpected. We observed repetition suppression in laser-evoked potentials and laser-induced gamma band oscillations, but not in laser-induced desynchronizations in the alpha and beta band. Critically, neither these EEG responses, nor the peripheral physiological data showed significant differences between the expectation conditions, with Bayesian analyses mostly providing evidence for an absence of effects. This indicates that repetition suppression to brief nociceptive laser stimuli is not driven by top-down factors, but rather mediated by other adaptation processes. Although this does not preclude an influence of predictive coding models in nociception, it suggests that when the nervous system receives highly precise input, its responses are less susceptible to influence from expectations.NEW & NOTEWORTHY We show that repetition suppression (RS; the diminished neural activity upon repeated stimulus presentation) can be observed in the nociceptive domain: brief laser stimuli lead to RS in event-related potentials and gamma oscillations. Importantly, nociceptive RS is not modulated by repetition probability and thus expectations regarding the occurrence of a repetition. This indicates that nociceptive responses are less prone to top-down expectations under conditions of limited sensory uncertainty, as established via precise laser stimuli. |
Kristina I. Pultsina; Tatiana A. Stroganova; Galina L. Kozunova; Andrey O. Prokofyev; Aleksandra S. Miasnikova; Anna M. Rytikova; Boris V. Chernyshev Atypical pupil-linked arousal induced by low-risk probabilistic choices, and intolerance of uncertainty in adults with ASD Journal Article In: Cognitive, Affective & Behavioral Neuroscience, vol. 25, no. 2, pp. 531–549, 2025. @article{Pultsina2025a,Adults with autism spectrum disorder (ASD) experience stress when operating in a probabilistic environment, even if it is familiar, but the underlying mechanisms remain unclear. Their decision-making may be affected by the uncertainty aversion implicated in ASD and associated with increased autonomic arousal. Previous studies have shown that in neurotypical (NT) people, decisions with predictably better outcomes are less stressful and elicit smaller pupil-linked arousal than those involving exploration. Here, in a sample of 46 high-functioning ASD and NT participants, using mixed-effects model analysis, we explored pupil-linked arousal and behavioral performance in a probabilistic reward learning task with a stable advantage of one choice option over the other. We found that subjects with ASD learned and preferred advantageous probabilistic choices at the same rate and to the same extent as NT participants, both in terms of choice ratio and response time. Although both groups exhibited similar predictive behaviors, learning to favor advantageous choices led to increased pupillary arousal for these choices in the ASD group, while it caused a decrease in pupillary arousal in the NT group. Moreover, greater pupil-linked arousal during decisions with higher expected value correlated with greater degree of self-reported intolerance of uncertainty in everyday life. Our results suggest that in a nonvolatile probabilistic environment, objectively good predictive abilities in people with ASD are coupled with elevated physiological stress and subjective uncertainty regarding the decisions with the best possible but still uncertain outcome that contributes to their intolerance of uncertainty. |
Hannah N. Rembrandt; Ellyn A. Riley Evidence of physiological changes associated with single-session pre-frontal tDCS: A pilot study Journal Article In: Frontiers in Human Neuroscience, vol. 19, pp. 1–17, 2025. @article{Rembrandt2025,Objective: Transcranial direct current stimulation (tDCS), a non-invasive, painless method of applying direct current electrical stimulation to specific areas of the brain, is an effective method for enhancing attention and post-stroke fatigue, as shown by behavioral improvements in post-stroke populations. While behavioral evidence supports this method, there is a paucity of physiological data corroboration of this improvement. The current study is designed to investigate if a single session of tDCS will improve attention and fatigue as shown by relevant physiological methods in persons with post-stroke aphasia. Methods: Ten participants (5 male; mean age: 62.8) engaged in two identically structured data collection sessions with at least a 3-day wash-out period between them. Sessions started with a sustained attention task with simultaneous electroencephalography (EEG) and pupillometry data collection, followed by an attention training program with simultaneous active or sham tDCS. Following tDCS, participants repeated the sustained attention task with simultaneous EEG and pupillometry data collection. Participants received active tDCS during one session, and sham tDCS during the other, with the order randomized. Results: No differences between conditions were found for either behavioral results from the sustained attention task (i.e., reaction time of correct responses; n = 9 p = 0.39) or EEG measured attention state data for any of the four attention states: no attention (n = 10 |
Veli-Matti Saarinen; Veikko Jousmäki Eye tracking in MEG Journal Article In: Attention, Perception, & Psychophysics, vol. 87, no. 1, pp. 238–244, 2025. @article{Saarinen2025,Magnetoencephalography (MEG) can measure brain activity in ms-level temporal resolution. MEG sensors are super sensitive devices for magnetic signals of the brain but are also prone to electromagnetic interferences. The MEG device is located inside the magnetically shielded room (MSR), and any monitoring device used inside the MSR requires special shielding and its location must be carefully selected to suppress electromagnetic interference. Eye-tracker measures eye movements, providing spatial location of the gaze, pupil diameters, and eye blinks. Eye tracking in MEG enables, for example, categorization of the MEG data based on gaze position and interactive stimulus using gaze position. Combining the methods together will require considering the electromagnetic interference for the MEG—that is, additional shielding, positioning of the eye tracker, and subject-specific issues related to make-up and eye-corrective lenses. |
Travis N. Talcott; John E. Kiat; Steven J. Luck; Nicholas Gaspelin Is covert attention necessary for programming accurate saccades? Evidence from saccade-locked event-related potentials Journal Article In: Attention, Perception, and Psychophysics, vol. 87, no. 1, pp. 172–190, 2025. @article{Talcott2025,For decades, researchers have assumed that shifts of covert attention mandatorily occur prior to eye movements to improve perceptual processing of objects before they are fixated. However, recent research suggests that the N2pc component—a neural measure of covert attentional allocation—does not always precede eye movements. The current study investigated whether the N2pc component mandatorily precedes eye movements and assessed its role in the accuracy of gaze control. In three experiments, participants searched for a letter of a specific color (e.g., red) and directed gaze to it as a response. Electroencephalograms and eye movements were coregistered to determine whether neural markers of covert attention preceded the initial shift of gaze. The results showed that the presaccadic N2pc only occurred under limited conditions: when there were many potential target locations and distractors. Crucially, there was no evidence that the presence or magnitude of the presaccadic N2pc was associated with improved eye movement accuracy in any of the experiments. Interestingly, ERP decoding analyses were able to classify the target location well before the eyes started to move, which likely reflects a presaccadic cognitive process that is distinct from the attentional process measured by the N2pc. Ultimately, we conclude that the covert attentional mechanism indexed by the N2pc is not necessary for precise gaze control. |
Natalie G. Wall; Oliver Smith; Linda Campbell; Carmel Loughland; Ulrich Schall Using EEG and eye tracking to evaluate an emotion recognition ipad app for autistic children Journal Article In: Clinical EEG and Neuroscience, pp. 1–11, 2025. @article{Wall2025c,Autism is a neurodevelopmental condition that impacts individuals' communication and social interaction skills. Autistic children often have smaller N170 amplitudes in response to faces than neurotypical children. Autistic children also avoid the salient areas of the face. Technology-based interventions have been developed to teach autistic children how to recognise facial expressions, but the results have exhibited considerable variability across studies. The current study explored the effectiveness of an iPad app designed to support autistic children in recognising facial expressions by examining how participants process facial information through event-related potentials (ERP) and eye-tracking recordings. ERPs and eye tracking were recorded from 20 neurotypical and 15 autistic children aged between 6 and 12 years. The results replicated previous work, with the autistic group having smaller N170 and Vertex Positive Potential amplitudes and more scan time off the face when compared to non-autistic children. Following the intervention, some changes were observed in facial feature scanning among autistic participants, characterised by increased time spent on the face and decreased fixations. These findings add to the work, indicating that eye tracking may be a valuable biomarker for intervention outcomes in autism. Further research into N170 as a biomarker is needed. |
Changying Yan Enhancing English language teaching quality evaluation via dynamic multimodal cognitive transfer models Journal Article In: International Journal of Information and Communication Technology, vol. 26, no. 32, pp. 121–141, 2025. @article{Yan2025c,This paper proposes a dynamic assessment method based on multimodal cognitive transfer modelling to address the limitations of static and unidimensional analysis in English teaching quality assessment. A two-channel LSTM-cognitive state space model is constructed by synchronously collecting four-dimensional data of speech, vision, text and physiological signals in the teaching scene, quantifying students' cognitive state transfer trajectories based on the ACT-R cognitive architecture in the knowledge transfer channel, and adopting a dynamic causal map to model the feedback mechanism of teachers' strategy adjustment in the teaching intervention channel. A time-varying weighted assessment function was designed to dynamically fuse cognitive state vectors with intervention intensity. In a 136-lesson experiment in 12 schools, the causal attribution rate of this method was improved by 41.2%, and the adoption rate of intervention suggestions reached 83.7%, which verified the effectiveness and universality for dynamic quality assessment of English teaching. |
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. |
Siqi Yuan; Chenyu Pang; Liang Wu; Li Yi; Kun Guo; Yong Hui Jiang; Yong Q. Zhang; Shihui Han Autism-like atypical face processing in Shank3 mutant dogs Journal Article In: Science Advances, vol. 11, no. 14, pp. 1–12, 2025. @article{Yuan2025b,Atypical face processing is a neurocognitive basis of social deficits in autism spectrum disorder (ASD) and a candidate cognitive marker for the disease. Although hundreds of risk genes have been identified in ASD, it remains unclear whether mutations in a specific gene may cause ASD-like atypical face processing. Dogs have acquired exquisite face processing abilities during domestication and may serve as an effective animal model for studying genetic associations of ASD-like atypical face processing. Here, we showed that dogs with Shank3 mutations exhibited behavioral and attentional avoidance of faces, contrasting with wild-type controls. Moreover, neural responses specific to faces (versus objects) recorded from the electrodes over the temporal cortex were significantly decreased and delayed in Shank3 mutants compared to wild-type controls. Cortical responses in the frontal/ parietal region underlying categorization of faces by species/breeds were reduced in Shank3 mutants. Our findings of atypical face processing in dogs with Shank3 mutations provide a useful animal model for studying ASD mechanisms and treatments. |
Rajamanickam Yuvaraj; Sampathraman Samyuktha; Jack Fogarty; Jun Song Huang; Samuel Tan; Wong Teck Kiong Automated boredom recognition using multimodal physiological signals Journal Article In: IEEE Transactions on Affective Computing, pp. 1–18, 2025. @article{Yuvaraj2025,Increasing access to affordable physiological sensors and computing devices has bolstered the development of emotionaware systems that recognize human emotions. However, to date, there has been little research testing machine-driven approaches to recognizing boredom, particularly in learning contexts. To address that, this study tests the automatic recognition of boredom induced during a video lecture using a multimodal recognition system relying on physiological signals. Electroencephalogram (EEG), electrocardiogram (ECG), galvanic skin response (GSR), and eye gaze data were recorded from 84 healthy adults (mean age = 26.90 ± 5.29) as they watched both non-boring and boring educational videos to control and induce boredom. Signal features were extracted from the physiological data and validated using Wilcoxon signed-rank tests prior to evaluating their utility for boredom recognition with three separate machine-learning classification techniques, namely Extreme Gradient Boosting (XGB), Random Forest (RF), and Gradient Boosting (GB), with leave-one-out cross-validation. Both unimodal and multimodal recognition systems were assessed by evaluating model performance using each physiological signal separately or in combination (using decision fusion). Multimodal recognition was found to enhance model performance when compared to any single modality, with the highest average boredom recognition accuracy of 88.56% ± 0.82% recorded for EEG + eye gaze modal fusion with RF. |
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. |
Xinni Zhang; Zhineng Lv; Xuesong Jin; Lijun Yun; Guihong Zhao; Zaiqing Chen Multimodal neurobehavioral integration in binocular color rivalry: Cortical-eye movement analysis under color, location, and combined stimuli Journal Article In: Frontiers in Neuroscience, vol. 19, pp. 1–22, 2025. @article{Zhang2025q,Objective: Color and spatial location are key cues influencing visual selection in binocular color rivalry, jointly modulating attentional resource allocation and prefrontal cortex (PFC) activity. However, how these cues—individually and in combination—regulate ocular behavior and PFC activation remains insufficiently understood and lacks systematic empirical investigation. Methods: This study integrates eye tracking, functional near-infrared spectroscopy (fNIRS), and reaction time measurements to systematically investigate the cortical and oculomotor response characteristics under a binocular rivalry paradigm, focusing on color rivalry (Color stimuli), spatial location (Location stimuli), and their combined (Color & Location stimuli). Results: The results revealed that Color stimuli elicited rapid saccades, significant pupil dilation, and a decrease in prefrontal HbO concentration. Location stimuli induced stable saccadic patterns and a typical biphasic HbR response in BA46. The combined Color & Location stimuli triggered significant changes in oculomotor behavior during the later processing stage, accompanied by a marked increase in HbO activation in BA10, suggesting its dominant role in multisensory integration and cognitive resource reallocation. Further coupling analyses showed a significant positive correlation between prefrontal HbO concentration and reaction time (r = 0.555, p < 0.01), and a significant negative correlation between HbO concentration and saccade amplitude (r = –0.376, p < 0.05), consistent with the theoretical predictions of the “neural efficiency–cognitive load trade-off” model. Task-dependent coupling relationship were also observed among oculomotor parameters and between eye movement and cerebral hemodynamic signals. Conclusion: Color stimuli induce rapid saccadic behavior and impose higher prefrontal load, Location stimuli engage a more efficient dorsal pathway, while Color & Location stimuli intensify resource rivalry and induce a processing bottleneck, manifested as prolonged reaction times co-occurring with heightened cortical activation. |
Floortje G. Bouwkamp; Floris P. Lange; Eelke Spaak Spatial predictive context speeds up visual search by biasing local attentional competition Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 1, pp. 28–42, 2025. @article{Bouwkamp2025,The human visual system is equipped to rapidly and implicitly learn and exploit the statistical regularities in our environment. Within visual search, contextual cueing demonstrates how implicit knowledge of scenes can improve search performance. This is commonly interpreted as spatial context in the scenes becoming predictive of the target location, which leads to a more efficient guidance of attention during search. However, what drives this enhanced guidance is unknown. First, it is under debate whether the entire scene (global context) or more local context drives this phenomenon. Second, it is unclear how exactly improved attentional guidance is enabled by target enhancement and distractor suppression. In the present magnetoencephalography experiment, we leveraged rapid invisible frequency tagging to answer these two outstanding questions. We found that the improved performance when searching implicitly familiar scenes was accompanied by a stronger neural representation of the target stimulus, at the cost specifically of those distractors directly surrounding the target. Crucially, this biasing of local attentional competition was behaviorally relevant when searching familiar scenes. Taken together, we conclude that implicitly learned spatial predictive context improves how we search our environment by sharpening the attentional field. |
2024 |
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. |
Tara L. Alvarez; Mitchell Scheiman; Suril Gohel; Farzin Hajebrahimi; Melissa Noble; Ayushi Sangoi; Chang Yaramothu; Christina L. Master; Arlene Goodman Effectiveness of treatment for concussion-related convergence insufficiency: The CONCUSS study protocol for a randomized clinical trial Journal Article In: PloS ONE, vol. 19, no. 11, pp. 1–23, 2024. @article{Alvarez2024,PURPOSE: To describe CONCUSS, a randomized clinical trial (RCT) designed to compare the following: the effectiveness of immediate office-based vergence/accommodative therapy with movement (OBVAM) to delayed OBVAM as treatments for concussion-related convergence insufficiency (CONC-CI) to understand the impact of time (watchful waiting), the effect of OBVAM dosage (12 versus 16 therapy sessions), and to investigate the underlying neuro-mechanisms of OBVAM on CONC-CI participants. METHODS: CONCUSS is an RCT indexed on https://clinicaltrials.gov/study/NCT05262361 enrolling 100 participants aged 11-25 years with medically diagnosed concussion, persistent post-concussive symptoms 4-24 weeks post-injury, and symptomatic convergence insufficiency. Participants will receive standard concussion care and will be randomized to either immediate OBVAM or delayed (by six weeks) OBVAM. At the Outcome 1 examination (week 7), clinical assessments of success as determined by changes in the near point of convergence (NPC), positive fusional vergence (PFV), and symptoms will be compared between the two treatment groups. After the Outcome 1 visit, those in the delayed group receive 16 visits of OBVAM, while those in the immediate OBVAM group receive four more therapy visits. Outcome 2 assessment will be used to compare both groups after participants receive 16 sessions of OBVAM. The primary measure is the between-group differences of the composite change in the NPC and PFV at the Outcome 1 visit. Secondary outcome measures include individual clinical measures, objective eye-tracking parameters, and functional brain imaging. CONCLUSIONS: Major features of the study design include formal definitions of conditions and outcomes, standardized diagnostic and treatment protocols, a delayed treatment arm, masked outcome examinations, and the incorporation of objective eye movement recording and brain imaging as outcome measures. CONCUSS will establish best practices in the clinical care of CONC-CI. The objective eye movement and brain imaging, correlated with the clinical signs and symptoms, will determine the neuro-mechanisms of OBVAM on CONC-CI. |
Elaine J. Anderson; Tessa M. Dekker; Mahtab Farahbakhsh; Nashila Hirji; D. Samuel Schwarzkopf; Michel Michaelides; Geraint Rees fMRI and gene therapy in adults with CNGB3 mutation Journal Article In: Brain Research Bulletin, vol. 215, pp. 1–10, 2024. @article{Anderson2024,Achromatopsia is an inherited retinal disease that affects 1 in 30,000–50,000 individuals and is characterised by an absence of functioning cone photoreceptors from birth. This results in severely reduced visual acuity, no colour vision, marked sensitivity to light and involuntary oscillations of the eyes (nystagmus). In most cases, a single gene mutation prevents normal development of cone photoreceptors, with mutations in the CNGB3 or CNGA3 gene being responsible for ∼80 % of all patients with achromatopsia. There are a growing number of studies investigating recovery of cone function after targeted gene therapy. These studies have provided some promise for patients with the CNGA3 mutation, but thus far have found limited or no recovery for patients with the CNGB3 mutation. Here, we developed colour-calibrated visual stimuli designed to isolate cone photoreceptor responses. We combined these with adapted fMRI techniques and pRF mapping to identify if cortical responses to cone-driven signals could be detected in 9 adult patients with the CNGB3 mutation after receiving gene therapy. We did not detect any change in brain activity after gene therapy when the 9 patients were analysed as a group. However, on an individual basis, one patient self-reported a change in colour perception, corroborated by improved performance on a psychophysical task designed to selectively identify cone function. This suggests a level of cone sensitivity that was lacking pre-treatment, further supported by a subtle but reliable change in cortical activity within their primary visual cortex. |
Reza Azadi; Emily Lopez; Jessica Taubert; Amanda Patterson; Arash Afraz Inactivation of face-selective neurons alters eye movements when free viewing faces Journal Article In: Proceedings of the National Academy of Sciences, vol. 121, no. 3, pp. 1–10, 2024. @article{Azadi2024a,During free viewing, faces attract gaze and induce specific fixation patterns corresponding to the facial features. This suggests that neurons encoding the facial features are in the causal chain that steers the eyes. However, there is no physiological evidence to support a mechanistic link between face- encoding neurons in high- level visual areas and the oculo- motor system. In this study, we targeted the middle face patches of the inferior temporal (IT) cortex in two macaque monkeys using an functional magnetic resonance imaging (fMRI) localizer. We then utilized muscimol microinjection to unilaterally suppress IT neural activity inside and outside the face patches and recorded eye movements while the animals free viewing natural scenes. Inactivation of the face- selective neurons altered the pattern of eye movements on faces: The monkeys found faces in the scene but neglected the eye contralateral to the inactivation hemisphere. These findings reveal the causal contribution of the high- level visual cortex in eye movements. |
Mareike Bacha-Trams; Gökce Ertas Yorulmaz; Enrico Glerean; Elisa Ryyppö; Karoliina Tapani; Eero Virmavirta; Jenni Saaristo; Iiro P. Jääskeläinen; Mikko Sams Sisterhood predicts similar neural processing of a film Journal Article In: NeuroImage, vol. 297, pp. 1–14, 2024. @article{BachaTrams2024,Relationships between humans are essential for how we see the world. Using fMRI, we explored the neural basis of homophily, a sociological concept that describes the tendency to bond with similar others. Our comparison of brain activity between sisters, friends and acquaintances while they watched a movie, indicate that sisters' brain activity is more similar than that of friends and friends' activity is more similar than that of acquaintances. The increased similarity in brain activity measured as inter-subject correlation (ISC) was found both in higher-order brain areas including the default-mode network (DMN) and sensory areas. Increased ISC could not be explained by genetic relation between sisters neither by similarities in eye-movements, emotional experiences, and physiological activity. Our findings shed light on the neural basis of homophily by revealing that similarity in brain activity in the DMN and sensory areas is the stronger the closer is the relationship between the people. |
Elise Beckers; Islay Campbell; Roya Sharifpour; Ilenia Paparella; Alexandre Berger; Jose Fermin Balda Aizpurua; Ekaterina Koshmanova; Nasrin Mortazavi; Puneet Talwar; Siya Sherif; Heidi I. L. Jacobs; Gilles Vandewalle Impact of repeated short light exposures on sustained pupil responses in an fMRI environment Journal Article In: Journal of Sleep Research, vol. 33, no. 4, pp. 1–14, 2024. @article{Beckers2024,Light triggers numerous non-image-forming, or non-visual, biological effects. The brain correlates of these non-image-forming effects have been investigated, notably using magnetic resonance imaging and short light exposures varying in irradiance and spectral quality. However, it is not clear whether non-image-forming responses estimation may be biased by having light in sequential blocks, for example, through a potential carryover effect of one light onto the next. We reasoned that pupil light reflex was an easy readout of one of the non-image-forming effects of light that could be used to address this issue. We characterised the sustained pupil light reflex in 13–16 healthy young individuals under short light exposures during three distinct cognitive processes (executive, emotional and attentional). Light conditions pseudo-randomly alternated between monochromatic orange light (0.16 melanopic equivalent daylight illuminance lux) and polychromatic blue-enriched white light of three different levels (37, 92, 190 melanopic equivalent daylight illuminance lux). As expected, higher melanopic irradiance was associated with larger sustained pupil light reflex in each cognitive domain. This result was stable over the light sequence under higher melanopic irradiance levels compared with lower ones. Exploratory frequency-domain analyses further revealed that sustained pupil light reflex was more variable under lower melanopic irradiance levels. Importantly, sustained pupil light reflex varied across tasks independently of the light condition, pointing to a potential impact of light history and/or cognitive context on sustained pupil light reflex. Together, our results emphasise that the distinct contribution and adaptation of the different retinal photoreceptors influence the non-image-forming effects of light and therefore potentially their brain correlates. |
Johanna Bergmann; Lucy S. Petro; Clement Abbatecola; Min S. Li; A. Tyler Morgan; Lars Muckli Cortical depth profiles in primary visual cortex for illusory and imaginary experiences Journal Article In: Nature Communications, vol. 15, no. 1, pp. 1–13, 2024. @article{Bergmann2024,Visual illusions and mental imagery are non-physical sensory experiences that involve cortical feedback processing in the primary visual cortex. Using laminar functional magnetic resonance imaging (fMRI) in two studies, we investigate if information about these internal experiences is visible in the activation patterns of different layers of primary visual cortex (V1). We find that imagery content is decodable mainly from deep layers of V1, whereas seemingly ‘real' illusory content is decodable mainly from superficial layers. Furthermore, illusory content shares information with perceptual content, whilst imagery content does not generalise to illusory or perceptual information. Together, our results suggest that illusions and imagery, which differ immensely in their subjective experiences, also involve partially distinct early visual microcircuits. However, overlapping microcircuit recruitment might emerge based on the nuanced nature of subjective conscious experience. |
Jack J. Blanchard; Jason F. Smith; Melanie E. Bennett; Ryan D. Orth; Christina L. G. Savage; Julie M. McCarthy; James A. Coan; Alexander J. Shackman Motivation and pleasure deficits undermine the benefits of social affiliation in psychosis Journal Article In: Clinical Psychological Science, vol. 12, no. 6, pp. 1195 –1217, 2024. @article{Blanchard2024,In psychotic disorders, motivation and pleasure (MAP) deficits are associated with decreased affiliation and heightened functional impairment. We leveraged a transdiagnostic sample enriched for psychosis and a multimethod approach to test the hypothesis that MAP deficits undermine the stress-buffering benefits of affiliation. Participants completed the social-affiliation-enhancement task (SAET) to cultivate affiliation with an experimental partner. Although the SAET increased perceived affiliation and mood, individuals with greater negative symptoms derived smaller emotional benefits from the partners, as indexed by self-report and facial behavior. We then used the handholding functional MRI paradigm, which combines threat anticipation with affiliative physical contact, to determine whether MAP deficits undermine the social regulation of distress. Individuals with greater MAP deficits showed diminished neural “benefits”—reduced dampening of threat-elicited activation—from affiliative touch in key frontoparietal nodes of the dorsal attention network. In short, MAP symptoms disrupt the emotional and neuroregulatory benefits of affiliation. |
Magdalena Bocha; Sabrina Karl; Isabella C. Wagnera; Lukas L. Lengersdorff; Ludwig Huberc; Claus Lamm Action observation reveals a network with divergent temporal and parietal cortex engagement in dogs compared with humans Journal Article In: Imaging Neuroscience, vol. 2, pp. 1–29, 2024. @article{Bocha2024,Action observation is a fundamental pillar of social cognition. Neuroimaging research has revealed a human and non-human primate action observation network (AON) encompassing frontotemporoparietal areas with links to the species' imitation tendencies and relative lobe expansion. Dogs (Canis familiaris) have good action perception and imitation skills and a less expanded parietal than temporal cortex, but their AON remains unexplored. We conducted a functional MRI study with 28 dogs and 40 humans and found functionally analogous involvement of somatosensory and temporal brain areas of both species' AONs and responses to transitive and intransitive action observation in line with their imitative skills. Employing a functional localizer, we also identified functionally analogous agent-responsive areas within both species' AONs. However, activation and task-based functional connectivity measures suggested significantly less parietal cortex involvement in dogs than in humans. These findings advance our understanding of the neural bases of action understanding and the convergent evolution of social cognition, with analogies and differences resulting from similar social environments and divergent brain expansion, respectively. |
Petra Borovska; Benjamin Haas Individual gaze shapes diverging neural representations Journal Article In: Proceedings of the National Academy of Sciences, vol. 121, no. 36, pp. 1–3, 2024. @article{Borovska2024,Complex visual stimuli evoke diverse patterns of gaze, but previous research suggests that their neural representations are shared across brains. Here, we used hyperalignment to compare visual responses between observers viewing identical stimuli. We find that individual eye movements enhance cortical visual responses but also lead to representational divergence. Pairwise differences in the spatial distribution of gaze and in semantic salience predict pairwise representational divergence in V1 and inferior temporal cortex, respectively. This suggests that individual gaze sculpts individual visual worlds. |
Andy Brendler; Max Schneider; Immanuel G. Elbau; Rui Sun; Taechawidd Nantawisarakul; Dorothee Pöhlchen; Tanja Brückl; A. K. Brem; E. B. Binder; A. Erhardt; J. Fietz; N. C. Grandi; Y. Kim; S. Ilić-Ćoćić; L. Leuchs; S. Lucae; T. Namendorf; J. Pape; L. Schilbach; I. Mücke-Heim; J. Ziebula; Michael Czisch; Philipp G. Sämann; Michael D. Lee; Victor I. Spoormaker In: Scientific Reports, vol. 14, no. 344, pp. 1–11, 2024. @article{Brendler2024,Major depressive disorder (MDD) is a devastating and heterogenous disorder for which there are no approved biomarkers in clinical practice. We recently identified anticipatory hypo-arousal indexed by pupil responses as a candidate mechanism subserving depression symptomatology. Here, we conducted a replication and extension study of these findings. We analyzed a replication sample of 40 unmedicated patients with a diagnosis of depression and 30 healthy control participants, who performed a reward anticipation task while pupil responses were measured. Using a Bayesian modelling approach taking measurement uncertainty into account, we could show that the negative correlation between pupil dilation and symptom load during reward anticipation is replicable within MDD patients, albeit with a lower effect size. Furthermore, with the combined sample of 136 participants (81 unmedicated depressed and 55 healthy control participants), we further showed that reduced pupil dilation in anticipation of reward is inversely associated with anhedonia items of the Beck Depression Inventory in particular. Moreover, using simultaneous fMRI, particularly the right anterior insula as part of the salience network was negatively correlated with depressive symptom load in general and anhedonia items specifically. The present study supports the utility of pupillometry in assessing noradrenergically mediated hypo-arousal during reward anticipation in MDD, a physiological process that appears to subserve anhedonia. |
Holly Bridge; Abigail Wyllie; Aaron Kay; Bailey Rand; Lucy Starling; Rebecca S. Millington-Truby; William T. Clarke; Jasleen K. Jolly; I. Betina Ip Neurochemistry and functional connectivity in the brain of people with Charles Bonnet syndrome Journal Article In: Therapeutic Advances in Ophthalmology, vol. 16, pp. 1–18, 2024. @article{Bridge2024,Background: Charles Bonnet syndrome (CBS) is a condition in which people with vision loss experience complex visual hallucinations. These complex visual hallucinations may be caused by increased excitability in the visual cortex that are present in some people with vision loss but not others. Objectives: We aimed to evaluate the association between γ-aminobutyric acid (GABA) in the visual cortex and CBS. We also tested the relationship among visually evoked responses, functional connectivity, and CBS. Design: This is a prospective, case-controlled, cross-sectional observational study. Methods: We applied 3-Tesla magnetic resonance spectroscopy, as well as task-based and resting state (RS) connectivity functional magnetic resonance imaging in six participants with CBS and six controls without CBS. GABA+ was measured in the early visual cortex (EVC) and in the lateral occipital cortex (LOC). Participants also completed visual acuity and cognitive tests, and the North-East Visual Hallucinations Interview. Results: The two groups were well-matched for age, gender, visual acuity and cognitive scores. There was no difference in GABA+ levels between groups in the visual cortex. Most participants showed the expected blood oxygenation level dependent (BOLD) activation to images of objects and the phase-scrambled control. Using a fixed effects analysis, we found that BOLD activation was greater in participants with CBS compared to controls. Analysis of RS connectivity with LOC and EVC showed little difference between groups. A fixed effects analysis showed a correlation between the extent of functional connectivity with LOC and hallucination strength. Conclusion: Overall, our results provide no strong evidence for an association between GABAergic inhibition in the visual cortex and CBS. We only found subtle differences in visual function and connectivity between groups. These findings suggest that the neurochemistry and visual connectivity for people with Charles Bonnet hallucinations are comparable to a sight loss population. Differences between groups may emerge when investigating subtle and transient changes that occur at the time of visual hallucinations. |
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. |
Yvonne Buschermöhle; Malte B. Höltershinken; Tim Erdbrügger; Jan Ole Radecke; Andreas Sprenger; Till R. Schneider; Rebekka Lencer; Joachim Gross; Carsten H. Wolters Comparing the performance of beamformer algorithms in estimating orientations of neural sources Journal Article In: iScience, vol. 27, no. 3, pp. 1–21, 2024. @article{Buschermoehle2024,The efficacy of transcranial electric stimulation (tES) to effectively modulate neuronal activity depends critically on the spatial orientation of the targeted neuronal population. Therefore, precise estimation of target orientation is of utmost importance. Different beamforming algorithms provide orientation estimates; however, a systematic analysis of their performance is still lacking. For fixed brain locations, EEG and MEG data from sources with randomized orientations were simulated. The orientation was then estimated (1) with an EEG and (2) with a combined EEG-MEG approach. Three commonly used beamformer algorithms were evaluated with respect to their abilities to estimate the correct orientation: Unit-Gain (UG), Unit-Noise-Gain (UNG), and Array-Gain (AG) beamformer. Performance depends on the signal-to-noise ratios for the modalities and on the chosen beamformer. Overall, the UNG and AG beamformers appear as the most reliable. With increasing noise, the UG estimate converges to a vector determined by the leadfield, thus leading to insufficient orientation estimates. |
Islay Campbell; Roya Sharifpour; Jose Fermin Balda Aizpurua; Elise Beckers; Ilenia Paparella; Alexandre Berger; Ekaterina Koshmanova; Nasrin Mortazavi; John Read; Mikhail Zubkov; Puneet Talwar; Fabienne Collette; Siya Sherif; Christophe Phillips; Laurent Lamalle; Gilles Vandewalle Regional response to light illuminance across the human hypothalamus Journal Article In: eLife, vol. 13, pp. 1–21, 2024. @article{Campbell2024d,Light exerts multiple non-image-forming biological effects on physiology including the stimulation of alertness and cognition. However, the subcortical circuitry underlying the stimulating impact of light is not established in humans. We used 7 Tesla functional magnetic resonance imaging to assess the impact of variations in light illuminance on the regional activity of the hypothalamus while healthy young adults (N=26; 16 women; 24.3 ± 2.9y) were completing two auditory cognitive tasks. We find that, during both the executive and emotional tasks, higher illuminance triggered an activity increase over the posterior part of the hypothalamus, which includes part of the tuberomamillary nucleus and the posterior part of the lateral hypothalamus. In contrast, increasing illuminance evoked a decrease in activity over the anterior and ventral parts of the hypothalamus, encompassing notably the suprachiasmatic nucleus and another part of the tuberomammillary nucleus. Critically, performance of the executive task was improved under higher illuminance and was negatively correlated with the activity of the posterior hypothalamus area. These findings reveal the distinct local dynamics of different hypothalamus regions that underlie the impact of light on cognition. They may suggest that light acts on the orexin and histamine system to affect the quality of wakefulness. ### Competing Interest Statement The authors have declared no competing interest. |
Qiongwen Cao; Michael Cohen; Akram Bakkour; Yuan Leong; Jean Decety Moral conviction interacts with metacognitive ability in modulating neural activity during sociopolitical decision-making Journal Article In: Cognitive, Affective, & Behavioral Neuroscience, pp. 1–20, 2024. @article{Cao2024a,The extent to which a belief is rooted in one's sense of morality has significant societal implications. While moral convictions can inspire positive collective action, they can also prompt dogmatism, intolerance, and societal divisions. These negative effects may be exacerbated by poor metacognition. There has been extensive research in social psychology about the characteristics of moral convictions, but their neural mechanisms and how they are incorporated into the valuation and decision-making process remain unclear. This study was designed to examine the neural mechanisms of decision-making on sociopolitical issues that vary on moral conviction, as well as how metacognitive abilities relate to these mechanisms. Participants (N = 44) underwent fMRI scanning while deciding on each trial which of two groups of political protesters they supported more. As predicted, stronger moral conviction was related to faster response times. Hemodynamic response in the anterior insula (aINS), anterior cingulate cortex (ACC), and lateral prefrontal cortex (lPFC) were elevated during decisions with higher moral conviction level, supporting both the emotional and cognitive dimensions of moral conviction. Functional connectivity between lPFC and vmPFC was also higher on trials higher in moral conviction, elucidating mechanisms through which moral conviction is incorporated into valuation. Support for protesters was positively associated with brain activity in regions involved in valuation (particularly vmPFC and amygdala). Metacognitive sensitivity, measured in a separate perceptual task, negatively correlated with parametric effects of moral conviction in the brain, providing new evidence that metacognition modulates responses to morally convicted issues. |
Benedetta Cecconi; Javier Montupil; Sepehr Mortaheb; Rajanikant Panda; Robert D. Sanders; Christophe Phillips; Naji Alnagger; Emma Remacle; Aline Defresne; Melanie Boly; Mohamed Ali Bahri; Laurent Lamalle; Steven Laureys; Olivia Gosseries; Vincent Bonhomme; Jitka Annen Study protocol: Cerebral characterization of sensory gating in disconnected dreaming states during propofol anesthesia using fMRI Journal Article In: Frontiers in Neuroscience, vol. 18, pp. 1–14, 2024. @article{Cecconi2024,Background: Disconnected consciousness describes a state in which subjective experience (i.e., consciousness) becomes isolated from the external world. It appears frequently during sleep or sedation, when subjective experiences remain vivid but are unaffected by external stimuli. Traditional methods of differentiating connected and disconnected consciousness, such as relying on behavioral responsiveness or on post-anesthesia reports, have demonstrated limited accuracy: unresponsiveness has been shown to not necessarily equate to unconsciousness and amnesic effects of anesthesia and sleep can impair explicit recollection of events occurred during sleep/sedation. Due to these methodological challenges, our understanding of the neural mechanisms underlying sensory disconnection remains limited. Methods: To overcome these methodological challenges, we employ a distinctive strategy by combining a serial awakening paradigm with auditory stimulation during mild propofol sedation. While under sedation, participants are systematically exposed to auditory stimuli and questioned about their subjective experience (to assess consciousness) and their awareness of the sounds (to evaluate connectedness/disconnectedness from the environment). The data collected through interviews are used to categorize participants into connected and disconnected consciousness states. This method circumvents the requirement for responsiveness in assessing consciousness and mitigates amnesic effects of anesthesia as participants are questioned while still under sedation. Functional MRI data are concurrently collected to investigate cerebral activity patterns during connected and disconnected states, to elucidate sensory disconnection neural gating mechanisms. We examine whether this gating mechanism resides at the thalamic level or results from disruptions in information propagation to higher cortices. Furthermore, we explore the potential role of slow-wave activity (SWA) in inducing disconnected consciousness by quantifying high-frequency BOLD oscillations, a known correlate of slow-wave activity. Discussion: This study represents a notable advancement in the investigation of sensory disconnection. The serial awakening paradigm effectively mitigates amnesic effects by collecting reports immediately after regaining responsiveness, while still under sedation. Ultimately, this research holds the potential to understand how sensory gating is achieved at the neural level. These biomarkers might be relevant for the development of sensitive anesthesia monitoring to avoid intraoperative connected consciousness and for the assessment of patients suffering from pathologically reduced consciousness. |