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!
2026 |
Benjamin G. Lowe; Alexandra Woolgar; Sophie Smit; Anina N. Rich Using EEG to detect lapses in sustained attention to moving stimuli Journal Article In: Cortex, vol. 195, pp. 1–14, 2026. @article{Lowe2026,Sustaining attention is effortful but crucial for daily life. Despite this, attentional lapses are common and can have fatal consequences (e.g., when driving). The spontaneous nature of these lapses makes studying their underlying phenomena elusive. As such, methods capable of determining when lapses have occurred may be fruitful research tools, with the potential to save lives if implemented within real-world settings. Here, we capitalised on a recent hierarchical classification method, which uses multivariate decoding to index how well human observers sustain their attention within a dynamic visual environment. We asked whether this method could be used to anticipate behavioural errors based on neural activity measured with electroencephalography (EEG |
Estelle Raffin; Roberto F. Salamanca-Giron; Krystel R. Huxlin; Olivier Reynaud; Loan Mattera; Roberto Martuzzi; Friedhelm C. Hummel Causal disconnectomics of motion perception networks: Insights from transcranial magnetic stimulation-induced BOLD responses Journal Article In: The Journal of Physiology, vol. 604, pp. 503–526, 2026. @article{Raffin2026,Understanding how focal perturbations trigger large-scale network reorganization is essential for uncovering the neural mechanisms that support perception and behaviour. Here we used a transcranial magnetic stimulation (TMS) perturbational approach by applying brief 10 Hz TMS to early visual areas (EVAs) or the medio-temporal (MT) area in healthy participants while recording concurrent functional magnetic resonance imaging (fMRI). TMS delivered during the early stages of motion processing specifically impaired direction discrimination at both sites,whereas disruption of the later processing phase impaired performances only for the MT condition. Despite a similar local increase in BOLD activity induced by EVA and MT stimulation, the broader network responses diverged significantly. Perturbation ofEVA elicited a more robust and efficient pattern of functional reorganization, manifesting as more constrained BOLD changes, consistent with greater resilience to focal disruption. In contrast behavioural impairments induced by MT stimulation were accompanied by a disorganized and less-efficient network configuration, characterized by smaller small-world properties and longer path lengths. The decrease in performances induced by MT stimulation scaled with lower clustering coefficients, implying a more random or decentralized network structure. These findings demonstrate that TMS-fMRI coupling provides a powerful framework for causally mapping the relationships between local neural perturbations, large-scale network dynamics and behavioural performance. |
Sara LoTemplio; Jack Silcox; David L. Strayer; Brennan R. Payne Single‐trial relationships between the error‐related negativity, pe, error‐related pupillary dilation response, and post‐error behavior Journal Article In: Psychophysiology, vol. 63, no. 1, 2026. @article{LoTemplio2026,The amplitude of the error‐related negativity (ERN) is known to be correlated with attention to task and general cognitive control abilities. However, previous research has struggled to consistently link ERN amplitude with behavioral accuracy or reaction time in the task from which the ERN is being measured. This lack of relationship could be due to many factors that are difficult to control for, so explorations of other converging measures to understand error‐processing and subsequent behavior adjustment are warranted. The current study examines how two other physiological markers of error‐processing—the phasic pupillary dilation response (PDR) and the positivity following an error (Pe)—relate to post‐error behavior. Additionally, we also examine relationships between the three physiological indices of error‐processing. In the study, EEG and pupillometry were simultaneously recorded while participants completed 24 blocks (50 trials each) of an Ericksen Flanker task. For post‐error accuracy, we found that on a single‐trial level, the amplitude of all three physiological error‐processing indices for error trials predicted post‐error accuracy. At the subject level, only the PDR predicted average post‐error accuracy. For post‐error slowing, at the single‐trial level, only the Pe predicted post‐error slowing, whereas only the ERN predicted post‐error slowing at the subject level. We also found that both the ERN and Pe correlated with PDR amplitude. This is consistent with our hypothesis that the Pe and PDR may share underlying neural mechanisms, but qualified by the fact that the ERN, which is not hypothesized to have shared neural mechanisms, also predicted unique variance in pupillary amplitude. Collectively, these results suggest that the PDR and Pe might represent promising indicators of post‐error behavior adjustment and highlight the need to examine relationships at multiple levels of analysis. |
Thomas Seacrist; Elizabeth A. Walshe; Shukai Cheng; Emily Brown; Charlotte Birnbaum; Victoria Kaufman; Flaura K. Winston; William C. Gaetz A novel paradigm for identifying eye-tracking metrics associated with cognitive control during driving through MEG neuroimaging Journal Article In: Transportation Research Part F: Traffic Psychology and Behaviour, vol. 116, pp. 1–13, 2026. @article{Seacrist2026,Understanding the neurocognitive underpinnings of driving behavior in adolescents is critical to improving road safety. To address this, we established a novel paradigm linking magnetoencephalography (MEG)-recorded frequency-specific brain activity to simulated driving performance, identifying periods of increased cognitive control. However, this initial paradigm did not incorporate eye-tracking – a potentially scalable proxy for cognitive control that could be leveraged by in-vehicle driver monitoring systems. This proof-of-concept study expands our paradigm by integrating eye-tracking to identify scanning behavior metrics associated with periods of increased cognitive control validated by MEG. Typically developing adolescents (n = 11; mean age = 15.1 ± 1.5 yrs) completed three driving tasks of varying cognitive demand, and MEG frequency specific analysis confirmed periods of high (Hi) and low (Lo) cognitive control via the established biomarker of frontal midline theta (FMT). Fixation count, fixation duration, horizontal/vertical mean gaze position, saccade amplitude, and horizontal/vertical spread of search were compared between Hi vs. Lo periods of cognitive control. Task-specific differences in fixation count (p < 0.05), mean gaze position (p < 0.01), saccade amplitude (p < 0.05), and spread of search (p < 0.01) were observed between Hi compared to Lo cognitive control periods. These differences corresponded to expected task-specific changes in scanning behavior that would accompany cognitive control over behavior, suggesting a signal that eye-tracking may serve as a proxy for underlying neurocognitive processes. This integrated approach demonstrates methodological rigor and offers a promising framework for further research and informing development of in-vehicle driver monitoring systems for detecting cognitive deficits in real time, with implications for enhancing teen driver safety. |
Wiktor Wicecławski; Jakub Paszulewicz ERP evidence of attentional selection outside of effective oculomotor range Journal Article In: Experimental Brain Research, vol. 244, no. 1, pp. 1–9, 2026. @article{Wiȩclawski2026,The close link between visual attention and the oculomotor system is well documented. Within the selection-for-action framework, two perspectives exist. According to Visual Attention Model (VAM) attention is seen as a prerequisite for successful movement execution, though it is considered a distinct cognitive and neural process. By contrast, the premotor theory of attention (PMTA) argues that the beneficial effects of attention are fully accounted for by the system's preparation for saccadic eye movements. From this standpoint, a central prediction emerges: attentional advantages should be confined to regions within the oculomotor range, since saccadic planning is not feasible outside those limits. A common way to examine this prediction is to present cues and targets in a hemifield beyond the oculomotor range, typically achieved by occluding one eye while abducting the other. Using this method, Smith et al. showed that in a visual search task, exogenous orienting is reduced in the temporal hemifield when the eye is abducted. They concluded that exogenous attentional orienting is constrained by the range of potential saccadic movements. In our study, we sought to replicate Smith et al.'s findings while extending the paradigm with EEG recordings—an approach not yet applied in this context. PMTA predicts that, under eye abduction, stimuli appearing in the temporal hemifield would yield diminished N2pc amplitudes. An ANOVA revealed no reduction of N2pc amplitude in the temporal hemifield. Taken together, our results support the growing body of evidence suggesting that visual attention is not strictly bound to the oculomotor range. |
2025 |
Jingnan Du; Vaibhav Tripathi; Maxwell L. Elliott; Joanna Ladopoulou; Wendy Sun; Mark C. Eldaief; Randy L. Buckner Within-individual precision mapping of brain networks exclusively using task data Journal Article In: Neuron, vol. 113, pp. 4069–4083, 2025. @article{Du2025,Precision mapping of brain networks within individuals prevailingly relies on functional connectivity analysis of resting-state data. Here, we explored whether networks can be estimated using only task data. Correlation matrices estimated from task data were similar to those derived from resting-state data. The largest factor affecting similarity was the amount of data. Precision networks estimated from task data showed strong spatial overlap with those derived from resting-state data and predicted the same triple functional dissociation in independent data. To illustrate novel possibilities enabled by the present methods, we mapped the detailed organization of thalamic association zones within individuals by pooling extensive resting-state and task data. We also demonstrated how task data can be used to estimate networks while simultaneously extracting task responses. Broadly, these findings suggest that there is an underlying, stable network architecture that is idiosyncratic to the individual and persists across task states. |
Mariana Ferreira; João Pedro Marques; Miguel Raimundo; Hugo Quental; Miguel Castelo-Branco Improvements induced by retinal gene therapy with voretigene neparvovec depend on visual cortical hemispheric dominance mechanisms Journal Article In: Communications Medicine, vol. 5, no. 1, pp. 1–9, 2025. @article{Ferreira2025,Background: RPE65-associated retinal degeneration (RPE65-RD) causes severe visual deficits. Gene therapy with AAV2-hRPE65v2 is a breakthrough but it is currently unknown which visual pathways benefit from treatment and if cortical mechanisms can amplify retinal improvements. Methods: In this within-subject design, ten patients with biallelic RPE65-RD underwent sub-retinal injection of AAV2-hRPE65v2. Psychophysical full-field stimulus threshold determination and functional magnetic resonance imaging were performed before and 12 months after treatment. Population receptive fields (pRF) were computed in V1 and visual responses assessed using contrast-reversed checkerboards (3 contrast levels). Results: Here we show significant improvement in light sensitivity at low-luminance and neural response enhancements under low-luminance conditions specifically in the right hemisphere, which is known to show dominance in attentional and visual pooling of spatial information. Changes in pRF size also reflect known hemispheric spatial asymmetries (left/right biased for local/global analysis, respectively). Conclusions: Our findings show a contribution of known early and high-level cortical dominance mechanisms on improvement, which constrain the effects of therapy and are therefore a target for neurorehabilitation. These findings provide insight into the limits of clinical benefits of gene therapy and suggest that neurorehabilitation approaches may be needed to enhance improvements, similarly to cochlear implants. |
Luise P. Graichen; Magdalena S. Linder; Lars Keuter; Ole Jensen; Christian F. Doeller; Claus Lamm; Tobias Staudigl; Isabella C. Wagner Entorhinal grid-like codes for visual space during memory formation Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–15, 2025. @article{Graichen2025,Eye movements, such as saccades, allow us to gather information about the environment and, in this way, can shape memory. In non-human primates, saccades are associated with the activity of grid cells in the entorhinal cortex. Grid cells are essential for spatial navigation, but whether saccade-based grid-like signals play a role in human memory formation is currently unclear. Here, human participants undergo functional magnetic resonance imaging and continuous eye gaze monitoring while studying scene images. Recognition memory is probed immediately thereafter. Results reveal saccade-based grid-like codes in the left entorhinal cortex that are specific to later remembered trials during study, a finding that we replicate with an independent data set. The grid-related effects are time-locked to activation increases in the frontal eye fields. Unexpectedly, lower saccade-based grid-like codes are associated with better subsequent recognition memory performance. Our findings suggest an entorhinal map of visual space that is timed with neural activity in oculomotor regions, and negatively associated with subsequent memory. Grid-like codes, entorhinal cortex, saccades, frontal eye fields (FEF), memory, functional magnetic resonance imaging (fMRI) |
Chanyuan Gu; Samuel A. Nastase; Zaid Zada; Ping Li Reading comprehension in L1 and L2 readers: Neurocomputational mechanisms revealed through large language models Journal Article In: npj Science of Learning, vol. 10, no. 1, pp. 1–13, 2025. @article{Gu2025,While evidence has accumulated to support the argument of shared computational mechanisms underlying language comprehension between humans and large language models (LLMs), few studies have examined this argument beyond native-speaker populations. This study examines whether and how alignment between LLMs and human brains captures the homogeneity and heterogeneity in both first-language (L1) and second-language (L2) readers. We recorded brain responses of L1 and L2 English readers of texts and assessed reading performance against individual difference factors. At the group level, the two groups displayed comparable model-brain alignment in widespread regions, with similar unique contributions from contextual embeddings. At the individual level, multiple regression models revealed the effects of linguistic abilities on alignment for both groups, but effects of attentional ability and language dominance status for L2 readers only. These findings provide evidence that LLMs serve as cognitively plausible models in characterizing homogeneity and heterogeneity in reading across human populations. |
Bao Li; Li Tong; Chi Zhang; Panpan Chen; Long Cao; Hui Gao; Zi Ya Yu; Lin Yuan Wang; Bin Yan An fMRI dataset on occluded image interpretation for human amodal completion research Journal Article In: Scientific Data, vol. 12, no. 1, pp. 1–10, 2025. @article{Li2025b,In everyday environments, partially occluded objects are more common than fully visible ones. Despite their visual incompleteness, the human brain can reconstruct these objects to form coherent perceptual representations, a phenomenon referred to as amodal completion. However, current computer vision systems still struggle to accurately infer the hidden portions of occluded objects. While the neural mechanisms underlying amodal completion have been partially explored, existing findings often lack consistency, likely due to limited sample sizes and varied stimulus materials. To address these gaps, we introduce a novel fMRI dataset,the Occluded Image Interpretation Dataset (OIID), which captures human perception during image interpretation under different levels of occlusion. This dataset includes fMRI responses and behavioral data from 65 participants. The OIID enables researchers to identify the brain regions involved in processing occluded images and examines individual differences in functional responses. Our work contributes to a deeper understanding of how the human brain interprets incomplete visual information and offers valuable insights for advancing both theoretical research and related practical applications in amodal completion fields. |
Tina T. Liu; Michael C. Granovetter; Anne Margarette Anne; Sophia Robert; Jason Z Fu; Christina Patterson; David C. Plaut; Marlene Behrmann Cross-sectional and longitudinal changes in category selectivity in visual cortex following pediatric cortical resection Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–18, 2025. @article{Liu2025p,The topographic organization of category-selective responses in human ventral occipitotemporal cortex (VOTC) and its relationship to regions subserving language functions is remarkably uniform across individuals. This arrangement is thought to result from the clustering of neurons responding to similar inputs, constrained by intrinsic architecture and tuned by experience. We examine the malleability of this organization in individuals with unilateral resection of VOTC during childhood for the management of drug-resistant epilepsy. In cross-sectional and longitudinal functional imaging studies, we compare the topography and neural representations of 17 category-selective regions in individuals with a VOTC resection, a ‘control patient' with a resection outside VOTC, and typically developing matched controls. We demonstrate both adherence to and deviation from the standard topography, particularly with respect to the hemispheric lateralization of category-selective regions, and uncover fine-grained competitive dynamics between word- and face-selectivity over time in the single, preserved VOTC. The findings elucidate the nature and extent of cortical plasticity and highlight the potential for remodeling of extrastriate architecture and function. |
Nasrin Mortazavi; Puneet Talwar; Ekaterina Koshmanova; Roya Sharifpour; Elise Beckers; Alexandre Berger; Islay Campbell; Ilenia Paparella; Fermin Balda; Ismael Dardour Hamzaoui; Christian Berthomier; Christine Bastin; Christophe Phillips; Pierre Maquet; Fabienne Collette; Mikhail Zubkov; Laurent Lamalle; Gilles Vandewalle REM sleep quality is associated with balanced tonic activity of the locus coeruleus during wakefulness Journal Article In: Journal of Biomedical Science, vol. 32, no. 1, pp. 1–13, 2025. @article{Mortazavi2025,Background: Animal studies established that the locus coeruleus (LC) plays important roles in sleep and wakefulness regulation. Whether it contributes to sleep variability in humans is not yet established. Here, we investigated if the in vivo activity of the LC is related to the variability in the quality of Rapid Eye Movement (REM) sleep. Methods: We assessed the LC activity of 34 healthy younger (~ 22y) and 18 older (~ 61y) individuals engaged in bottom-up and top-down cognitive tasks using 7-Tesla functional Magnetic Resonance Imaging (fMRI). We further recorded their sleep electroencephalogram (EEG) to evaluate associations between LC fMRI measures and REM sleep EEG metrics. Results: Theta oscillation energy during REM sleep was positively associated with LC response in the top-down task. In contrast, REM sleep theta energy was negatively associated with LC activity in older individuals during the bottom-up task. Importantly, sigma oscillations power immediately preceding a REM sleep episode was positively associated with LC activity in the top-down task. Conclusions: LC activity during wakefulness was related to REM sleep intensity and to a transient EEG change preceding REM sleep, a feature causally related to LC activity in animal studies. The associations depend on the cognitive task, suggesting that a balanced level of LC tonic activity during wakefulness is required for optimal expression of REM sleep. The findings may have implications for the high prevalence of sleep complaints reported in aging and for disorders such as insomnia, Alzheimer's, and Parkinson's disease, for which the LC may play pivotal roles through sleep. |
Renana Storm; Viktoria Wrobel; Antonia Frings; Andreas Sprenger; Christoph Helmchen In: Scientific Reports, vol. 15, no. 1, pp. 1–11, 2025. @article{Storm2025,Persistent postural-perceptual dizziness (PPPD) is often preceded by vestibular disorders. We applied galvanic vestibular stimulation (GVS) and related stimulus-evoked activity to individual ratings of perceived motion for each stimulus and to perceived egomotion thresholds by GVS and behavioural parameters outside the scanner: levels of functional disability by standardized questionnaires, visual motion coherence, passive egomotion perception by chair rotation and quantitative postural stability. We hypothesized that the preceding vestibular disorder predisposes to abnormal brain excitability by vestibular stimulation. All participants showed normal vestibular function tests on quantitative testing. GVS with different intensities was applied to 28 patients and 28 age- and gender-matched healthy participants (HC) in the scanner. After each stimulus, participants rated their perceived level of egomotion. GVS perception threshold was significantly lower in PPPD patients. Contrasting stimulus-identical GVS against a sham stimulus, group comparison revealed a stronger activation in the patient's supramarginal gyrus, insular cortex (operculum 3), and vermis. This stronger excitability was not related to the individual threshold of perceived egomotion by GVS. Patients rated GVS-evoked egomotion intensity by identical GVS intensities larger than HC but neural activity did not correlate with individual ratings of perceived egomotion by GVS. As GVS evoked larger egomotion and larger brain activation in patients, the ratio of brain activity to egomotion perception was not different between groups. GVS-evoked insular activity increased with the level of PPPD-related disability and postural imbalance. The larger activation in multisensory cortical vestibular network indicates a sensitization to vestibular stimuli eliciting egomotion perception which increases with levels of PPPD disability. It seems to reflect a sensory-neural amplification rather than an abnormal sensory-perceptual scaling. |
Ekin Tünçok; Marisa Carrasco; Jonathan Winawer Spatial attention selectively alters visual cortical representation during target anticipation Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–19, 2025. @article{Tuencok2025,Attention enables us to efficiently and flexibly interact with the environment by prioritizing specific image locations and features in preparation for responding to stimuli. Using a concurrent psychophysics–fMRI experiment, we investigate how covert spatial attention modulates responses in human visual cortex before target onset and how it affects subsequent behavioral performance. Performance improves at cued locations and worsens at uncued locations compared to distributed attention, demonstrating a selective processing tradeoff. Pre-target BOLD responses in cortical visual field maps reveal two key changes: First, a stimulus-independent baseline shift, with increases near cued locations and decreases elsewhere, paralleling behavioral results. Second, a shift in population receptive field centers toward the attended location. Both effects increase in higher visual areas. Together, these findings reveal that spatial attention has large effects on visual cortex prior to target appearance, altering neural response properties across multiple visual field maps and enhancing performance through anticipatory mechanisms. |
Zinong Yang; Stephanie D. Williams; Ewa Beldzik; Stephanie Anakwe; Emilia Schimmelpfennig; Laura D. Lewis Attentional failures after sleep deprivation are locked to joint neurovascular, pupil and cerebrospinal fluid flow dynamics Journal Article In: Nature Neuroscience, pp. 2526–2536, 2025. @article{Yang2025e,Sleep deprivation rapidly disrupts cognitive function and in the long term contributes to neurological disease. Why sleep deprivation has such profound effects on cognition is not well understood. Here we use simultaneous fast fMRI–EEG to test how sleep deprivation modulates cognitive, neural and fluid dynamics in the human brain. We demonstrate that attentional failures during wakefulness after sleep deprivation are tightly orchestrated in a series of brain–body changes, including neuronal shifts, pupil constriction and cerebrospinal fluid (CSF) flow pulsations, pointing to a coupled system of fluid dynamics and neuromodulatory state. CSF flow and hemodynamics are coupled to attentional function within the awake state, with CSF pulsations following attentional impairment. The timing of these dynamics is consistent with a vascular mechanism regulated by neuromodulatory state. The attentional costs of sleep deprivation may thus reflect an irrepressible need for rest periods driven by a central neuromodulatory system that regulates both neuronal and fluid physiology. |
Mrugank Dake; Clayton E. Curtis Perturbing human V1 degrades the fidelity of visual working memory Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–8, 2025. @article{Dake2025,Decades of macaque research established the importance of prefrontal cortex for working memory. Surprisingly, recent human neuroimaging studies demonstrated that the contents of working memory can be decoded from primary visual cortex (V1). However the necessity of this mnemonic information remains unknown and contentious. Here we provide causal evidence that transcranial magnetic stimulation targeting human V1 disrupted the fidelity of visual working memory. Errors increased only for targets remembered in the portion of the visual field disrupted by stimulation. Moreover, concurrently measured electroencephalography confirmed that stimulation disrupted not only memory behavior, but neurophysiological signatures of working memory. These results change the question from whether visual cortex is necessary for working memory to what mechanisms it uses to support memory. Moreover, they point to models in which the mechanisms supporting working memory are distributed across brain regions, including sensory areas that here we show are critical for memory storage. |
Burcu Bayram; David Meijer; Roberto Barumerli; Michelle Spierings; Robert Baumgartner; Ulrich Pomper Bayesian prior uncertainty and surprisal elicit distinct neural patterns during sound localization in dynamic environments Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–18, 2025. @article{Bayram2025,Estimating the location of a stimulus is a key function in sensory processing, and widely considered to result from the integration of prior information and sensory input according to Bayesian principles. A deviation of sensory input from the prior elicits surprisal, depending on the uncertainty of the prior. While this mechanism is increasingly understood in the visual domain, much less is known about its implementation in audition, especially regarding spatial localization. Here, we combined human EEG with computational modeling to study auditory spatial inference in a noisy, volatile environment and analyzed behavioral and neural patterns associated with prior uncertainty and surprisal. First, our results demonstrate that participants indeed used prior information during periods of stable environmental statistics, but showed evidence of surprisal and discarded prior information following environmental changes. Second, we observed distinct EEG activity patterns associated with prior uncertainty and surprisal in both the time- and time–frequency domain, which are in line with previous studies using visual tasks. Third, these EEG activity patterns were predictive of our participants' sound localization error, response uncertainty, and prior bias on a trial-by-trial basis. In summary, our work provides novel behavioral and neural evidence for Bayesian inference during dynamic auditory localization. |
Lixiang Chen; Radoslaw Martin Cichy; Daniel Kaiser Representational shifts from feedforward to feedback rhythms index phenomenological integration in naturalistic vision Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–5, 2025. @article{Chen2025f,How does the brain integrate complex and dynamic visual inputs into phenomenologically seamless percepts? Previous results demonstrate that when visual inputs are organized coherently across space and time, they are more strongly encoded in feedback-related alpha rhythms, and less strongly in feedforward-related gamma rhythms. Here, we tested whether this representational shift from feedforward to feedback rhythms is linked to the phenomenological experience of coherence. In an Electroencephalography (EEG) study, we manipulated the degree of spatiotemporal coherence by presenting two segments from the same video across visual hemifields, either synchronously or asynchronously (with a delay between segments). We asked participants whether they perceived the stimulus as coherent or incoherent. When stimuli were presented at the perceptual threshold (i.e., when the same stimulus was judged as coherent 50% of times), perception co-varied with stimulus coding across alpha and gamma rhythms: When stimuli were perceived as coherent, they were represented in alpha activity; when stimuli were perceived as incoherent, they were represented in gamma activity. Whether the same visual input is perceived as coherent or incoherent thus depends on representational shifts between feedback-related alpha and feedforward-related gamma rhythms. |
Andrew W. Corcoran; Arthur Le Coz; Jakob Hohwy; Thomas Andrillon When your heart isn't in it anymore: Cardiac correlates of task disengagement Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–16, 2025. @article{Corcoran2025,Neuroscience is beginning to uncover the role of interoceptive feedback in perception, learning, and decision-making; however, the relation between spontaneous visceral and cognitive dynamics has received surprisingly little scrutiny. Here, we investigate how subjective, physiological, and behavioural indicators of arousal and attentional state vary in relation to ongoing cardiac activity and brain-heart coupling. Electroencephalogram, electrocardiogram, and pupillometric records were obtained from 65 adults during the performance of a sustained attention to response task (SART). Thought probes were intermittently administered during the SART to collect subjective reports of attentional state (on-task, mind-wandering, mind-blanking) and vigilance level (alertness vs. sleepiness). Mind-wandering and mind-blanking reports increased in frequency with time-on-task and were accompanied by decreases in alertness and pupil-linked arousal, but evinced distinct psychophysiological and behavioural profiles: While mind-wandering was associated with greater heart-rate variability and late modulation of the heartbeat-evoked potential, mind-blanking was characterised by more profound decreases in heart-rate, pupil size, and brain-heart coupling. Lower heart-rate predicted decreased vigilance and pupil size, in addition to slower, less-biased responses; increased heart-rate variability predicted more impulsive behaviour and pupil dilation. Together, these findings reveal that cardiac parameters and brain-heart connectivity measures afford complementary information about arousal states and attentional dynamics during task performance. |
Katharina Duecker; Kimron L. Shapiro; Simon Hanslmayr; Benjamin J. Griffiths; Yali Pan; Jeremy M. Wolfe; Ole Jensen Guided visual search is associated with target boosting and distractor suppression in early visual cortex Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–11, 2025. @article{Duecker2025,Visual attention paradigms have revealed that neural excitability in higher-order visual areas is modulated according to a priority map guiding attention towards task-relevant locations. Neural activity in early visual regions, however, has been argued to be modulated based on bottom-up salience. Here, we combined Magnetoencephalography (MEG) and Rapid Invisible Frequency Tagging (RIFT) in a classic visual search paradigm to study feature-guidance in early human visual cortex. Our results demonstrate evidence for both target boosting and distractor suppression when the participants were informed about the task-relevant and -irrelevant colour (guided search) compared to when they were not (unguided search). These results conceptually replicated using both a magnitude-squared coherence approach and a General Linear Model based on a single-trial measure of the RIFT response. The present findings reveal that feature-guidance in visual search affects neuronal excitability as early as primary visual cortex, possibly contributing to a priority-map-based mechanism. |
Matthias Grabenhorst; David Poeppel; Georgios Michalareas Neural signatures of temporal anticipation in human cortex represent event probability density Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–17, 2025. @article{Grabenhorst2025,Temporal prediction is a fundamental function of neural systems. Recent results show that humans anticipate future events by calculating probability density functions, rather than hazard rates. However, direct neural evidence for this hypothesized mechanism is lacking. We recorded neural activity using magnetoencephalography as participants anticipated auditory and visual events distributed in time. We show that temporal anticipation, measured as reaction times, approximates the event probability density function, but not hazard rate. Temporal anticipation manifests as spatiotemporally patterned activity in three anatomically and functionally distinct parieto-temporal and sensorimotor cortical areas. Each of these areas revealed a marked neural signature of anticipation: Prior to sensory cues, activity in a specific frequency range of neural oscillations, spanning alpha and beta ranges, encodes the event probability density function. These neural signals predicted reaction times to imminent sensory cues. These results demonstrate that supra-modal representations of probability density across cortex underlie the anticipation of future events. |
Jason Helbing; Dejan Draschkow; Melissa L. H. Võ Incidental encoding of objects during search is stronger than intentional memorization due to increased recollection rather than familiarity Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 12, pp. 2538–2557, 2025. @article{Helbing2025,Most memory is not formed deliberately but as a by-product of natural behavior. These incidental representations, when generated during visual search, can be stronger than intentionally memorized content (search superiority effect). However, it is unknown if the search superiority effect is purely quantitative (stronger memory) or also driven by differences in the degrees of recollection and familiarity, two hallmark processes supporting recognition memory. Here, we use signal detection modeling, introspective judgments, event-related EEG potentials, and eye tracking measures to answer this question. In a preregistered study, 30 participants searched for objects in scenes and intentionally memorized others before completing a surprise recognition memory test. Behavioral data from remember-know judgments and receiver operating characteristics indicate that search targets were more often recollected compared with intentionally memorized objects, whereas the two tasks did not lead to differences in familiarity. Surprisingly, the neural signatures did not fully align with the behavioral findings regarding recollection and familiarity. That is, both search targets and intentionally memorized objects elicited a more positive-going mid-frontal negativity peaking at around 400 msec post stimulus onset (FN400), which is associated with familiarity, as well as a more positive-going parietal late component (LPC), indicative of recollection. Both components showed no differences between tasks, indicating equal contributions of recollection and familiarity to remembering searched and memorized objects. Furthermore, the LPC was, as expected, sensitive to differences between recollected and familiar objects when these were intentionally memorized, but it was not affected by these differences for searched objects. Overall, our findings indicate that search superiority relies predominantly on increased recollection. The fact that established neural markers of recollection (LPC) behaved as anticipated for intentionally memorized objects but carried no predictive power for incidentally memorized objects implies that memories established in more ecologically valid tasks might involve neural processes different from those activated in commonly used settings that are more reductionist. |
Dorottya Hetenyi; Joost Haarsma; Peter Kok Contents of visual predictions oscillate at alpha frequencies Journal Article In: The Journal of Neuroscience, vol. 45, no. 49, pp. 1–12, 2025. @article{Hetenyi2025,Predictions of future events have a major impact on how we process sensory signals. However, it remains unclear how the brain keeps predictions online in anticipation of future inputs. Here, we combined magnetoencephalography (MEG) and multivariate decoding techniques to investigate the content of perceptual predictions and their frequency characteristics. Thirty-two participants (23 female) were engaged in a shape discrimination task, while auditory cues predicted which specific shape would likely appear. Frequency analysis revealed significant oscillatory fluctuations of predicted shape representations in the pre-stimulus window in the alpha band (10–11 Hz). Furthermore, we found that this stimulus-specific alpha power was linked to expectation effects on shape discrimination behavior. Our findings demonstrate that sensory predictions are embedded in pre-stimulus alpha oscillations and modulate subsequent perceptual performance, providing a neural mechanism through which the brain deploys perceptual predictions. |
Xin Huang; Hezul Tin Yan Ng; Chien Ho Lin; Ming Yan; Olaf Dimigen; Werner Sommer; Urs Maurer How the dominant reading direction changes parafoveal processing: A combined EEG/eye-tracking study Journal Article In: Psychophysiology, vol. 62, no. 12, pp. 1–22, 2025. @article{Huang2025e,Reading directions vary across writing systems. Through long-term experience, readers adjust their visual systems to the dominant reading direction in their writing systems. However, little is known about the neural correlates underlying these adjustments because different writing systems do not just differ in reading direction, but also in visual and linguistic properties. Here, we took advantage of the fact that Chinese is read to different degrees in left-to-right or top-to-bottom directions in different regions. We investigated visual word processing in participants from Taiwan (both top-to-bottom and left-to-right directions) and from mainland China (only left-to-right direction). We used combined EEG/eye-tracking with a saccade-contingent parafoveal preview manipulation to investigate how the dominant reading direction shapes neural visual processing while participants read 5-word lists. Fixation-related potentials (FRPs) showed a reduced late N1 effect (preview positivity), but this effect was modulated by prior experience with a specific reading direction. Results replicated previous findings that valid previews facilitate visual word processing, as indicated by reduced FRP activation. Critically, the results provide the first neuroelectric evidence that this facilitation effect depends on experience with a given reading direction. The findings provide insight into how cultural experience shapes the way people process visual information and demonstrate how a person's everyday visual experience can influence how the brain processes parafoveal information. |
Dirk Kerzel Electrophysiological evidence for the optimal tuning of attention Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–14, 2025. @article{Kerzel2025,Optimal tuning of attention refers to shifts in goal-driven attention that increase the difference between the representation of the target and nontarget features. Evidence for optimal tuning comes from studies measuring the memory representation of the target and, to a lesser degree, from studies measuring attentional selectivity. In one study on attentional selectivity, cueing effects were found to be greater for cue colors deviating away from the nontarget color compared to cue colors deviating toward the nontarget color, suggesting that participants' search goal was optimally tuned. To address alternative accounts, we measured event-related potentials (ERPs) elicited by different cue colors at posterior electrodes PO7/PO8. We found that ERPs associated with attentional orienting (N1pc) or selection (N2pc) were larger for cue colors deviating away from the nontarget color, which is consistent with the optimal tuning of attention. In contrast, the results are difficult to reconcile with alternative accounts such as rapid disengagement or object updating. Further, we aimed to evaluate contributions from sensory adaptation by analyzing the Ppc component, a lateralized ERP in the P1 time range. Two control conditions, however, suggested that the Ppc was more likely driven by imbalanced saliency than sensory adaptation. |
Guoyang Liu; Yueyuan Zheng; Michelle Hei Lam Tsang; Yazhou Zhao; Janet H. Hsiao Understanding the role of eye movement pattern and consistency during face recognition through EEG decoding Journal Article In: npj Science of Learning, vol. 10, no. 1, pp. 1–13, 2025. @article{Liu2025i,Eye movement patterns and consistency during face recognition are both associated with recognition performance. We examined whether they reflect different mechanisms through EEG decoding. Eighty-four participants performed an old-new face recognition task with eye movement pattern and consistency quantified using eye movement analysis with hidden Markov models (EMHMM). Temporal dynamics of neural representation quality for face recognition were assessed through decoding old vs new faces using a support vector machine classifier. Results showed that a more eye-focused pattern was associated with higher decoding accuracy in the high-alpha band, reflecting better neural representation quality. In contrast, higher eye movement consistency was associated with shorter latency of peak decoding accuracy in the high-alpha band, which suggested more efficient neural representation development, in addition to higher ERP decoding accuracy. Thus, eye movement patterns are associated with neural representation effectiveness, whereas eye movement consistency reflects neural representation development efficiency, unraveling different aspects of cognitive processes. |
Marie Loescher; Patrick Haggard; Catherine Tallon-Baudry Interoception vs. exteroception: Cardiac interoception competes with tactile perception, yet also facilitates self-relevance encoding Journal Article In: PNAS, vol. 122, no. 49, pp. 1–12, 2025. @article{Loescher2025,Internal bodily signals, notably the heartbeat, influence our perception of the external world—but the nature of this influence remains unclear. Different frameworks, originating in opposing views of the function of interoception, have developed largely in parallel. One line of evidence (Internal/External Competition) indicates that interoceptive and exteroceptive inputs compete for neural resources. Another line (Self-related Facilitation) shows a link between interoceptive and self-related processing, which might include computing the self-relevance of exteroceptive inputs. We contrasted these accounts within a single experimental task for which they yielded distinct predictions. We measured heartbeat-evoked potentials (HEPs, a measure of cardiac interoception) with electroencephalogram and manipulated the self-relevance of an audio-tactile stimulus by placing the audio source either inside or outside the peripersonal space immediately around the body. On the one hand, prestimulus HEP amplitudes over the somatosensory cortex were linked to slower reaction times and affected audio-tactile stimulus-evoked responses in the same area, indicating competition for shared neural resources. On the other hand, prestimulus HEPs over integrative sensorimotor and default-mode network regions facilitated stimulus self-relevance encoding, both in reaction times and audio-tactile evoked responses. Importantly, Competition and Facilitation effects were spatially and statistically independent from each other. We therefore reconcile the two views by showing the coexistence of two independent mechanisms: one that allocates neural resources to either internal bodily signals or the external world, and another by which interoception and exteroception are combined to determine the self-relevance of external signals. Our results highlight the multidimensionality of HEPs and of internal states more generally. |
Elisabet Parés-Pujolràs; Simon P. Kelly; Peter R. Murphy Dissociable encoding of evolving beliefs and momentary belief updates in distinct neural decision signals Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–14, 2025. @article{ParesPujolras2025,Making accurate decisions in noisy environments requires integrating evidence over time. Studies of simple perceptual decisions in static environments have identified two human neurophysiological signals that evolve with similar integration dynamics, with one - the centroparietal positivity - appearing to compute the running integral and continuously feed it to the other - motor beta lateralisation. However, it remains unknown whether and how these signals serve more distinct functional roles in more complex scenarios. Here, we use a volatile expanded judgement task that dissociates raw sensory information, belief updates, and the evolving belief itself. We find that motor beta lateralisation traces the evolving belief across stimuli, while the centroparietal positivity locally encodes the belief updates associated with each individual stimulus. These results suggest a flexible computational hierarchy where context-dependent belief updates can be computed sample-by-sample at an intermediate processing level to modify downstream belief representations for protracted decisions about discrete stimuli. |
Hame Park; Ayelet Arazi; Bharath Chandra Talluri; Marco Celotto; Stefano Panzeri; Alan A. Stocker; Tobias H. Donner Confirmation bias through selective readout of information encoded in human parietal cortex Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–15, 2025. @article{Park2025,Decision-makers often process new evidence selectively, depending on their current beliefs about the world. We asked whether such confirmation biases result from biases in wthe encoding of sensory evidence in the brain, or alternatively in the utilization of encoded evidence for behavior. Human participants estimated the source of a sequence of visual-spatial evidence samples while we measured cortical population activity with magnetoencephalography. Halfway through the sequence, participants were prompted to judge the more likely source category. We find that processing of subsequent evidence depends on its consistency with the previously chosen category. Evidence encoded in parietal cortex contributes more to the estimation report when that evidence is consistent with the previous choice compared to when it contradicts that choice. Our results indicate that information contradicting pre-existing beliefs has little impact on subsequent behavior, despite being precisely encoded in the brain. This provides room for deliberative control to counteract confirmation biases. |
Katrina R. Quinn; Florian Sandhaeger; Nima Noury; Ema Zezelic; Markus Siegel Abstract choice representations during stable choice-response associations Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–8, 2025. @article{Quinn2025,An increasing body of evidence has demonstrated neural representations of choices independent of the motor actions used to report them – so-called abstract choices. However, it remains unclear whether such representations arise due to dynamic changes in choice-response associations or reflect a general property of decision-making. Here, we show that in the human brain, choices are represented abstractly even when choice-response associations remain stable over time. We recorded neural activity using magnetoencephalography while participants performed a motion discrimination task, with choice-response mappings held constant within blocks. We found neural information about participants' perceptual choices independent of both motor response and visual stimulus. Choice information increased during the stimulus and peaked after the response. Moreover, choice and response information showed distinct cortical distributions, with choice-related signals strongest in frontoparietal regions. Thus, abstract choice representations are not limited to dynamic or action-independent contexts and may be a general feature of decision-making. |
Alia Seedat; Alex Lepauvre; Jay Jeschke; Urszula Gorska-Klimowska; Marcelo Armendariz; Katarina Bendtz; Simon Henin; Rony Hirschhorn; Tanya Brown; Erika Jensen; Csaba Kozma; David Mazumder; Stephanie Montenegro; Leyao Yu; Niccolò Bonacchi; Diptyajit Das; Kyle Kahraman; Praveen Sripad; Fatemeh Taheriyan; Orrin Devinsky; Patricia Dugan; Werner Doyle; Adeen Flinker; Daniel Friedman; Wendell Lake; Michael Pitts; Liad Mudrik; Melanie Boly; Sasha Devore; Gabriel Kreiman; Lucia Melloni Open multi-center intracranial electroencephalography dataset with task probing conscious visual perception Journal Article In: Scientific Data, vol. 12, no. 1, pp. 1–14, 2025. @article{Seedat2025,We introduce an intracranial EEG (iEEG) dataset collected as part of an adversarial collaboration between proponents of two theories of consciousness: Global Neuronal Workspace Theory and Integrated Information Theory. The data were recorded from 38 patients undergoing intracranial monitoring of epileptic seizures across three research centers using the same experimental protocol. Participants were presented with suprathreshold visual stimuli belonging to four different categories (faces, objects, letters, false fonts) in three orientations (front, left, right view), and for three durations (0.5, 1.0, 1.5 s). Participants engaged in a non-speeded Go/No-Go target detection task to identify infrequent targets with some stimuli becoming task-relevant and others task-irrelevant. Participants also engaged in a motor localizer task. The data were checked for its quality and converted to Brain Imaging Data Structure (BIDS). The de-identified dataset contains demographics, clinical information, electrode reconstruction, behavioral performance, and eye-tracking data. We also provide code to preprocess and analyze the data. This dataset holds promise for reuse in consciousness science and vision neuroscience to answer questions related to stimulus processing, target detection, and task-relevance, among many others. |
Dixit Sharma; Bart Krekelberg Predicting spiking activity from scalp EEG Journal Article In: Journal of Neural Engineering, vol. 22, no. 6, pp. 1–16, 2025. @article{Sharma2025,Objective. Despite decades of electroencephalography (EEG) research, the relationship between EEG and underlying spiking dynamics remains unclear. This limits our ability to infer neural dynamics reflected in intracranial signals from EEG, a critical step to bridge electrophysiological findings across species and to develop non-invasive brain–machine interfaces (BMIs). In this study, we aimed to estimate spiking activity in the visual cortex using non-invasive scalp EEG. Approach . We recorded spiking activity from a 32-channel floating microarray permanently implanted in parafoveal V1 and scalp-EEG in a male macaque monkey. While the animal fixated, the screen flickered at different temporal frequencies to induce steady-state visual evoked potentials. We analyzed the relationship between the V1 multi-unit spiking activity envelope (MUAe) and EEG frequency bands to predict MUAe at each time point from EEG. We extracted instantaneous spectrotemporal features of the EEG signal, including phase, amplitude, and phase-amplitude coupling of its frequency bands. Main results . Although the relationship between these spectrotemporal features and the V1 MUAe was complex and frequency-dependent, they were reliably predictive of the MUAe. Specifically, in a linear regression predicting MUAe from EEG, each EEG feature (phase, amplitude, coupling) contributed to model predictions. In addition, we found that MUAe predictions were better in shallow than deep cortical layers, and that the phase of stimulus frequency further improved MUAe predictions. Significance. Our study shows that a comprehensive account of spectrotemporal features of non-invasive EEG provides information on underlying spiking activity beyond what is available when only the amplitude or phase of the EEG signal is considered. This demonstrates the richness of the EEG signal and its complex relationship with neural spiking activity and suggests that using more comprehensive spectrotemporal signatures could improve BMI applications. |
Qiao Songlin; Xuemei Xia; Jing Chen; Matteo Valsecchi Attentional tracking reduces cortical alpha oscillations Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–14, 2025. @article{Songlin2025,The premotor theory of attention suggests that both overt and covert attentional orienting are governed by similar mechanisms and neural structures, a concept extensively investigated in paradigms involving shifts in attention and gaze towards peripheral targets. Previous studies have found a strong link between cortical alpha oscillations and overt smooth pursuit of a target. However, the relationship between alpha oscillations and covert tracking of peripheral moving stimuli remains unclear. To address this, we asked 16 observers to maintain fixation while covertly attending to a visual stimulus moving along the horizontal meridian at varying speeds (2, 6, or 12 °/s), within either the left or right hemifield. We simultaneously recorded both eye movements and EEG data. Our results revealed that alpha power was significantly reduced when observers tracked a target that moved further in the periphery, independent of its speed. These findings confirm that the distribution of alpha power is sensitive to the allocation of covert attention during tracking. This suggests a tight link between the attentional processes involved in covert tracking and overt pursuit of a moving target, supporting the premotor theory of attention. |
Caleb Stone; Jason B. Mattingley; Dragan Rangelov Neural mechanisms of metacognitive improvement under speed pressure Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–12, 2025. @article{Stone2025,The ability to accurately monitor the quality of one's choices, or metacognition, improves under speed pressure, possibly due to changes in post-decisional evidence processing. Here, we investigate the neural processes that regulate decision-making and metacognition under speed pressure using time-resolved analyses of brain activity recorded using electroencephalography. Participants performed a motion discrimination task under short and long response deadlines and provided a metacognitive rating following each response. Behaviourally, participants were faster, less accurate, and showed superior metacognition with short deadlines. These effects were accompanied by a larger centro-parietal positivity (CPP), a neural correlate of evidence accumulation. Crucially, post-decisional CPP amplitude was more strongly associated with participants' metacognitive ratings following errors under short relative to long response deadlines. Our results suggest that superior metacognition under speed pressure may stem from enhanced metacognitive readout of post-decisional evidence. |
Zhongbin Su; Xiaolin Zhou; Stefan Pollmann; Lihui Wang Dynamic face-related eye movement representations in the human ventral pathway Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–12, 2025. @article{Su2025c,Multiple brain areas along the ventral pathway have been known to represent face images. Here, in a magnetoencephalography (MEG) experiment, we show dynamic representations of face-related eye movements in the ventral pathway in the absence of image perception. Participants followed a dot presented on a uniform background, the movement of which represented gaze tracks acquired previously during their free-viewing of face and house pictures. We found a dominant role of the ventral stream in representing face-related gaze tracks, starting from the orbitofrontal cortex (OFC) and anterior temporal lobe (ATL), and extending to the medial temporal and ventral occipitotemporal cortex. Our findings show that the ventral pathway represents the gaze tracks used to explore faces, by which top-down prediction of face category in OFC and ATL may guide, via the medial temporal cortex or directly, face perception in the ventral occipitotemporal cortex. |
Lijuan Wang; Steven Frisson; Yali Pan; Ole Jensen Fast hierarchical processing of orthographic and semantic parafoveal information during natural reading Journal Article In: Nature Communications, vol. 16, no. 1, pp. 1–12, 2025. @article{Wang2025f,In reading, information from parafoveal words is extracted before direct fixation; however, it is debated whether this processing is restricted to orthographic features or also encompasses semantics. Moreover, the neuronal mechanisms supporting parafoveal processing remain poorly understood. We co-registered MEG and eye-tracking data in a natural reading paradigm to uncover the timing and brain regions involved in parafoveal processing. Representational similarity analysis revealed that parafoveal orthographic neighbours (e.g., “writer” vs. “waiter”) showed higher representational similarity than non-neighbours (e.g., “writer” vs. “police”), emerging ~68 ms after fixation onset on the preceding word (e.g., “clever”) in the visual word form area. Similarly, parafoveal semantic neighbours (e.g., “writer” vs. “author”) exhibited increased representational similarity at ~137 ms in the left inferior frontal gyrus. Importantly, the degree of orthographic and semantic parafoveal processing was correlated with individual reading speed. Our findings suggest fast hierarchical processing of parafoveal words across distinct brain regions, enhancing reading efficiency. |
Xin Wang; Shitao Chen; Keyang Wang; Liyu Cao Predicted action-effects shape action representation through pre-activation of alpha oscillations Journal Article In: Communications Biology, vol. 8, no. 1, pp. 1–11, 2025. @article{Wang2025n,Actions are typically accompanied by sensory feedback (or action-effects). Action-effects, in turn, influence the action. Theoretical accounts of action control assume a pre-activation of action-effects prior to action execution. Here we show that when participants were asked to report the time of their voluntary keypress using the position of a fast-rotating clock hand, a predictable action-effect (i.e. a 250 ms delayed sound after keypress) led to a shift of visuospatial attention towards the clock hand position of action-effect onset, thus demonstrating an influence of action-effects on action representation. Importantly, the attention shift occurred about 1 second before the action execution, which was further preceded and predicted by a lateralisation of alpha oscillations in the visual cortex. Our results indicate that when the spatial location is the key feature of action-effects, the neural implementation of the action-effect pre-activation is achieved through alpha lateralisation. |
Xiaojuan Xue; Gilles Pourtois Neurophysiological evidence for emotional attention modulation depending on goal relevance Journal Article In: Scientific Reports, vol. 15, no. 1, pp. 1–16, 2025. @article{Xue2025b,Although threat-related stimuli can capture attention automatically, recent findings have challenged this assumption by showing that goal rather than threat can be prioritized and eventually guide attentional control. In this study, we used high density electroencephalography (EEG) in 40 participants while peripheral emotional faces (either fear or happiness) were either goal-relevant or irrelevant during a dot-probe task (DPT). The use of peripheral vision was established by eye-tracking. Both the face specific N170 component and the subsequent Early Posterior Negativity (EPN) were enhanced by fear at the cue level, yet the latter one only when fear was goal relevant. Importantly, we found that early on following target onset at the P1 level, both value and goal relevance drove spatial attention and interacted with each other such that when they were goal-relevant, fearful faces captured attention less than when they were not. These results suggest that emotional attention is flexible and it can be influenced by the goal relevance of emotion. Moreover, they shed light on the electrophysiological manifestations of this flexibility and dovetail with the assumption that sensory gain control effects occurring in the visual cortex depending on attentional control are multiplexed and determined by both value and goal. |
Hao Zhang; Yiqing Hu; Yang Li; Shuangyu Zhang; Xiao Li Li; Chenguang Zhao Simultaneous dataset of brain, eye and hand during visuomotor tasks Journal Article In: Scientific Data, vol. 12, no. 1, pp. 1–15, 2025. @article{Zhang2025f,Visuomotor integration is a complex skill set encompassing many fundamental abilities, such as visual search, attention monitoring, and motor control. To explore the dynamic interplay between visual inputs and motor outputs, it is necessary to simultaneously record multiple brain activities with high temporal and spatial resolution, as well as to record implicit and explicit behaviors. However, there is a lack of public datasets that provide simultaneous multiple modalities during a visual-motor task. Functional near-infrared spectroscopy and electroencephalography to record brain activity simultaneously facilitate more precise capture of the complex visuomotor of brain mechanisms. Additionally, by employing a combined eye movement and manual response, it is possible to fully evaluate the effects of visuomotor outputs from implicit and explicit dimensions. We recorded whole-brain EEG (34 electrodes) and fNIRS (44 channels) covering the frontal and parietal cortex along with eye movements, behavior sampling, and operant behavior. The dataset underwent rigorous synchronization, quality control to highlight the effectiveness of our experiments and to demonstrate the high quality of our multimodal data framework. |
Panpan Chen; Chi Zhang; Bao Li; Li Tong; Long Cao; Ziya Yu; Bin Yan In: Cerebral Cortex, vol. 35, no. 11, pp. 1–11, 2025. @article{Chen2025i,Visual perceptual decoding of facial expressions is a key focus in affective neuroscience. Developing a mapping model between visual content and signals is crucial for decoding. Most previous visual decoding models focused on brain responses to static images, neglecting temporal-dynamic feature modeling. Additionally, they input all visual cortices as a whole into the model, overlooking that visual information flows bidirectionally between the lower and higher visual cortices based on bottom-up and top-down visual mechanisms, thus hard to capture bidirectional information between visual regions located in different spatial positions. Here, we present a spatio-temporal bidirectional long short-term memory-based model to decode 3 categories of facial expressions from multi-time functional magnetic resonance imaging data. Specifically, we used the spatio-temporal bidirectional long short-term memory module with the ability to simulate time series to grasp the temporal-dependence from visual cortices, and its forward and backward directions simulate bidirectional information flow between visual cortices to capture the bidirectional spatial information. Experimental outcomes indicate that the mean decoding accuracy employing beta estimates of multi-time response signals (Repetition TimesTR1-6 1 to 6) from 5 participants is significantly higher than that of other time points signals, unidirectional connections, and publicly available models. These results reveal that our model captures temporal-dependencies and bidirectional spatial information from the visual cortices, enhancing decoding performance. |
Matthew Lehet; Beier Yao; Ivy F. Tso; Vaibhav A. Diwadkar; Jessica Fattal; Jacqueline Bao; Katharine N. Thakkar Altered effective connectivity within a thalamocortical corollary discharge network in individuals with schizophrenia Journal Article In: Schizophrenia Bulletin, vol. 51, no. 6, pp. 1637–1650, 2025. @article{Lehet2025,Study Design: Here, we modeled fMRI data using dy- namic causal modeling (DCM) to examine patient-control differences in effective connectivity evoked by a double-step (DS) task (30 SZ, 29 HC). The interrogated network was formed from a combination of (1) functionally identified FEF and IPS regions that robustly responded on DS trials and (2) anatomically identified thalamic regions involved in CD transmission. We also examined the relationship between clinical symptoms and effective connectivity parameters associated with task modulation of network pathways. Study Results: Network connectivity was indeed modulated by the DS task, which involves CD transmission. More importantly, we found reduced effective connectivity from thalamus to IPS in SZ, which was further correlated with passivity symptom severity. Conclusions: These results reaffirm the importance of IPS and thalamocortical connections in oculomotor CD signaling and provide mechanistic insights into CD alterations and consequently agency disturbances in schizophrenia. |
Alexander J. Shackman; Jason F. Smith; Ryan D. Orth; Christina L. G. Savage; Paige R. Didier; Julie M. McCarthy; Melanie E. Bennett; Jack J. Blanchard Blunted ventral striatal reactivity to social reward is associated with more severe motivation and pleasure deficits in psychosis Journal Article In: Schizophrenia Bulletin, vol. 51, no. 6, pp. 1620–1636, 2025. @article{Shackman2025,Background and Hypothesis: Among individuals living with psychotic disorders, social impairment is common, debilitating, and challenging to treat. While the roots of this impairment are undoubtedly complex, converging lines of evidence suggest that social motivation and pleasure (MAP) deficits play a central role. Yet most neuroimaging studies have focused on monetary rewards, precluding decisive inferences. Study Design: Here we leveraged parallel social and monetary incentive delay functional magnetic resonance imaging paradigms to test whether blunted reactivity to social incentives in the ventral striatum - a key component of the distributed neural circuit mediating appetitive motivation and hedonic pleasure - is associated with more severe MAP symptoms in a transdiagnostic adult sample enriched for psychosis. To maximize ecological validity and translational relevance, we capitalized on naturalistic audiovisual clips of an established social partner expressing positive feedback. Study Results: Although both paradigms robustly engaged the ventral striatum, only reactivity to social incentives was associated with clinician-rated MAP deficits. This association remained significant when controlling for other symptoms, binary diagnostic status, or striatal reactivity to monetary incentives. Follow-up analyses suggested that this association predominantly reflects diminished activation during the presentation of social reward. Conclusions These observations provide a neurobiologically grounded framework for conceptualizing the social-anhedonia symptoms and social impairments that characterize many individuals living with psychotic disorders and underscore the need to develop targeted intervention strategies. |
Tahnée Engelen; Teresa Schuhmann; Alexander T. Sack; Catherine Tallon-Baudry Cardiac, respiratory, and gastric rhythms independently modulate motor corticospinal excitability in humans Journal Article In: PLoS Biology, vol. 23, no. 11, pp. 1–21, 2025. @article{Engelen2025,Interoception refers to the brain's sensing of internal body state and encompasses various bodily systems, notably the cardiac, respiratory, and gastric rhythms. Beyond their roles in physiological regulation and emotional states, each of these visceral rhythms has been shown to influence brain activity and cognition, prompting for the development of various interpretative functional frameworks. However, both experimental data and functional hypothesis leave it unclear whether and how each visceral rhythm acts simultaneously and independently on brain activity. Here, we address this question by measuring in human participants how the corticospinal excitability of the motor system varies with the phase of each of the three visceral rhythms. We applied single pulse transcranial magnetic stimulation (TMS) over the hand region in primary motor cortex to elicit Motor Evoked Potentials (MEPs), whose amplitude reflects corticospinal excitability, and tested whether MEP amplitude depends on the phase of the simultaneously measured cardiac, respiratory, and gastric rhythms. All three visceral rhythms were coupled to motor excitability with similar effect sizes at the group level. However, we found no relation between coupling strengths: participants displaying high coupling with one organ did not necessarily display high coupling to the other organs. These results indicate that independent mechanisms could underly the coupling between the cardiac, respiratory, and gastric rhythms and motor excitability. We further introduce the concept of individual interoceptive profiles and show that such interoceptive profiles obtained from objective coupling strength measures were not explained by self-reported awareness of the organ. Altogether, our results call for refined specifications of the frameworks offering a functional or clinical interpretation of viscera-brain coupling taking into account both independent mechanisms and individual interoceptive profiles. |
Allyson Copeland; Brennan R. Payne Co-registered eye-movements and brain potentials reveal multiple effects of context across the visual field in natural reading Journal Article In: Psychophysiology, vol. 62, no. 11, 2025. @article{Copeland2025,This study investigates how expectancy and plausibility influence behavioral and neural measures of language processing during naturalistic reading comprehension. Prior event-related potential (ERP) studies show evidence of distinct post-N400 positivities to violations of semantic expectancy and plausibility using artificial serial presentation but have yet to establish these phenomena during naturalistic reading. Therefore, we recorded simultaneous eye movements and EEG while participants read highly constraining sentences with expected, unexpected (but plausible), and anomalous target words. Time locked to the pre-target word, we observed a contextually graded parafoveal N400 effect. The N400 was facilitated (i.e., reduced) when the word was subsequently fixated, suggesting trans-saccadic integration of semantic features. At target fixation, we also observed a late anteriorly distributed positivity to unexpected target words and a posteriorly distributed positivity to anomalous target words, effects that were not clearly present when time locked to the pre-target word. Eye-tracking (ET) measures show that readers were sensitive to both expectancy and plausibility at target fixation. In conclusion, we show that readers can begin accessing semantic information in parafoveal vision, but higher-level semantic processing may require the orchestration of both parafoveal and foveal representations. |
Jiahong Cui; Nianqiu Shen; Hongjun Chen; Hongyu Zhou; Lei Hu; Wenbo Yu; Zhihan Liu; Daisuke Sawamura; Yuxuan Wang; Fengyu Cong Cross-modal semantic and non-semantic distraction impairs auditory working memory: Behavioral and ERP evidence Journal Article In: Biological Psychology, vol. 202, pp. 1–10, 2025. @article{Cui2025a,Visual distraction often disrupts auditory working memory, but it remains unclear whether semantic and non-semantic distractors interfere through similar or distinct neural mechanisms. This study used event-related potentials (ERPs) to examine the neural dynamics of semantic and non-semantic visual distractors during an auditory working memory task. Thirty nine healthy adults performed a paced auditory serial addition task (PASAT) while exposed to either semantic (digits) or non-semantic (abstract symbols) visual distractors. Behavioral results showed that both distractor types impaired accuracy, with semantic distractors producing longer reaction times and higher omission rates. ERP analyses revealed that both distractor types elicited enhanced P200 amplitudes and prolonged N200 and P300 latencies, reflecting shared early attentional capture and delayed stimulus evaluation. Semantic distractors further induced stronger N200 negativity at frontal sites and shortened P200 latencies, suggesting rapid semantic access and increased conflict detection demands, whereas only non-semantic distractors reduced P300 amplitudes, indicating reactive resource reallocation. Importantly, greater reductions in P300 amplitude were associated with slower responses in the semantic distractor condition. These findings demonstrate that semantic and non-semantic distractors engage partially overlapping but functionally distinct neural processes, emphasizing the importance of distractor content and processing stage in models of cross-modal cognitive control. |
Margaret Jane Moore; Amanda K. Robinson; Jason B. Mattingley Expectation dynamically modulates the representational time course of objects and locations Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–16, 2025. @article{Moore2025a,Past work has demonstrated that predictive information modulates how the brain responds to visual stimuli, but it is not yet clear how the brain integrates different types of predictive information to facilitate efficient perception. Here, we aim to explore how expectations about upcoming stimulus identities (“what” information) and upcoming stimulus locations (“where” information) modulate the directionality and occurrence of prediction effects in brain activity. Participants (n = 40) viewed real-world object images in rapid serial visual presentation (RSVP) streams which were predictable in terms of both object identity and stimulus location. Multivariate pattern analyses of electroencephalography (EEG) data were used to quantify and compare the degree of information represented in neural activity when stimuli were random (unpredictable), expected, or unexpected in terms of identity and location. Decoding accuracy for expected locations was significantly reduced relative to random locations between 160 and 238 ms post-onset. However, this effect subsequently reversed with decoding accuracy for expected locations becoming higher than accuracy for random locations between 273 and 430 ms. This temporally dynamic effect was not replicated within analyses decoding object identity. However, consistent evidence for reduced decoding of unexpected relative to random stimuli in later time windows (>250 ms) post-onset was identified across both stimulus types (e.g. objects and locations). These results are critically important when considered in the context of predictive coding research as they highlight important complexities in how predictability modulates neural responses. |
Matt Oxner; Dirk Moorselaar; Matthias M. Müller; Jan Theeuwes A flash in the pan? Distractor suppression cannot be inferred from the early lateralized positivity Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 11, pp. 2351–2369, 2025. @article{Oxner2025,Humans excel at avoiding distraction in visual environments, successfully filtering out repeated salient distractors that could otherwise capture attention. A recent theoretical perspective posits a mechanism whereby such distractors can be proactively suppressed, reducing their impact on attentional deployment. Electrophysiological evidence for this view comes from the distractor positivity (PD), a neural component associated with distractor handling. The PD has been observed at early latencies (< 200 msec) following distractor appearance, a timing interpreted as reflecting distractor suppression before attentional capture. However, the relationship between this “early PD” and distractor suppression remains fundamentally correlational. This raises critical questions about the extent to which this neural marker exclusively indexes mechanisms of suppression, as opposed to being driven by other factors confounded with distractor presence, such as stimulus salience. We tested the specificity of this early positivity to distractor handling across three experiments employing visual search tasks. Participants were presented with unique color singletons serving as distractors, targets, or task-irrelevant items. The early lateralized positivity was elicited by salient color distractors, but also appeared in response to all other salient singletons, including those that could not be proactively suppressed. Our findings indicate that the early positivity is not unique to suppressed distractors—instead, it likely reflects sensory imbalance between visual hemifields or salience tagging in response to lateralized stimuli. Consequently, we argue that the “early PD” does not provide definitive evidence for proactive distractor suppression, as its association with distractor presence appears to be incidental rather than causal. |
Paul Schmid; Christoph Reichert; Mandy V. Bartsch; Stefan Dürschmid Broadband high-frequency activity initializes distractor suppression Journal Article In: Cerebral Cortex, vol. 35, no. 11, pp. 1–14, 2025. @article{Schmid2025,Selective attention requires fast and accurate distractor suppression. We investigated if broadband high-frequency activity (BHA; 80–150 Hz), indicative of local neuronal population dynamics in early sensory cortices, indexes rapid processing of distracting information. In a first experiment we tested whether BHA distinguishes targets from distracting information in a visual search paradigm using tilted gratings as targets and distractors. In a second experiment, we examined whether BHA distractor processing can be trained by statistical regularities. In both experiments, BHA preceded the target enhancement (NT) and distractor suppression (PD; 1–40 Hz) event-related field (ERF) components and distinguished between targets and distractors. Only the BHA but not ERF component amplitude correlated with participants' performance and was higher for lateral distractors versus lateral targets. Furthermore, BHA predicted the strength of the PD. These results indicate that BHA initiates stimulus discrimination via distractor suppression. |
Paul Justin Connor Smith; Niko A. Busch Spontaneous alpha-band lateralization extends persistence of visual information in iconic memory by modulating cortical excitability Journal Article In: The Journal of Neuroscience, vol. 45, no. 48, pp. 1–10, 2025. @article{Smith2025c,Pre-stimulus alpha oscillations in the visual cortex modulate neuronal excitability, influencing sensory processing and decision-making. While this relationship has been demonstrated mostly in detection tasks with low visibility stimuli, interpretations of such effects can be ambiguous due to biases, making it difficult to clearly distinguish between perception-related and decision-related effects. In this study, we investigated how spontaneous fluctuations in pre-stimulus alpha power affect iconic memory, a high-capacity, ultra-short visual memory store. Data from 49 healthy adults (34 female and 15 male) was analyzed. We employed a partial report task, where a brief display of six stimuli was followed by a report cue indicating the target stimulus. In this paradigm, accuracy at short stimulus-cue onset asynchronies (SOAs) is typically high, reflecting the initial availability of sensory information, but it rapidly declines at intermediate SOAs due to the decay of the iconic memory trace, stabilizing at a low asymptote at long SOAs, representing the limited capacity of short-term memory. Crucially, performance in this task is constrained by the temporal persistence of sensory information, not by low visibility or response bias. We found that strong pre-stimulus alpha power enhanced performance by amplifying initial stimulus availability without affecting the speed of iconic decay. This effect partially reflects stronger pre-stimulus alpha power in the hemisphere ipsilateral to the to-be-reported target, likely suppressing neuronal excitability of neurons coding irrelevant stimuli. Our findings underscore the role of alpha oscillations in modulating neuronal excitability and visual perception, independent of decision-making strategies implicated in prior studies. |
Hyunwoo Gu; Joonwon Lee; Sungje Kim; Jaeseob Lim; Hyang Jung Lee; Heeseung Lee; Min Jin Choe; Dong Yoo; Jun Hwan (Joshua) Ryu; Sukbin Lim; Sang Hun Lee Attractor dynamics of working memory explain a concurrent evolution of stimulus-specific and decision-consistent biases in visual estimation Journal Article In: Neuron, vol. 113, no. 20, pp. 3476–3490, 2025. @article{Gu2025a,Sensory evidence tends to be fleeting, often unavailable when we categorize or estimate world features. To overcome this, our brains sustain sensory information in working memory (WM). Although keeping that information accurate while acting on it is vital, humans display two canonical biases: estimates are biased toward a few stimuli (“stimulus-specific bias”) and prior decisions (“decision-consistent bias”). Integrative—especially neural mechanistic—accounts of these biases remain scarce. Here, we identify drift dynamics toward discrete attractors as a common source of both biases in orientation estimation, with decisions further steering memory states. Behavior and neuroimaging data reveal how these biases co-evolve through the decision-steered attractor dynamics. Task-optimized recurrent neural networks suggest neural mechanisms that enable categorical decisions to emerge from WM for continuous stimuli while updating their trajectory, warping decision-consistent biases under stimulus-specific drift. |
Marc M. Himmelberg; Yuna Kwak; Marisa Carrasco; Jonathan Winawer Unpacking the V1 map: Differential covariation of preferred spatial frequency and cortical magnification across spatial dimensions Journal Article In: PLoS Computational Biology, vol. 21, no. 10, pp. 1–27, 2025. @article{Himmelberg2025,Primary visual cortex (V1) has long served as a model system for understanding cortical organization. Although its structural and functional properties vary markedly across its surface, patterns of covariation suggest possible underlying constancies. Such constancies would imply that V1 is composed of multiple identical units whose visual properties differ only due to differences in their inputs. To test this, we used fMRI to investigate how V1 cortical magnification and preferred spatial frequency covary with eccentricity and polar angle, measured in 40 observers. V1 cortical magnification and preferred spatial frequency were strongly correlated across eccentricity and around polar angle, however their relation differed between these dimensions: they were proportional across eccentricity but not polar angle. The constant ratio of cortical magnification to preferred spatial frequency when measured as a function of eccentricity suggests a shared underlying cause of variation in the two properties, e.g., the gradient of retinal ganglion cell density across eccentricity. In contrast, the deviation from proportionality around polar angle implies that cortical variation differs from that in retina along this dimension. Thus, a constancy hypothesis is supported for one of the two spatial dimensions of V1, highlighting the importance of examining the full 2D-map to understand how V1 is organized. |
Aakash Agrawal; Stanislas Dehaene From retinotopic to ordinal coding: Dissecting the cortical stages of visual word recognition Journal Article In: PNAS, vol. 122, no. 43, pp. 1–11, 2025. @article{Agrawal2025,Fluent reading requires the brain to precisely encode the positions of letters within words, distinguishing for instance FORM and FROM across variations in size, position, and font. Early visual areas, however, are known to encode retinotopic positions, and how these representations get transformed into a position-invariant neural code remains unclear. Building upon a computational model of reading, we used 7T functional MRI and magnetoencephalography (MEG) to reveal a cortical hierarchy in which early visual areas (V1–V4) predominantly encode retinotopic information, whereas higher-level regions, including the visual word form area, transition to an ordinal letter-position code. MEG analyses confirm that retinotopic encoding emerges early (60 to 200 ms), followed by a shift toward ordinal representations in later time windows (220 to 450 ms). Despite this transition, word position remained a dominant factor across all time points, suggesting a concurrent coding of both retinotopic and abstract positional information. These findings uncover the spatiotemporal dynamics by which the human brain transforms visual input into structured prelexical representations, shedding light on the cortical stages of reading and their developmental and clinical implications. |
Natasa Ganea; Richard N. Aslin; David J. Lewkowicz Covert attention modulates the SSVEP in a paradigm suitable for infants and young children Journal Article In: Attention, Perception, & Psychophysics, vol. 87, no. 7, pp. 2085–2104, 2025. @article{Ganea2025,Attention and visual gaze are usually tightly linked. Sometimes, however, we attend covertly to peripheral events without redirecting our gaze from the event that first attracted our overt attention. Despite evidence in adults that the steady-state visual evoked potential (SSVEP) varies with modulation of covert attention, paradigms used with adults are not suitable for use with infants and young children who cannot be instructed to perform tasks that dissociate overt from covert attention. Here, we provide evidence from a paradigm suitable for infants and young children that when gaze remains fixed on a central flickering visual stimulus while covert attention is directed briefly to the peripheral visual field, the SSVEP response undergoes significant attenuation. Signal-to-noise ratio (SNR) and intertrial coherence (ITC) measures of the SSVEP response to the central stimulus were lower when participants covertly deployed their attention to the peripheral stimulus than when central gaze and attention were aligned. Crucially, SNR was a more robust measure of attentional modulation than ITC, even though both measures were significantly correlated. Moreover, a 6 Hz flicker of the central stimulus resulted in a more reliable measure of attentional modulation than 12 Hz, and the inclusion of higher harmonics did not improve the reliability of either the SNR or the ITC measures. Our paradigm is unique in that it relies on short (2 s) response epochs, validates eye position during rapid shifts of covert attention, and makes it possible to obtain SSVEP measures of covert attention from infants, young children, and special populations. |
Cameron Smith; Daniel H. Baker Neural correlates of the deployment of spatial attention, and their modulation by repetitive movements Journal Article In: PLoS One, vol. 20, pp. 1–18, 2025. @article{Smith2025,The deployment of spatial attention generates distinct neural signatures that can be detected at the scalp. Here, we use multivariate pattern analysis of EEG data to decode the deployment of spatial attention, and ask if this is modulated by repetitive movements. ‘Stimming' movements (also known as repetitive stereotypies), are widely reported in autism, but also present in some neurotypical individuals. Stimming has historically been viewed as a problematic behaviour, but many individuals claim that stimming benefits attention. We first validated our paradigm (a Posner-style cueing design), demonstrating above-chance classification of cue direction from around 300 ms post-cue onset. We then investigated whether stimming modulates decoding accuracy and task performance. Our results, consisting of data primarily from neurotypical participants, do not suggest that stimming has a negative impact on an individual's ability to attend, unless the individual does not typically engage in stimming behaviours. This suggests interventions aiming to reduce stimming behaviours are not necessarily warranted and highlights the need for further research into the potential benefits of stimming specifically within the autistic population. Future research might also consider the potential overlap between autistic stimming and the fidgeting behaviours which are characteristic of ADHD, to help understand the significant overlaps between the characteristics of the two conditions. |
Dirk Moorselaar; Jan Theeuwes; Stefan Van der Stigchel Alpha-band activity tracks reflexive changes in the breadth of the zoom lens of attention Journal Article In: The Journal of Neuroscience, vol. 45, no. 44, pp. 1–11, 2025. @article{Moorselaar2025,Spatial attention is often conceptualized as a flexible "zoom lens" that can dynamically adjust its focus, but most evidence stems from studies of voluntary attention. Our study investigates whether involuntary, reflexive attention exhibits similar adaptability in attentional scope. Using behavioral and electroencephalographic (EEG) experiments with exogenous cues of varying spatial extent, we examined how attentional gradients dynamically adjust when attention is involuntarily captured. Male and female human participants performed visual search tasks preceded by narrow- or broad-cue displays at different onset asynchronies. We applied inverted encoding models to alpha-band neural activity to precisely track the locus and breadth of attentional tuning. Across experiments, we found that reflexive attentional gradients flexibly adapt to match cue characteristics. Behaviorally, narrow cues yielded progressively sharper attentional gradients compared with broad cues, with differences emerging over time. Critically, EEG analyses revealed that alpha-band activity tracked these dynamic adjustments, with differences in spatial selectivity emerging rapidly (±200 ms postcue) and continuing to evolve. Contrary to previous suggestions that involuntary attention primarily influences response efficiency, our results demonstrate that exogenous cues modulate attentional resources across the visual field at early processing stages. |
Sumiya Sheikh Abdirashid; Tomas Knapen; Serge O. Dumoulin The precision of attention controls attraction of population receptive fields Journal Article In: Journal of Vision, vol. 25, no. 11, pp. 1–15, 2025. @article{Abdirashid2025,We alter our sampling of visual space not only by where we direct our gaze, but also by where and how we direct our attention. Attention attracts receptive fields toward the attended position, but our understanding of this process is limited. Here we show that the degree of this attraction toward the attended locus is dictated not just by the attended position, but also by the precision of attention. We manipulated attentional precision while using 7T functional magnetic resonance imaging to measure population receptive field (pRF) properties.Participants performed the same color-proportion detection task either focused at fixation (0.1° radius) or distributed across the entire display (>5° radius). We observed blood oxygenation level-dependent response amplitude increases as a function of the task, with selective increases in foveal pRFs for the focused attention task and vice versa for the distributed attention task. Furthermore, cortical spatial tuning changed as a function of attentional precision. Specifically, focused attention more strongly attracted pRFs toward the attended locus compared with distributed attention. This attraction also depended on the degree of overlap between a pRF and the attention field. A Gaussian attention field model with an offset on the attention field explained our results. Together, our observations indicate the spatial distribution of attention dictates the degree of its resampling of visual space. |
Ecem Altan; Catherine A. Morgan; Steven C. Dakin; D. Samuel Schwarzkopf Spatial frequency adaptation modulates population receptive field sizes Journal Article In: eLife, vol. 13, pp. 1–29, 2025. @article{Altan2025,The spatial tuning of neuronal populations in the early visual cortical regions is related to the spatial frequency (SF) selectivity of neurons. However, there has been no direct investigation into how this relationship is reflected in population receptive field (pRF) sizes despite the common application of pRF mapping in visual neuroscience. We hypothesised that adaptation to high/low SF would decrease the sensitivity of neurons with respectively small/large receptive field sizes, resulting in a change in pRF sizes as measured by functional magnetic resonance imaging (fMRI). To test this hypothesis, we first quantified the SF aftereffect using a psychophysical paradigm where human observers made SF judgments following adaptation to high/low SF noise patterns. We then incorporated the same adaptation technique into a standard pRF mapping procedure to investigate the spatial tuning of the early visual cortex following SF adaptation. Results showed that adaptation to a low/high SF resulted in smaller/larger pRFs, respectively, as hypothesised. Our results provide the most direct evidence to date that the spatial tuning of the visual cortex, as measured by pRF mapping, is related to the SF selectivity of visual neural populations. This has implications for various domains of visual processing, including size perception and visual acuity. |
Ilona M. Bloem; Leah Bakst; Joseph T. McGuire; Sam Ling Dynamic estimation of the attentional field from visual cortical activity Journal Article In: eLife, vol. 14, pp. 1–21, 2025. @article{Bloem2025,Navigating around the world, we must adaptively allocate attention to our surroundings based on anticipated future stimuli and events. This allocation of spatial attention boosts visuocortical representations at attended locations and locally enhances perception. Indeed, spatial attention has often been analogized to a ‘spotlight' shining on the item of relevance. Although the neural underpinnings of the locus of this attentional spotlight have been relatively well studied, less is known about the size of the spotlight: to what extent can the attentional field be broadened and narrowed in accordance with behavioral demands? In this study, we developed a paradigm for dynamically estimating the locus and spread of covert spatial attention, inferred from visuocortical activity using fMRI in humans. We measured BOLD activity in response to an annulus while participants (four female, four male) used covert visual attention to determine whether more numbers or letters were present in a cued region of the annulus. Importantly, the width of the cued area was systematically varied, calling for different sizes of the attentional spotlight. The deployment of attention was associated with an increase in BOLD activity in corresponding retinotopic regions of visual areas V1–V3. By modeling the visuocortical attentional modulation, we could reliably recover the cued location, as well as a broadening of the attentional modulation with wider attentional cues. This modeling approach offers a useful window into the dynamics of attention and spatial uncertainty. |
Laura Doll; Sabine Heiland; Alexander Gutschalk A role of pupil-linked arousal, cingulo-insular cortex, and intralaminar thalamus for auditory near-threshold perception Journal Article In: Journal of Cognitive Neuroscience, vol. 37, no. 9, pp. 1391–1415, 2025. @article{Doll2025,The perception of near-threshold tones varies strongly across trials, likely because of fluctuations in sustained attention or arousal. We used parallel fMRI and pupillometry to study the role of attention networks for the detection of near-threshold tones in three phases: (1) passive listening, (2) active detection of salient tones, and (3) active detection of near-threshold tones. Results confirmed previous findings from magnetoencephalography that auditory cortex activity and pupil-dilation responses for near-threshold tones were only observed when task-relevant, stronger for hit trials, but also present for miss trials. We then sought which attention-related areas show a similar response pattern, and found it in insular cortex, anterior midcingulate cortex, and inferior precentral sulcus. Moreover, activity in the insula was already stronger for hit than miss trials in the prestimulus interval. Activity for hit trials was also observed in a number of subcortical nuclei, including thalamus, periaqueductal gray, locus coeruleus, and the colliculi. Like insula, activity in the intralaminar nuclei of the thalamus additionally showed activity for miss trials and stronger activity for hit trials in the baseline. Finally, BOLD activity correlated to spontaneous pupil fluctuations was evaluated and revealed biphasic activation and deactivation in a widespread cortical network, with a maximum 3 sec and minimum 7 sec after pupil dilation. The cortical networks included insula, anterior midcingulate cortex, retro-splenial, and sensory cortex. Overall, these data identify the cingulo-insular network and the intralaminar thalamic nuclei as potential sources of fluctuations in auditory cortex activity in the context of near-threshold tone detection. |
Natalia Ladyka-Wojcik; Zhong Xu Liu; Jennifer D. Ryan In: Imaging Neuroscience, vol. 3, pp. 1–18, 2025. @article{LadykaWojcik2025,Memory and gaze behavior are intricately linked, guiding one another to extract information and create mental representations of our environment for subsequent retrieval. Recent findings from functional neuroimaging and computational modeling suggest that reciprocal interactions between the extended hippocampal system and visuo-oculomotor regions are functionally relevant for building these mental representations during visual exploration. Yet, evidence for the directionality of information flow during encoding within this reciprocal architecture in humans is limited. In the current study, we used dynamic causal modeling (DCM) to give a non-invasive account for the directional influences between these systems when new memories are created. Here, we provide novel evidence demonstrating how unrestricted, naturalistic visual exploration induces changes in this connectivity. Subsequent memory retrieval performance was also predicted by the pattern of connectivity modulated by unrestricted visual exploration, identifying the mechanism underlying a rich history of previous work linking increased gaze behavior during encoding to later memory. Together, these findings suggest that gaze behavior shapes the ways in which brain dynamics within and between the hippocampal system and early visual cortex unfold during encoding in humans. Importantly, these directional interactions support the building of coherent, lasting mental representations. |
Vivian C. Paulun; Raghavendra Rao T. Pramod; Joshua B. Tenenbaum; Nancy Kanwisher Dissociable cortical regions represent things and stuff in the human brain Journal Article In: Current Biology, vol. 35, pp. 1–9, 2025. @article{Paulun2025,Extensive prior work has identified regions of the human brain associated with visual perception of objects (lateral occipital complex [LOC]) and their physical properties and interactions (“frontoparietal physics network” [FPN]). However, this work has nearly exclusively tested the response of these regions to rigid objects. Deformable or nonsolid substances, or “stuff,” including liquids such as water or honey and granular materials such as sand or snow, are of similar importance in everyday life but have different physical properties and invite different actions. Little is known about the brain basis of stuff perception. Here, we scan participants with functional MRI (fMRI) while they view videos of rigid and non-rigid objects (“things”) and liquid and granular substances (stuff). We find double dissociations between the processing of things and stuff within both the ventral and dorsal visual pathways. These findings suggest that distinct mental algorithms are engaged when we perceive things and stuff, as they are in artificial physics engines. |
Addison D. N. Billing; Eleanor S. Smith; Robert J. Cooper; Rebecca P. Lawson Maternal anxiety shapes prediction error responses in the infant brain Journal Article In: Neurophotonics, vol. 12, no. 03, pp. 1–16, 2025. @article{Billing2025,SIGNIFICANCE: Postnatal maternal anxiety affects a substantial number of new mothers and is linked to long-term risk for anxiety in their offspring. Yet, the neural mechanisms through which postnatal maternal anxiety influences early cognitive development remain unclear. We investigated whether postnatal maternal anxiety shapes how infant brains respond to unexpected events-prediction errors-which are central to learning in uncertain environments. AIM: We examined prediction error processing in 6- to 8-month-old infants using high-density diffuse optical tomography and eye-tracking. We hypothesized that neural responses in the medial prefrontal cortex (mPFC) would vary with maternal anxiety levels. APPROACH: Infants viewed audiovisual events where expected outcomes were occasionally omitted, eliciting prediction errors. Hemodynamic responses in the frontal cortex were analyzed using a general linear model, with trial-by-trial gaze data as a parametric modulator. Maternal anxiety was measured using the state-trait anxiety inventory. RESULTS: Prediction error responses were localized to the mPFC and were only detectable when controlling for infant attention using eye-tracking. Cortical activation in response to unexpected stimuli was significantly enhanced in infants of mothers with higher trait anxiety. CONCLUSION: Our findings suggest that maternal anxiety modulates prediction error processing in the infant brain, potentially shaping early sensitivity to environmental unpredictability and conferring risk for later anxiety. |
Debadatta Dash; Fumiaki Iwane; William Hayward; Roberto F Salamanca-Giron; Marlene Bönstrup; Ethan R Buch; Leonardo G Cohen Sequence action representations contextualize during early skill learning Journal Article In: eLife, vol. 13, pp. 1–25, 2025. @article{Dash2025,Activities of daily living rely on our ability to acquire new motor skills composed of precise action sequences. Here, we asked in humans if the millisecond-level neural representation of an action performed at different contextual sequence locations within a skill differentiates or remains stable during early motor learning. We first optimized machine learning decoders predictive of sequence-embedded finger movements from magnetoencephalographic (MEG) activity. Using this approach, we found that the neural representation of the same action performed in different contextual sequence locations progressively differentiated—primarily during rest intervals of early learning (offline)—correlating with skill gains. In contrast, representational differentiation during practice (online) did not reflect learning. The regions contributing to this representational differentiation evolved with learning, shifting from the contralateral pre- and post-central cortex during early learning (trials 1–11) to increased involvement of the superior and middle frontal cortex once skill performance plateaued (trials 12–36). Thus, the neural substrates supporting finger movements and their representational differentiation during early skill learning differ from those supporting stable performance during the subsequent skill plateau period. Representational contextualization extended to Day 2, exhibiting specificity for the practiced skill sequence. Altogether, our findings indicate that sequence action representations in the human brain contextually differentiate during early skill learning, an issue relevant to brain-computer interface applications in neurorehabilitation. |
Jan Willem Gee; Niels A. Kloosterman; Anke Braun; Tobias H. Donner Catecholamines reduce choice history biases in perceptual decision making Journal Article In: PLoS Biology, vol. 23, pp. 1–15, 2025. @article{Gee2025,Theoretical accounts postulate that the catecholaminergic neuromodulator noradrenaline shapes cognition and behavior by reducing the impact of prior expectations on learning, inference, and decision-making. A ubiquitous effect of dynamic priors on perceptual decisions under uncertainty is choice history bias: the tendency to systematically repeat, or alternate, previous choices, even when stimulus categories are presented in a random sequence. Here, we directly test for a causal impact of catecholamines on these priors. We pharmacologically elevated catecholamine levels in human participants through the application of the noradrenaline reuptake inhibitor atomoxetine. We quantified the resulting changes in observers' history biases in a visual perceptual decision task. Choice history biases in this task were highly idiosyncratic, tending toward choice repetition or alternation in different individuals. Atomoxetine decreased these biases (toward either repetition or alternation) compared to placebo. Behavioral modeling indicates that this bias reduction was due to a reduced bias in the accumulation of sensory evidence, rather than of the starting point of the accumulation process. Atomoxetine had no significant effect on other behavioral measures tested, including response time and choice accuracy. Atomoxetine and variations of pupil-linked arousal at slower and faster timescales had analogous effects on choice history bias. We conclude that catecholamines reduce the impact of a specific form of prior on perceptual decisions. |
Ziyue Hu; Dominic M. D. Tran; Reuben Rideaux Multimodal evidence challenges the effectiveness of probabilistic cueing for establishing sensory expectations Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–19, 2025. @article{Hu2025b,Predictive coding theories posit a reduction in error-signaling neural activity when incoming sensory input matches existing expectations—a phenomenon termed expectation suppression. However, the empirical evidence for expectation suppression, as well as its underlying neural mechanism, is contentious. A further aspect of predictive coding that remains untested is how predictions are integrated across sensorimotor domains. To investigate these two questions, we employed a novel cross-domain probabilistic cueing paradigm, where participants were presented with both visual and motor cues within a single trial. These cues manipulated the orientation and temporal expectancy of target stimuli with 75% validity. Participants completed a reproduction task where they rotated a bar to match the orientation of the target stimulus while their neural and pupil responses were respectively measured via electroencephalography and eye tracking. Our results showed a consistent, feature-unspecific effect of motor expectancy across multiple measures, while evidence for visual expectancy was limited. However, neither motor nor visual expectancy modulated the fidelity of sensory representations. These results indicate that violations of temporal expectancy in the current study may reveal the brain's intrinsic sensitivity to temporal regularities in the natural settings, rather than feature-specific predictions. In contrast, the absence of visual expectancy effects in both neural and pupillometry results adds to a growing body of evidence questioning the effectiveness of probabilistic cueing paradigms for establishing expectations capable of altering sensory representations. Due to null findings in the visual and sensory representation analyses, we did not further investigate cross-domain prediction integration. |
Sandra Klonteig; Elise S. Roalsø; Brage Kraft; Torgeir Moberget; Eva Hilland; Peyman Mirtaheri; Rune Jonassen In: Journal of Behavior Therapy and Experimental Psychiatry, vol. 88, pp. 1–12, 2025. @article{Klonteig2025,Background: Attentional bias (AB) is characterized by preferential cognitive and emotional processing of mood-congruent stimuli and considered a central mechanism in mood disorders. Considerable research has focused on improving AB measures to enhance mechanistic understanding and clinical utility. The present study examines psychometric properties of a range of AB measures with a multimodal setup. Methods: A nonclinical sample of 62 women aged 20–30 years completed the facial dot-probe task while behavioral responses (reaction time), eye-gaze patterns (eye tracking), and electrical brain potentials (electroencephalography) were recorded. AB metrics from four types of AB measures – traditional, response-based, dwell time, and the N2pc component– were examined with internal consistency and short-term test-retest calculations. AB metrics with an internal consistency score over .4 were considered reliable, and their validity were explored by examining relations to depression and anxiety symptoms. In addition, the consistency between reliable metrics across trials were examined. Results: Findings show that traditional AB metrics exhibited no degree of reliability, whereas response-based and dwell time metrics overall demonstrated better internal consistencies. Response-based metrics also had higher test-retest reliability in all but one metric. The previously reported reliability of the N2pc component was not observed. As for validity, no linear associations were found between the reliable measures, depression, and anxiety. There were no relations between metrics across trials. Conclusions: This study provides insights for future AB research, emphasizing the potential of novel metrics over traditional ones and the use of multimodal setups to develop reliable and potentially hybrid measurements for clinical assessment. |
Baiwei Liu; Siyang Kong; Freek Ede Microsaccades strongly modulate but do not directly cause the EEG N2pc marker of spatial attention Journal Article In: PLoS biology, vol. 23, no. 9, pp. 1–17, 2025. @article{Liu2025c,The N2pc is a popular human-neuroscience marker of covert and internal spatial attention that occurs 200-300 ms after being prompted to shift attention-a time window also characterized by the spatial biasing of small fixational eye movements known as microsaccades. Here, we show how co-occurring microsaccades profoundly modulate N2pc amplitude during top-down shifts of spatial attention in both perception and working memory. At the same time, we show that a significant-albeit severely weakened-N2pc can still be established in the absence of co-occurring microsaccades. Moreover, despite the strong modulation of the N2pc by microsaccade presence and direction, the N2pc does not align to the precise timing of microsaccades, ruling out that the observed N2pc modulations by microsaccades are a direct artifact of microsaccade-related eye-muscle activity, corneo-retinal dipole movement, or visual inputs moving over the retina. Thus, while microsaccades strongly modulate N2pc amplitude, microsaccades themselves are not a prerequisite for, nor a direct cause of, the N2pc. |
Sreenivasan Meyyappan; Mingzhou Ding; George R. Mangun Hierarchical organization of human visual feature attention control Journal Article In: The Journal of Neuroscience, vol. 45, no. 39, pp. 1–13, 2025. @article{Meyyappan2025,Attention can be deployed in advance of visual stimuli based on features such as color or direction of motion. This anticipatory feature-based attention involves top-down neural control signals from the frontoparietal network that bias visual cortex to enhance attended information and suppress distraction. For example, anticipatory attention control can enable effective selection based on stimulus color while ignoring distracting information about stimulus motion. Anticipatory attention can also be focused more narrowly, for example, to select specific colors or motion directions that define task-relevant aspects of the stimuli. One important question that remains open is whether anticipatory attention control first biases broad feature dimensions such as color versus motion before biasing the specific feature attributes (e.g., blue vs green). To investigate this, we recorded EEG activity during a task where human participants of either sex were cued to either attend to a motion direction (up or down) or a color (blue or green) on a trial-by-trial basis. Applying multivariate decoding approaches to the EEG alpha band activity (8-12 Hz) during attention control (cue-target interval), we observed significant decoding for both the attended dimensions (motion vs color) and specific feature attributes (up vs down; blue vs green). Importantly, the temporal onset of the dimension-level biasing (motion vs color) preceded that of the attribute-level biasing (up vs down and blue vs green). These findings demonstrate that the top-down control of feature-based attention proceeds in a hierarchical fashion, first biasing the broad feature dimension, and then narrowing to the specific feature attribute. |
Matthias Mittner; Josephine Maria Groot A novel method for modeling tonic and phasic pupil dynamics in humans Journal Article In: Behavior Research Methods, vol. 57, no. 9, pp. 1–13, 2025. @article{Mittner2025,The human pupil is a widely used physiological metric in psychology and neuroscience. Changes in pupil diameter (PD) are thought to reflect changes in locus coeruleus-norepinephrine (LC/NE) activity, which is associated with cognitive and behavioral optimization. Here, we present a novel algorithm to decompose the pupil signal into its tonic and phasic components. We evaluate the utility and validity of the algorithms using both artificially generated data and an existing dataset from a fast-paced finger-tapping task. Results show that the novel algorithm outperforms traditional approaches on simulated data. We further demonstrate that our algorithm provides more conclusive evidence for relationships between mind wandering reports and pupil predictors compared to traditional window-averaging. Finally, we demonstrate that the novel and traditional estimates contain distinct information regarding neuroimaging correlates and task performance. |
Brian Nestor; Ayah Elaboudi; Sara Milligan; Elizabeth R. Schotter Parafoveally perceived orthographic cues facilitate foveal semantic processing: Evidence from event-related potentials Journal Article In: Brain and Language, vol. 268, pp. 1–10, 2025. @article{Nestor2025,Readers extract information from words viewed parafoveally, but it is unclear whether this processing is limited to orthography or if it extends to lexico-semantic content. In the current ERP study, we measured the N400 responses to words that were perceived parafoveally and/or foveally using the RSVP-with-flankers paradigm and a parafoveal masking manipulation. We compared anomalous orthographically related (neighbor) and unrelated (non-neighbor) words to expected words to determine whether the N400 responses were driven by orthographic and/or semantic processing. We observed a large parafoveal N400 effect in response to the non-neighbors (versus expected), and a smaller, later parafoveal N400 for neighbors, suggesting that the parafoveal response is largely orthographic in nature. We also observed a significant reduction in foveal N400 magnitude when non-neighbor words were previously visible parafoveally (but not for the foveal N400 response to neighbors), suggesting that facilitation of foveal processing is driven by parafoveal detection of orthographic violations. |
Richard Schweitzer; Thomas Seel; Jörg Raisch; Martin Rolfs Early visual signatures and benefits of intra-saccadic motion streaks Journal Article In: PLoS Computational Biology, vol. 21, no. 9, pp. 1–37, 2025. @article{Schweitzer2025a,Eye movements routinely induce motion streaks as they shift visual projections across the retina at high speeds. To investigate the visual consequences of intra-saccadic motion streaks, we co-registered eye tracking and EEG while gaze-contingently shifting target objects during saccades, presenting either continuous, 'streaky' or apparent, step-like motion in four directions. We found significant reductions of secondary saccade latency, as well as improved decoding of the post-saccadic target location from the EEG signal when motion streaks were available. These signals arose as early as 50 ms after saccade offset and had a clear occipital topography. Using a physiologically plausible visual processing model, we provide evidence that the target's motion trajectory is coded in orientation-selective channels and that speed of gaze correction was linked to the visual dynamics arising from the combination of saccadic and target motion, providing a parsimonious explanation of the behavioral benefits of intra-saccadic motion streaks. |
David Clewett; Ringo Huang; Lila Davachi Locus coeruleus activation “resets” hippocampal event representations and separates adjacent memories Journal Article In: Neuron, vol. 113, no. 15, pp. 2521–2535.e1–e8, 2025. @article{Clewett2025,Memories reflect the ebb and flow of experiences, capturing distinct events from our lives. Using a combination of functional magnetic resonance imaging (fMRI), neuromelanin imaging, and pupillometry, we show that arousal and locus coeruleus (LC) activation segment continuous experiences into discrete memories. As sequences unfold, encountering a context shift or event boundary triggers pupil-linked arousal and LC processes that predict later memory separation. Boundaries, furthermore, promote temporal pattern separation within the left hippocampal dentate gyrus, which correlates with heightened LC responses to those same transition points. Unlike transient LC effects, indirect structural and functional markers of elevated background LC activation correlate with reduced arousal-related LC and pupil responses at boundaries, suggesting that hyperarousal disrupts event segmentation. Our findings support the idea that arousal mechanisms initiate a neural and memory “reset” in response to significant changes, fundamentally shaping the episodes that define episodic memory. |
Marco Ninghetto; Georgios A. Keliris; Kamil Szulborski; Tomasz Gałecki; Bartosz Kossowski; Daan Panneman; Frans P. M. Cremers; Monika Ołdak; Jacek P. Szaflik; Kalina Burnat Cortical response to transient and long-term visual field loss Journal Article In: Cerebral Cortex, vol. 35, no. 8, pp. 1–13, 2025. @article{Ninghetto2025,In the visual cortices, receptive fields (RFs) are arranged in a gradient from small sizes in the center of the visual field to the largest sizes at the periphery. Using functional magnetic resonance imaging (fMRI) mapping of population RFs, we investigated RF adaptation in V1, V2, and V3 in patients after long-term photoreceptor degeneration affecting the central (Stargardt disease [STGD]) and peripheral (Retinitis Pigmentosa [RP]) regions of the retina. In controls, we temporarily limited the visual field to the central 10° to model peripheral loss. The central loss experienced by STGD patients led to an increase in RF size in the dorsal subdivisions of V1, V2, and V3. In contrast, peripheral loss in RP patients led to a bilateral increase in population RF sizes in V1 but a decrease in V2. Transient peripheral loss in controls led to an increase in RF size in V1 and a decrease in V2 and V3, regardless of the dorsal-ventral division of the cortical representation. Our findings suggest a dorsal-ventral difference in RF size in response to central visual field loss, likely reflecting the functional relevance of these divisions within the cortical representations of the visual field. |
Wendy Sun; Anne Billot; Jingnan Du; Xiangyu Wei; Rachel A. Lemley; Mohammad Daneshzand; Aapo Nummenmaa; Randy L. Buckner; Mark C. Eldaief Precision network modeling of transcranial magnetic stimulation across individuals suggests therapeutic targets and potential for improvement Journal Article In: Human Brain Mapping, vol. 46, no. 11, pp. 1–23, 2025. @article{Sun2025,Higher-order cognitive and affective functions are supported by large-scale networks in the brain. Dysfunction in different networks is proposed to associate with distinct symptoms in neuropsychiatric disorders. However, the specific networks targeted by current clinical transcranial magnetic stimulation (TMS) approaches are unclear. While standard-of-care TMS relies on scalp-based landmarks, recent FDA-approved TMS protocols use individualized functional connectivity with the subgenual anterior cingulate cortex (sgACC) to optimize TMS targeting. Leveraging previous work on precision network estimation and modeling of the TMS electric field (E-field), we asked whether various clinical TMS approaches target different functional networks between individuals. Results revealed that modeled homotopic scalp positions (left F3 and right F4) target different networks within and across individuals, and right F4 generally favors a right-lateralized control network. TMS coil positions over the dorsolateral prefrontal cortex (dlPFC) zone anticorrelated with the sgACC most frequently target a network coupled to the ventral striatum (reward circuitry) but largely miss that network in some individuals. We further illustrate how modeling can be used to retrospectively assess the estimated targets achieved in prior TMS sessions and also used to prospectively provide coil positions that can target distinct closely localized dlPFC network regions with spatial selectivity and maximal E-field intensity. In a final study, precision targeting was found to be feasible in participants with Major Depressive Disorder using data derived from a single low-burden MRI session suggesting the methods are applicable to translational efforts where limiting patient burden and ensuring robustness are critical. |
Zhaohuan Ding; Wenbo Ma; Leixiao Feng; Mingsha Zhang; Xiaoli Li Quantifying task-locked information transmission between cortical areas with TMS-EEG Journal Article In: NeuroImage, vol. 317, pp. 1–10, 2025. @article{Ding2025c,Objective: This study aims to develop TMS-EEG (Transcranial magnetic stimulation combined with EEG) technology to detect task-locked neural network activation and dynamically quantify information transmission. Approach: 30 participants performed visually guided gap saccade tasks while TMS-EEG data were recorded, with the TMS pulses delivered to prefrontal cortex (PFC) and posterior parietal cortex (PPC) at different task stages. The directed transfer function (DTF) method was applied to TMS-EEG data to indicate the information flow. By analyzing the channel combinations associated with the PFC and PPC, we calculated differences in information flow within the alpha, beta, and gamma frequency bands to determine whether TMS-EEG could quantitatively characterize the direction of information flow between cortical areas. Main results: Analysis of eye tracker data revealed that all participants successfully performed the saccade task, with a correct rate exceeding 90 %. The mean saccade latency was 132.25 ± 22.59 ms after target appearance. Stimulation of the PFC and PPC revealed significant differences in information flow in the gamma bands at different time points. Specifically, during the preparatory period, the C3 electrode acts as a hub for incoming information from O1, later transitioning to send information towards F4 and O1 post-target. Then, P3 emerges as a hub, sending data towards P4, with connectivity between them intensifying post 100 ms from the target's appearance. Significance: This study utilized DTF values derived from TMS-EEG to characterize information flow between cortical areas during the gap saccade task. This approach provides a novel method for quantifying dynamic changes in connectivity and causality between cortical areas during task processing. |
Wenbo Ma; Zhaohuan Ding; Leixiao Feng; Xiaoli Li; Mingsha Zhang The role of prefrontal and posterior parietal cortex in generating multiple step saccades Journal Article In: Neuroscience Bulletin, vol. 41, no. 8, pp. 1418–1428, 2025. @article{Ma2025,While multiple step saccades (MSS) are occasionally reported in the healthy population, they are more evident in patients with Parkinson's disease (PD). Therefore, MSS has been suggested as a biological marker for the diagnosis of PD. However, the lack of clarity on the neural mechanism underlying the generation of MSS largely impedes their application in the clinic. We have proposed recently that MSS are triggered by the discrepancy between desired and executed saccades. Accordingly, brain regions involved in saccadic planning and execution might play a role in the generation of MSS. To test this hypothesis, we explored the role of the prefrontal (PFC) and posterior parietal cortex (PPC) in generating MSS by conducting two experiments: electroencephalographic recording and single-pulse transcranial magnetic stimulation in the PFC or PPC of humans while participants were performing a gap saccade task. We found that the PFC and PPC are involved in the generation of MSS. |
Sajjad Karimi; Masoud Nateghi; Gabriela I. Cestero; Lina Chitadze; Deepanshi; Yi Yang; Juhee H. Vyas; Chuoqi Chen; Zeineb Bouzid; Cem O. Yaldiz; Nicholas Harris; Rachel Bull; Bradly T. Stone; Spencer K. Lynn; Bethany K. Bracken; Omer T. Inan; J. Douglas Bremner; Reza Sameni Prescreening depression using wearable electrocardiogram and photoplethysmogram data from a psycholinguistic experiment Journal Article In: Physiological Measurement, vol. 46, no. 8, pp. 1–20, 2025. @article{Karimi2025,Objective. Depression is a prevalent mental health disorder that significantly impacts well-being and quality of life. This study investigates the relationship between depression and cardiovascular function, exploring time-series features derived from electrocardiogram (ECG) and photoplethysmogram (PPG) data as potential biomarkers for depression prescreening. Approach. As part of a comprehensive psycholinguistic experiment, we collected data from 60 individuals, including both healthy participants and those with varying levels of depression, assessed using the Beck Depression Inventory-II (BDI-II) and the Patient Health Questionnaire-9 (PHQ-9). Bimodal features derived from both ECG and PPG data were used to develop machine learning models for depression risk classification, employing classifiers such as random forest, XGBoost, logistic regression, and support vector machines (SVMs). Additionally, regression models were built to predict depression severity based on ECG- and PPG-derived biomarkers. Main results. Key findings indicate that short-term variability (SD1) features in the ECG RR interval, peripheral systolic and diastolic phases from the PPG, and pulse duration significantly differ between healthy individuals and those at risk of depression. SVM achieved the best classification performance, with an area under the ROC curve of 0.83 ± 0.11 for BDI-II-based classification and 0.78 ± 0.11 for PHQ-9-based classification. SHapley Additive exPlanations analysis consistently identified systolic-SD1 and RR-SD1 as key predictors. Regression analysis further supported the role of cardiovascular features in assessing depression severity, yielding a mean absolute error of 10.18 for BDI-II and 5.27 for PHQ-9 score regression. Significance. This study demonstrates the feasibility of using wearable ECG and PPG technologies for depression prescreening. The findings suggest that cardiac activity-based biomarkers can contribute to the development of cost-effective, objective, and non-invasive tools for mental health assessment, complementing traditional diagnostic methods. |
Longxia Lou; Ximei Feng; Zehao Liang; Zhi-fang Liu; Zhijun Zhang Contextual plausibility effects among older adults in Chinese free-view reading: Co-registration of eye-tracking and electroencephalography Journal Article In: Perceptual and Motor Skills, pp. 1–22, 2025. @article{Lou2025,With regard to how aging affects contextual plausibility in Chinese natural silent reading, research remains limited. To address the methodological limitations inherent in using eye-tracking measures and event-related potentials separately, we aimed to apply co-registration of eye-tracking with electroencephalography (EEG) in Chinese natural silent reading. Old and young adults were asked to read sentences that contained either semantically congruent or semantically violated words. We failed to replicate any differences in contextual plausibility effects between the older and younger adults on eye-tracking measures of early word processing (including first fixation duration, gaze duration, probability of skipping, and refixation). However, larger plausibility effects for the old adult readers were observed in the measures of regression in probability and total reading time. No reliable age-related differences in plausibility effects were observed for brain response amplitudes in the parafoveal and foveal intervals (from −400 to 200 ms). Both eye-tracking and EEG data demonstrated that contextual plausibility in early word processing was preserved among the old adults, while the Chinese old adult readers made more efforts to reanalyze semantic information in relation to free-view reading. |
Jose Mora-Quiroga; Juan Pablo Abril-Ronderos; Marisol R. Lamprea Reciprocal competition between cognitive tasks and emotional processing revealed by EEG and eye tracking Journal Article In: International Journal of Psychophysiology, vol. 214, pp. 1–10, 2025. @article{MoraQuiroga2025,Processing of engaging stimuli triggers increases in visual exploration, enhances the centroparietal late positive potential (LPP) and produce larger posterior alpha desynchronization (a-ERD), suggesting orienting and attention allocation. It has been shown that a salient arousing image in the background of a cognitive task consumes processing resources from the superimposed task, producing a deleterious effect on performance. On the other hand, experiments designed to change the focus of attention during the processing of emotional stimuli have shown a reduction of the emotional response, indicating an attentional competition between significant stimuli simultaneously presented. This research aimed to describe the competition between the performance on a cognitive task presented in a reduced space of the image (1.2 %) and the processing of emotional images displayed at the background using EEG and Eye Tracking. Results showed that the superimposition of the task had an early attractor effect, evidenced by an above-chance decoding accuracy (about 180 ms since the image onset) and an enhancement in the N1 component. This engagement in the task reduced the processing of the images as evidenced by a decrease in the LPP amplitudes, an enhancement in the a-ERD and a greater dwell-time over the task. Additional analysis showed that the unpleasant pictures were visually explored and emotionally processed after participants responded to the task. In conclusion, the present study supports the competition model of selective attention, highlighting the dominance of top-down control in shaping perceptual and cognitive processing. |
Hamideh Norouzi; Mohammad Reza Daliri Microstate dynamics in working memory: Exploring spatial information coding of stimulus and behavioral performance Journal Article In: Brain and Behavior, vol. 15, no. 8, pp. 1–10, 2025. @article{Norouzi2025a,Background: EEG microstate analysis provides insights into the spatial and temporal dynamics of brain activity during cognitive tasks. The four canonical microstates (classes A, B, C, and D) have been widely reported and associated with various cognitive functions. However, the relationship between microstate parameters and behavioral responses in cognitive functions, such as working memory (WM), has not been sufficiently investigated. This study investigates how microstate dynamics relate to WM performance during a memory-guided saccade (MGS) task. Methods EEG and Eye-tracking data were recorded from participants performing an MGS task at two target eccentricities (near and far). Saccade error was used as a behavioral index of WM performance. Microstate parameters (occurrence, coverage, duration, and transition probability) were computed for the four canonical microstates during the trials. Results Our analysis revealed a significant reduction in the coverage of microstate C, often associated with the default mode network, during the memory maintenance interval compared to baseline. Moreover, a notable increase was observed in the duration of microstate D, considering polarity during the memory interval, which could be related to the frontoparietal control network (FPCN). Notably, the transition probability (TP) from D+ to D- during the memory duration correlated with saccade errors, indicating a behavioral predictive capacity. Furthermore, we identified distinct patterns of microstate D transitions to other microstates that differed significantly between the near and far target conditions, suggesting a functional role in spatial coding. Conclusion Microstate dynamics, particularly those of microstate D, play a dual role in spatial WM by supporting information coding and predicting behavioral accuracy. The polarity-specific transitions within microstate D provide a neural signature of WM performance, with implications for understanding network-level mechanisms underlying spatial memory and saccade control. |
Verónica Mäki-Marttunen; Sander Tjalling Nieuwenhuis Neuromodulatory influences on propagation of traveling waves along the unimodal-transmodal gradient Journal Article In: Cerebral Cortex, vol. 35, no. 7, pp. 1–11, 2025. @article{MaekiMarttunen2025,Understanding the factors underlying brain activity fluctuation is important to understand the flexible nature of the brain and cognition. Growing evidence indicates that functional magnetic resonance imaging (fMRI) activity travels as waves around global signal peaks following a unimodal-transmodal gradient. This may explain the organization of brain activity into functional networks, but why the strength of integration between networks fluctuates is uncertain. Given that arousal-related neuromodulatory systems affect network integration and that traveling waves are modulated by arousal, we aimed to assess the hypothesis that an increase in neuromodulatory tone can affect network integration by modulating the speed of propagation of traveling waves. We tested this hypothesis using pharmacological fMRI/pupil measurements during rest and tasks. Atomoxetine, which increases extracellular catecholamine levels, was associated with faster traveling waves, and faster traveling waves correlated with more network integration. We also examined temporal variations in pupil size, a signature of transient changes in neuromodulatory activity, and found that the periods of traveling waves were characterized by larger pupil size. Our results suggest that neuromodulatory tone affects traveling wave propagation, and that this arousal-modulated propagation shapes integrated functional connectivity features, highlighting specific effects of prolonged and transient neuromodulatory influences on slow brain dynamics. |
Michele Bevilacqua; Fabienne Windel; Elena Beanato; Pauline Menoud; Sarah Zandvliet; Nicola Ramdass; Lisa Fleury; Julie Hervé; Krystel R. Huxlin; Friedhelm C. Hummel; Estelle Raffin Pathway-dependent brain stimulation responses indicate motion processing integrity after stroke Journal Article In: Brain, vol. 148, no. 7, pp. 2361–2372, 2025. @article{Bevilacqua2025,Homonymous hemianopia (HH), a common visual impairment resulting from occipital lobe lesions, affects a significant number of stroke survivors. Intensive perceptual training can foster recovery, possibly by enhancing surviving visual pathways. This study employed cortico-cortical paired associative stimulation (ccPAS) to induce associative plasticity within the residual and bidirectional primary visual cortex (V1)-middle temporal area (MT) pathways in stroke patients. We used ccPAS, which is thought to tap into Hebbian-like spike-timing dependent plasticity, over a motion processing pathway in stroke patients to transiently improve visual motion discrimination in their blind field. Sixteen stroke patients participated in this double-blind, crossover study comparing the effects of bidirectional ccPAS (V1-to-MT or MT-to-V1) on motion discrimination and EEG-Granger Causality. Additionally, we explored potential multimodal sources of inter-individual variability. Results showed that MT-to-V1 ccPAS enhanced motion direction discrimination, but the expected electrophysiological increase in top-down MT-to-V1 inputs was observed only in patients who showed improvement in motion discrimination. Good responders to MT-V1 ccPAS also demonstrated improved functional coupling between the cortical motion pathway and other relevant areas in the visual network, as well as more preserved ipsilesional V1-MT structural integrity. These findings indicate that targeted ccPAS can effectively engage functionally relevant residual visual pathways in stroke-affected brains, potentially offering new avenues for patient stratification and visual recovery strategies. |
Srishty Aggarwal; Supratim Ray Changes in Higuchi fractal dimension across age in healthy human EEG are anticorrelated with changes in oscillatory power and 1/f slope Journal Article In: European Journal of Neuroscience, vol. 62, no. 2, pp. 1–17, 2025. @article{Aggarwal2025,Nonlinear dynamical methods such as Higuchi fractal dimension (HFD) are often used to study the complexities of brain activity. In human electroencephalogram (EEG), although power in the gamma band (30–70 Hz) and the slope of the power spectral density (PSD) have been shown to reduce with healthy ageing, there are conflicting findings regarding how HFD and other measures of complexity vary with ageing. Further, the dependence of HFD on features obtained from PSD (such as gamma power and slope) has not been thoroughly probed. To address these issues, we computed time- and frequency-resolved HFD for EEG data collected from an elderly population (N = 217), aged between 50 and 88 years, for baseline (BL) eyes open state and during a fixation task in which visual grating stimuli that induce strong gamma oscillations were presented. During BL, HFD increased with age for frequencies up to 150 Hz but surprisingly showed an opposite trend at higher frequencies. Interestingly, this change in HFD was opposite to the age-related change in PSD 1/f slope. Further, stimulus-related changes in HFD were anticorrelated with the changes in oscillatory power. However, stimulus- and age-related changes in HFD persisted even after normalization with surrogates, showing the effect of nonlinear dynamics on HFD. Further, age classification using HFD was slightly better than classification using spectral features (power and slope). Therefore, HFD could be jointly sensitive to various spectral features as well as some nonlinearities not captured using spectral analysis, which could enhance our understanding of brain dynamics underlying healthy ageing. |
Canhuang Luo; Edward F. Ester Traveling waves link human visual and frontal cortex during working memory–guided behavior Journal Article In: PNAS, vol. 122, no. 30, pp. 1–9, 2025. @article{Luo2025,Traveling waves guide the spatial propagation of neural activity and computational processes across the brain. Traveling waves could contribute to the control of memory-guided behaviors by flexibly organizing the timing and direction of interactions between brain regions responsible for storing memory content with those responsible for producing task-relevant behaviors. Using short-term memory as a test case, we report evidence supporting this possibility. Analyses of human EEG data revealed forward- and backward-propagating traveling waves linking visual and motor brain areas around the time of a memory-guided behavior. These waves predicted intra- and interindividual differences in task performance, could not be explained by eye movements or passive volume conduction, and were absent when participants planned but could not execute a memory-guided behavior. These findings implicate traveling waves as a mechanism in the initiation and control of memory-guided behaviors. |
Sabine Prantner; Alejandro Espino-Payá; M. Carmen Pastor; Cristina Giménez-García; Rafael Ballester-Arnal; Markus Junghoefer In: Psychophysiology, vol. 62, no. 7, pp. 1–21, 2025. @article{Prantner2025,Gender identity and sexual orientation form fundamental characteristics of an individual's sexual identity and relate to patterns of physiological and neural activity involved in processing erotic or explicit sexual stimuli. To investigate this, we used high-density magnetoencephalography (MEG) to measure brain responses of hetero- and homosexual women and men to opposite- and same-sex erotic images, as well as sexually explicit images. Additionally, we administered pupillometry and subjective measures of hedonic valence and emotional arousal. Erotic versus sexually explicit stimuli initially resulted in enhanced pupil dilation and stronger neural activity in the extended visual cortex, but at later times, reverse effects were found. Our results further showed that perceived affect varied by gender and sexual orientation, with significant group effects. Pupil measurements revealed differences in dilation depending on opposite- and same-sex erotic and sexually explicit images and participant groups. Similarly, effects of stimuli content were found for the neural activity. The findings suggest that preferred versus non-preferred stimuli are subjectively processed in a category-specific way, especially in hetero- and homosexual males as well as homosexual women compared to heterosexual women, and indicate a sensitivity to sexual images in affective-motivational and reward areas of the brain. To conclude, subjective, visual, and neural responses to sexually relevant stimuli seem partly dependent on gender and sexual orientation but predominanly indicate influences of stimulus content. |
Ankita Sengupta; Devarajan Sridharan Reward expectation yields distinct effects on sensory processing and decision making in the human brain Journal Article In: PLoS Biology, vol. 23, no. 7, pp. 1–33, 2025. @article{Sengupta2025,Reward expectation robustly guides both attention and decisions. Yet, whether common or distinct mechanisms mediate each of these processes remains unknown. Previous studies have often conflated the effect of reward expectation on sensory processing and decision-making because locations selected for sensory prioritization (sensitivity effects) were also prioritized for decisions (criterion effects). Here, we identify distinct forms of reward expectation that separably control spatial attention and decisional biases in human cortex. Sensitivity and criterion were independently modulated when expected rewards varied across locations (“space-specific”) or choices (“choice-specific”), respectively. Only sensitivity, not criterion, modulations reflected a limited, conserved attentional resource. Established neural and physiological signatures of attention, including gain modulation of event-related potentials, alpha-band power lateralization, and eye-movement biases, were elicited only by space-specific reward modulation. By contrast, neural correlates of decisional biases, including pre-stimulus alpha power suppression, selectively accompanied choice-specific reward modulation. Attention-related neural markers predicted sensitivity modulation by space-specific reward expectation but not criterion modulation by choice-specific reward expectation, indicating their distinct underlying mechanisms. Our findings uncover fundamentally dissociable behavioral and neural underpinnings of reward expectation effects on sensory and decisional selection, with critical implications for understanding how reward, attention, and choice are linked in the human brain. they must also choose to act quickly and pick off this fruit, to thwart their conspecifics. In other words, reward expectation can influence not only how attention is engaged but also how choices are made. Yet, these two effects have been frequently conflated in laboratory tasks. Here, with a task that decouples reward expectation's effects on attention from those on decision-making we uncover their distinct neural correlates. Our results show how reward shapes attention and biases choices independently in the human brain. |
Jing Zhu; Yuanlong Li; Changlin Yang; Hanshu Cai; Xiaowei Li; Bin Hu Transformer-based fusion model for mild depression recognition with EEG and pupil area signals Journal Article In: Medical and Biological Engineering and Computing, vol. 63, no. 7, pp. 2011–2027, 2025. @article{Zhu2025c,Early detection and treatment are crucial for the prevention and treatment of depression; compared with major depression, current researches pay less attention to mild depression. Meanwhile, analysis of multimodal biosignals such as EEG, eye movement data, and magnetic resonance imaging provides reliable technical means for the quantitative analysis of depression. However, how to effectively capture relevant and complementary information between multimodal data so as to achieve efficient and accurate depression recognition remains a challenge. This paper proposes a novel Transformer-based fusion model using EEG and pupil area signals for mild depression recognition. We first introduce CSP into the Transformer to construct single-modal models of EEG and pupil data and then utilize attention bottleneck to construct a mid-fusion model to facilitate information exchange between the two modalities; this strategy enables the model to learn the most relevant and complementary information for each modality and only share the necessary information, which improves the model accuracy while reducing the computational cost. Experimental results show that the accuracy of the EEG and pupil area signals of single-modal models we constructed is 89.75% and 84.17%, the precision is 92.04% and 95.21%, the recall is 89.5% and 71%, the specificity is 90% and 97.33%, the F1 score is 89.41% and 78.44%, respectively, and the accuracy of mid-fusion model can reach 93.25%. Our study demonstrates that the Transformer model can learn the long-term time-dependent relationship between EEG and pupil area signals, providing an idea for designing a reliable multimodal fusion model for mild depression recognition based on EEG and pupil area signals. |
Maggie E. Zink; Leslie Zhen; Jacie R. McHaney; Jennifer Klara; Kimberly Yurasits; Victoria E. Cancel; Olivia Flemm; Claire Mitchell; Jyotishka Datta; Bharath Chandresekaran; Aravindakshan Parthasarathy Increased listening effort and cochlear neural degeneration underlie speech-in-noise deficits in normal-hearing middle-aged adults Journal Article In: eLife, vol. 13, pp. 1–27, 2025. @article{Zink2025,Middle age represents a critical period of accelerated brain changes and provides a window for early detection and intervention in age-related neurological decline. Hearing loss is a key early marker of such decline and is linked to numerous comorbidities in older adults. Yet, ~10% of middle-aged individuals who report hearing difficulties show normal audiograms. Cochlear neural degeneration (CND) could contribute to these hidden hearing deficits, though its role remains unclear due to a lack of objective diagnostics and uncertainty regarding its perceptual outcomes. Here, we employed a cross-species design to examine neural and behavioral signatures of CND. We measured envelope following responses (EFRs) – neural ensemble responses to sound originating from the peripheral auditory pathway – in young and middle-aged adults with normal audiograms and compared these responses to young and middle-aged Mongolian gerbils, where CND was histologically confirmed. We observed near-identical changes in EFRs across species that were associated with CND. Behavioral assessments revealed age-related speech-in-noise deficits under challenging conditions, while pupil-indexed listening effort increased with age even when behavioral performance was matched. Together, these results demonstrate that CND contributes to speech perception difficulties and elevated listening effort in midlife, which may ultimately lead to listening fatigue and social withdrawal. |
Jasmine Pan; Louis N. Vinke; Joseph T. McGuire; Sam Ling Heterogeneous effects of cognitive arousal on the contrast response in human visual cortex Journal Article In: Journal of Neuroscience, vol. 45, no. 25, pp. 1–14, 2025. @article{Pan2025a,While animal studies have found that arousal states modulate visual responses, direct evidence for effects of arousal on human vision remains limited. Here, we used fMRI to examine effects of cognitive arousal on the gain of contrast response functions (CRFs) in human visual cortex. To measure CRFs, we measured BOLD responses in early visual cortex (V1–V3) while participants (n = 20, 14 females and 6 males) viewed stimuli that parametrically varied in contrast. To induce different cognitive arousal states, participants solved auditory arithmetic problems categorized as either Easy (low arousal) or Hard (high arousal). We found diversity in the modulatory effects across individuals: some individuals exhibited enhanced neural response with increased arousal, whereas others exhibited the opposite effect—a decrease in response with increased arousal. The pattern of overall BOLD modulation showed within-individual stability and was correlated with the degree of arousal-driven change in pupil size. Individuals who exhibited larger increases in pupil size with the arousal manipulation tended to show greater arousal-related decreases in visuocortical responses. We speculate that the polarity of the modulatory effect by cognitive arousal may relate to individual differences in cognitive effort expended in the high-difficulty condition, with individuals reaching different points on an underlying non-monotonic function. |
Gabriel Wainstein; Christopher J. Whyte; Kaylena A. Ehgoetz Martens; Eli J. Müller; Vicente Medel; Britt Anderson; Elisabeth Stöttinger; James Danckert; Brandon R. Munn; James M. Shine Evidence from pupillometry, fMRI, and RNN modelling shows that gain neuromodulation mediates task-relevant perceptual switches Journal Article In: eLife, vol. 13, pp. 1–26, 2025. @article{Wainstein2025,Perceptual updating has been hypothesised to rely on a network reset modulated by bursts of ascending neuromodulatory neurotransmitters, such as noradrenaline, abruptly altering the brain's susceptibility to changing sensory activity. To test this hypothesis at a large-scale, we analysed an ambiguous figures task using pupillometry and functional magnetic resonance imaging (fMRI). Behaviourally, qualitative shifts in the perceptual interpretation of an ambiguous image were associated with peaks in pupil diameter, an indirect readout of phasic bursts in neuromodulatory tone. We further hypothesised that stimulus ambiguity drives neuromodulatory tone, leading to heightened neural gain, hastening perceptual switches. To explore this hypothesis computationally, we trained a recurrent neural network (RNN) on an analogous perceptual categorisation task, allowing gain to change dynamically with classification uncertainty. As predicted, higher gain accelerated perceptual switching by transiently destabilising the network's dynamical regime in periods of maximal uncertainty. We leveraged a low-dimensional readout of the RNN dynamics to develop two novel macroscale predictions: perceptual switches should occur with peaks in low-dimensional brain state velocity and with a flattened egocentric energy landscape. Using fMRI, we confirmed these predictions, highlighting the role of the neuromodulatory system in the large-scale network reconfigurations mediating adaptive perceptual updates. |
Ralph Andrews; Michael C. Melnychuk; Catherine N. Moran; David P. McGovern; Alexa Holfelder; Sarah Moran; Paul M. Dockree Arousal and sustained attention fluctuate differently with respiration in younger and older adults Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–21, 2025. @article{Andrews2025,Respiration is being increasingly recognised as both synchronising its dynamics with external events and modulating internal psychophysiological states. However, the extent to which these effects stem from a respiratory modulation of attention remains underexplored. Here, we leverage differing attentional strategies of younger (YA) and older adults (OA)—OA exhibited greater focus during a simple contrast change detection task—to examine their relationship with respiratory phase-locking behaviour. OA exhibited stronger phase-locking of their respiratory cycle to task-relevant events compared with YA. Notably, participants appeared to actively adjust their breathing so that late exhalation phases coincided with target presentation, despite variable inter-target intervals. To characterise this target-locked respiratory phase window, we analysed pupil diameter and EEG frequency-power as indices of arousal and attention. Pupil diameter, frontal delta and theta, posterior alpha, and steady-state visually evoked potential (SSVEP) amplitude all varied significantly over the respiratory cycle, suggesting that arousal was enhanced for respiratory phases aligned with target expectancy and attenuated outside these phases. OA showed stronger respiratory modulation of delta, theta, and alpha, whereas YA showed stronger modulation of pupil diameter and SSVEP. We interpret these findings as evidence that respiration shapes attentional fluctuations, expanding and contracting the vigilant state across the respiratory cycle through interactions with arousal and attentional systems. Further, the age-dependent quality of attention which is applied to a task has implications for the degree of respiratory phase-locking and how physiological signatures of arousal and attention are modulated. |
Cora Fischer; Jochen Kaiser; Christoph Bledowski A direct neural signature of serial dependence in working memory Journal Article In: eLife, vol. 13, pp. 1–25, 2025. @article{Fischer2025a,Serial dependence describes the phenomenon that current object representations are attracted to previously encoded and reported representations. While attractive biases have been observed reliably in behavior, a direct neural correlate has not been established. Previous studies have either shown a reactivation of past information without observing a neural signal related to the bias of the current information, or a repulsive distortion of current neural representations contrasting the behavioral bias. The present study recorded neural signals with magnetoencephalography (MEG) during a working memory task to identify neural correlates of serial dependence. Participants encoded and memorized two sequentially presented motion directions per trial, one of which was later retro-cued for report. Multivariate analyses provided reliable reconstructions of both motion directions. Importantly, the reconstructed directions in the current trial were attractively shifted toward the target direction of the previous trial. This neural bias mirrored the behavioral attractive bias, thus reflecting a direct neural signature of serial dependence. The use of a retro-cue task in combination with MEG allowed us to determine that this neural bias emerged at later, post-encoding time points. This timing suggests that serial dependence in working memory affects memorized information during read-out and reactivation processes that happen after the initial encoding. |
Katharina Lingelbach; Jochem W. Rieger Neurophysiological basis of emotional face perception and working memory load in a dual-task MEG study Journal Article In: Human Brain Mapping, vol. 46, no. 8, pp. 1–25, 2025. @article{Lingelbach2025,Research on the neurophysiological effects of emotional face processing, working memory (WM) load, and their interaction in dual-tasks remains scarce. Therefore, we conducted a combined magnetoencephalography eye-tracking study with 47 participants. The dual-task temporally interleaved a facial emotion discrimination task with a visuo-spatial n-back task. Source-space cluster analyzes of event-related magnetic fields (ERFs) and oscillations revealed significant main effects of emotional expression and WM load. During emotion discrimination, enhanced ERFs for negative facial expressions located across the insula, ACC, and face-specific occipital regions suggest amplified emotion processing but also the recruitment of attentional control mechanisms. During the n-back phase, emotional faces did not affect evoked responses when they were task-irrelevant. Interaction trends in pupil dilation indicated that emotion-specific processing is diminished under high WM load. During the n-back phase, increased WM load reduced alpha and low beta oscillations in temporo- and parieto-occipital areas. In addition, reduced target fixations in the presence of negative facial distractors indicated a tendency toward emotion-specific interference. Furthermore, sustained increased WM load affected perceived valence, pupil size, and reaction time in both subtasks. A convergence of neurophysiological, physiological, and behavioural findings points to specific processing modes with greater resource depletion for negative expressions and high WM load in the dual-task. In conclusion, the study advanced our understanding of (a) circumstances under which emotional faces modulate ERFs in a dual-task, (b) mechanisms underlying emotion discrimination, (c) interaction effects of emotional expression and WM load in gaze behavior, as well as (d) how WM-related oscillatory alpha and beta power is affected by increasing load. |
Amir Norouzpour; Tawna L. Roberts Repeated measures analysis for steady-state evoked potentials Journal Article In: Computers in Biology and Medicine, vol. 191, pp. 1–10, 2025. @article{Norouzpour2025,Introduction: Brain response to repetitive stimuli generates steady-state evoked potentials (ssEP) that vary depending on the experimental conditions. To analyze these responses, Fourier measurements extracted from ssEP data require statistical techniques to differentiate neural responses across various experimental conditions within the same participant(s). In this study, we introduce new statistical methods to compare multiple dependent clusters of discrete Fourier measurements corresponding to multiple experimental conditions. Methods: We present two statistics: 1) The first statistic is derived from repeated measures analysis of variance (ANOVA) for complex numbers, used to compare multiple dependent circular clusters of Fourier estimates under the assumption of equal variance across the clusters. 2) The second statistic is employed when either the assumption of circularity within the clusters or the assumption of equal variance across the clusters is violated. In this case, we derive the statistic from the rank-sum Friedman test to compare multiple related clusters of complex numbers. Results: We demonstrated the validity of the statistics using simulated and empirical ssEP data. Our methods offer robust statistical tools that maintain a constant Type-I error of 0.05 in all conditions, including equal or unequal variance-covariance matrix of the real and imaginary components of Fourier estimates across the circular and elliptical clusters, even in the presence of outliers in the dataset. Furthermore, our statistics demonstrate a lower Type-II error compared to repeated measures multivariate analysis of variance (rmMANOVA). Conclusion: The statistical methods enable us to compare multiple dependent clusters of Fourier estimates corresponding to multiple experimental conditions within the same participant(s), whether or not the variance is equal across the circular or elliptical clusters, even with outliers in the dataset. |
Daniel J. Pearce; Gerard M. Loughnane; Trevor T. J. Chong; Nele Demeyere; Jason B. Mattingley; Margaret J. Moore; Peter W. New; Redmond G. O'Connell; Megan H. O'Neill; Dragan Rangelov; Renerus J. Stolwyk; Sam S. Webb; Shou Han Zhou; Méadhbh B. Brosnan; Mark A. Bellgrove Target selection signals causally influence human perceptual decision-making Journal Article In: The Journal of Neuroscience, vol. 45, no. 24, pp. 1–13, 2025. @article{Pearce2025,The ability to form decisions is a foundational cognitive function which is impaired across many psychiatric and neurological conditions. Understanding the neural processes underpinning clinical deficits may provide insights into the fundamental mechanisms of decision-making. The N2c has been identified as an EEG signal indexing the efficiency of early target selection, which subsequently influences the timing of perceptual reports through modulating neural evidence accumulation rates. Evidence for the contribution of the N2c to human decision-making however has thus far come from correlational research in neurologically healthy individuals. Here, we capitalized on the superior temporal resolution of EEG to show that unilateral brain lesions in male and female humans were associated with specific deficits in both the timing and strength of the N2c in the damaged hemisphere, with corresponding deficits in the timing of perceptual reports contralaterally. The extent to which the N2c influenced clinical deficits in perceptual reporting speed depended on neural rates of evidence accumulation. This work provides causal evidence that the N2c indexes an early, hemisphere-specific process supporting human decision-making. This noninvasive EEG marker could be used to monitor novel approaches for remediating clinical deficits in perceptual decision-making across a range of brain disorders. |
Luis D. Ramirez; Feiyi Wang; Sam Ling Attention alters population spatial frequency tuning Journal Article In: Journal of Neuroscience, vol. 45, no. 25, pp. 1–12, 2025. @article{Ramirez2025,Spatial frequency (SF) selectivity serves as a fundamental building block within the visual system, determining what we can and cannot see. Attention is theorized to augment the visibility of items in our environment by changing how we process SFs. However, the specific neural mechanisms underlying this effect remain unclear, particularly in humans. Here, we used functional magnetic resonance imaging to measure voxel-wise population SF tuning (pSFT), which allowed us to examine how attention alters the SF response profiles of neural populations in the early visual cortex (V1–V3). In the scanner, participants (five female, three male) were cued to covertly attend to one of two spatially competing letter streams, each defined by low or high SF content. This task promoted feature-based attention directed to a particular SF, as well as the suppression of the irrelevant stream's SF. Concurrently, we measured pSFT in a task-irrelevant hemifield to examine how the known spatial spread of feature-based attention influenced the SF tuning properties of neurons sampled within a voxel. We discovered that attention elicited attractive shifts in SF preference, toward the attended SF. This suggests that attention can profoundly influence populations of SF preference across the visual field, depending on task goals and native neural preferences. |
Andrea I. Costantino; Benjamin O. Pelzer; Mark A. Williams; Matthew J. Crossley Partial information transfer from peripheral visual streams to foveal visual streams may be mediated through local primary visual circuits Journal Article In: NeuroImage, vol. 311, pp. 1–14, 2025. @article{Costantino2025,Visual object recognition is driven through the what pathway, a hierarchy of visual areas processing features of increasing complexity and abstractness. The primary visual cortex (V1), this pathway's origin, exhibits retinotopic organization: neurons respond to stimuli in specific visual field regions. A neuron responding to a central stimulus will not respond to a peripheral one, and vice versa. However, despite this organization, task-relevant feedback about peripheral stimuli can be decoded in unstimulated foveal cortex, and disrupting this feedback impairs discrimination behavior. The information encoded by this feedback remains unclear, as prior studies used computer-generated objects ill-suited to dissociate different representation types. To address this knowledge gap, we investigated the nature of information encoded in periphery-to-fovea feedback using real-world stimuli. Participants performed a same/different discrimination task on peripherally displayed images of vehicles and faces. Using fMRI multivariate decoding, we found that both peripheral and foveal V1 could decode images separated by low-level perceptual models (vehicles) but not those separated by semantic models (faces). This suggests the feedback primarily carries low-level perceptual information. In contrast, higher visual areas resolved semantically distinct images. A functional connectivity analysis revealed foveal V1 connections to both peripheral V1 and later-stage visual areas. These findings indicate that while both early and late visual areas may contribute to information transfer from peripheral to foveal processing streams, higher-to-lower area transfer may involve information loss. |
Marvin R. Maechler; Eunhye Choe; Patrick Cavanagh; Peter J. Kohler; Peter U. Tse Hemifield specificity of attention response functions during multiple-object tracking Journal Article In: Journal of Neuroscience, vol. 45, no. 19, pp. 1–12, 2025. @article{Maechler2025,The difficulty of tracking multiple moving objects among identical distractors increases with the number of tracked targets. Previous research has shown that the number of targets tracked (i.e., load) modulates activity in brain areas related to visuospatial attention, giving rise to so-called attention response functions (ARFs). While the hemifield/hemispheric effects of spatial attention (e.g., hemispatial neglect, hemifield capacity limits) are well described, it had not previously been tested whether a hemispheric or hemifield imbalance exists among ARFs. By recording blood oxygenation level-dependent activity from human brains (n = 19, female and male) in a multiple-object tracking paradigm, we show that the number of tracked objects modulates activity in a large network of areas bilaterally. A significant effect of contralateral load was found in earlier areas throughout the dorsal and ventral visual streams, while the effects of ipsilateral load emerged in later areas. Both contra- and ipsilateral load significantly influenced activity in the parietal and frontal lobes, specifically the dorsal attention network. In addition, some brain regions in the occipital lobe were significantly more sensitive to contralateral than ipsilateral load. Our results are consistent with findings showing that a diverse set of brain areas contributes to tracking multiple targets. In particular, we extend the canonical view of load-based ARFs to include hemifield bias. Given the hemifield-specific nature of speed and capacity limits to multiple-object tracking, we conjecture that areas that show a strong hemifield preference may impose a bottleneck on processing that results in limits on the capacity and speed of tracking. |
Julia Elina Stocker; Antonia Schulz; Ina Thome; Jens Sommer; Jonas Rabeneck; Kristin Marie Rusch; Olaf Steinsträter; Andreas Jansen Interhemispheric integration in the neural face perception network: Does stimulus location matter? Journal Article In: Imaging Neuroscience, vol. 3, pp. 1–16, 2025. @article{Stocker2025,The neural mechanisms underlying hemispheric lateralization can be investigated using neuroimaging methods and modelling techniques. In some experiments, sensory information is initially presented exclusively to one hemisphere, for example, by displaying a visual stimulus in the periphery of the contralateral hemifield. This experimental design enables, among other things, a comparison of competing theories of interhemispheric integration (e.g., interhemispheric inhibition vs. interhemispheric recruitment). However, the underlying neural models for peripheral stimulation may differ from those for central stimulation and, therefore, may not adequately describe the mechanisms associated with typical, foveal stimulus processing. To address this question, the present functional magnetic resonance imaging (fMRI) study analysed the influence of stimulus location (peripheral vs. central) on neural network connectivity, particularly interhemispheric transfer, as determined by dynamic causal modelling (DCM), for a face perception task. Face and object images were presented either peripherally or centrally to a group of healthy volunteers (N = 17). By contrasting brain activations for faces against objects, we identified bilateral face-sensitive regions, such as the left and right fusiform face area (FFA) and the occipital face area (OFA). Additionally, we extracted the bilateral primary visual cortex (V1) as the input region for our neural models. We constructed five increasingly complex models that differed only in their modulatory connectivity. Bayesian model averaging (BMA) was employed to average the parameters across all models, enabling the calculation of interhemispheric transfer difference (i.e., left-to-right minus right-to-left modulatory connectivity parameter) and the strength of interhemispheric transfer between bilateral OFA and FFA regions. Our findings demonstrate that interhemispheric integration depends on stimulus location. Peripheral presentations of faces induce different connectivity patterns compared with centrally depicted faces. Specifically, we observed larger interhemispheric transfer differences for peripheral face stimuli compared with central stimuli. In conclusion, peripheral and central presentations of faces modulate the face processing network differently, with left and right visual field presentations yielding asymmetrical connectivity patterns. Since faces are preferentially processed via the fovea, the typical face processing network likely aligns more closely with activation patterns elicited by central stimuli. In contrast, connectivity patterns triggered by peripheral stimulation may represent an atypical processing style and cannot be directly compared with those activated by central stimuli. |
Markus Conci; Leonie Nowack; Paul C. J. Taylor; Kathrin Finke; Hermann J. Müller Right parietal rTMS induces bidirectional effects of selective attention upon object integration Journal Article In: Brain Sciences, vol. 15, no. 5, pp. 1–19, 2025. @article{Conci2025,Background/Objectives: Part-to-whole object completion and search guidance by salient, integrated objects has been proposed to require attentional resources, as shown by studies of neglect patients suffering from right-parietal brain damage. The current study was performed to provide further causal evidence for the link between attention and object integration. Methods: Healthy observers detected targets in the left and/or right hemifields, and these targets were in turn embedded in various Kanizsa-type configurations that systematically varied in the extent to which individual items could be integrated into a complete, whole object. Moreover, repetitive transcranial magnetic stimulation (rTMS) was applied over the right intraparietal sulcus (IPS) and compared to both active and passive baseline conditions. Results: The results showed that target detection was substantially facilitated when the to-be detected item(s) were fully embedded in a salient, grouped Kanizsa figure, either a unilateral triangle or a bilateral diamond. However, object groupings in one hemifield did not facilitate target detection to the same extent when there were bilateral targets, one inside the (triangle) grouping and the other outside of the grouped object. These results extend previous findings from neglect patients. Moreover, a subgroup of observers was found to be particularly sensitive to IPS stimulation, revealing neglect-like extinction behavior with the single-hemifield triangle groupings and bilateral targets. Conversely, a second subgroup showed the opposite effect, namely an overall, IPS-dependent improvement in performance. Conclusions: These explorative analyses show that the parietal cortex, in particular IPS, seems to modulate the processing of object groupings by up- and downregulating the deployment of attention to spatial regions were to-be-grouped items necessitate attentional resources for object completion. |