EyeLink Usability / Applied Publications
All EyeLink usability and applied research publications up until 2023 (with some early 2024s) are listed below by year. You can search the publications using keywords such as Driving, Sport, Workload, etc. You can also search for individual author names. If we missed any EyeLink usability or applied article, please email us!
2022 |
Kumari Liza; Supratim Ray Local interactions between steady-state visually evoked potentials at nearby flickering frequencies Journal Article In: Journal of Neuroscience, vol. 42, no. 19, pp. 3965–3974, 2022. @article{Liza2022, Steady-state visually evoked potentials (SSVEPs) are widely used to index top-down cognitive processing in human electroencephalogram (EEG) studies. Typically, two stimuli flickering at different temporal frequencies (TFs) are presented, each producing a distinct response in the EEG at its flicker frequency. However, how SSVEP responses in EEGs are modulated in the presence of a competing flickering stimulus just because of sensory interactions is not well understood. We have previously shown in local field potentials (LFPs) recorded from awake monkeys that when two overlapping full-screen gratings are counterphased at different TFs, there is an asymmetric SSVEP response suppression, with greater suppression from lower TFs, which further depends on the relative orientations of the gratings (stronger suppression and asymmetry for parallel compared with orthogonal gratings). Here, we first confirmed these effects in both male and female human EEG recordings. Then, we mapped the response suppression of one stimulus (target) by a competing stimulus (mask) over a much wider range than the previous study. Surprisingly, we found that the suppression was not stronger at low frequencies in general, but systematically varied depending on the target TF, indicating local interactions between the two competing stimuli. These results were confirmed in both human EEG and monkey LFP and electrocorticogram (ECoG) data. Our results show that sensory interactions between multiple SSVEPs are more complex than shown previously and are influenced by both local and global factors, underscoring the need to cautiously interpret the results of studies involving SSVEP paradigms.SIGNIFICANCE STATEMENT Steady-state visually evoked potentials (SSVEPs) are extensively used in human cognitive studies and brain-computer interfacing applications where multiple stimuli flickering at distinct frequencies are concurrently presented in the visual field. We recently characterized interactions between competing flickering stimuli in animal recordings and found that stimuli flickering slowly produce larger suppression. Here, we confirmed these in human EEGs, and further characterized the interactions by using a much wider range of target and competing (mask) frequencies in both human EEGs and invasive animal recordings. These revealed a new "local" component, whereby the suppression increased when competing stimuli flickered at nearby frequencies. Our results highlight the complexity of sensory interactions among multiple SSVEPs and underscore the need to cautiously interpret studies involving SSVEP paradigms. |
Angela Radetz; Markus Siegel Spectral fingerprints of cortical neuromodulation Journal Article In: Journal of Neuroscience, vol. 42, no. 18, pp. 3836–3846, 2022. @article{Radetz2022, Pupil size has been established as a versatile marker of noradrenergic and cholinergic neuromodulation, which has profound effects on neuronal processing, cognition, and behavior. However, little is known about the cortical control and effects of pupil-linked neuromodulation. Here, we show that pupil dynamics are tightly coupled to temporally, spectrally, and spatially specific modulations of local and large-scale cortical population activity in the human brain. We quantified the dynamics of band-limited cortical population activity in resting human subjects using magnetoencephalography and investigated how neural dynamics were linked to simultaneously recorded pupil dynamics. Our results show that pupil-linked neuromodulation does not merely affect cortical population activity in a stereotypical fashion. Instead, we identified three frontal, precentral, and occipitoparietal networks, in which local population activity with distinct spectral profiles in the theta, beta, and alpha bands temporally preceded and followed changes in pupil size. Furthermore, we found that amplitude coupling at;16 Hz in a large-scale frontoparietal network predicted pupil dynamics. Our results unravel network-specific spectral fingerprints of cortical neuromodulation in the human brain that likely reflect both the causes and effects of neuromodulation. |
Arno Libert; Arne Van Den Kerchove; Benjamin Wittevrongel; Marc M. Van Hulle Analytic beamformer transformation for transfer learning in motion-onset visual evoked potential decoding Journal Article In: Journal of Neural Engineering, vol. 19, pp. 1–16, 2022. @article{Libert2022, Objective. While decoders of electroencephalography-based event-related potentials (ERPs) are routinely tailored to the individual user to maximize performance, developing them on populations for individual usage has proven much more challenging. We propose the analytic beamformer transformation (ABT) to extract phase and/or magnitude information from spatiotemporal ERPs in response to motion-onset stimulation. Approach. We have tested ABT on 52 motion-onset visual evoked potential (mVEP) datasets from 26 healthy subjects and compared the classification accuracy of support vector machine (SVM), spatiotemporal beamformer (stBF) and stepwise linear discriminant analysis (SWLDA) when trained on individual subjects and on a population thereof. Main results. When using phase- and combined phase/magnitude information extracted by ABT, we show significant improvements in accuracy of population-trained classifiers applied to individual users (p < 0.001). We also show that 450 epochs are needed for a correct functioning of ABT, which corresponds to 2 min of paradigm stimulation. Significance. We have shown that ABT can be used to create population-trained mVEP classifiers using a limited number of epochs. We expect this to pertain to other ERPs or synchronous stimulation paradigms, allowing for a more effective, population-based training of visual BCIs. Finally, as ABT renders recordings across subjects more structurally invariant, it could be used for transfer learning purposes in view of plug-and-play BCI applications. |
Ilmari Kurki; Aapo Hyvärinen; Linda Henriksson Dynamics of retinotopic spatial attention revealed by multifocal MEG Journal Article In: NeuroImage, vol. 263, pp. 1–13, 2022. @article{Kurki2022, Visual focal attention is both fast and spatially localized, making it challenging to investigate using human neuroimaging paradigms. Here, we used a new multivariate multifocal mapping method with magnetoencephalography (MEG) to study how focal attention in visual space changes stimulus-evoked responses across the visual field. The observer's task was to detect a color change in the target location, or at the central fixation. Simultaneously, 24 regions in visual space were stimulated in parallel using an orthogonal, multifocal mapping stimulus sequence. First, we used univariate analysis to estimate stimulus-evoked responses in each channel. Then we applied multivariate pattern analysis to look for attentional effects on the responses. We found that attention to a target location causes two spatially and temporally separate effects. Initially, attentional modulation is brief, observed at around 60–130 ms post stimulus, and modulates responses not only at the target location but also in adjacent regions. A later modulation was observed from around 200 ms, which was specific to the location of the attentional target. The results support the idea that focal attention employs several processing stages and suggest that early attentional modulation is less spatially specific than late. |
Timo L. Kvamme; Mesud Sarmanlu; Christopher Bailey; Morten Overgaard In: Neuroscience, vol. 482, pp. 1–17, 2022. @article{Kvamme2022, Spontaneous neural oscillations are key predictors of perceptual decisions to bind multisensory sig- nals into a unified percept. Research links decreased alpha power in the posterior cortices to attention and audio- visual binding in the sound-induced flash illusion (SIFI) paradigm. This suggests that controlling alpha oscillations would be a way of controlling audiovisual binding. In the present feasibility study we used MEG- neurofeedback to train one group of subjects to increase left/right and another to increase right/left alpha power ratios in the parietal cortex. We tested for changes in audiovisual binding in a SIFI paradigm where flashes appeared in both hemifields. Results showed that the neurofeedback induced a significant asymmetry in alpha power for the left/right group, not seen for the right/left group. Corresponding asymmetry changes in audiovisual binding in illusion trials (with 2, 3, and 4 beeps paired with 1 flash) were not apparent. Exploratory analyses showed that neurofeedback training effects were present for illusion trials with the lowest numeric disparity (i.e., 2 beeps and 1 flash trials) only if the previous trial had high congruency (2 beeps and 2 flashes). Our data suggest that the relation between parietal alpha power (an index of attention) and its effect on audiovisual binding is dependent on the learned causal structure in the previous stimulus. The present results suggests that low alpha power biases observers towards audiovisual binding when they have learned that audiovisual signals orig- inate from a common origin, consistent with a Bayesian causal inference account of multisensory perception. |
Timo L. Kvamme; Mesud Sarmanlu; Morten Overgaard Doubting the double-blind: Introducing a questionnaire for awareness of experimental purposes in neurofeedback studies Journal Article In: Consciousness and Cognition, vol. 104, pp. 1–13, 2022. @article{Kvamme2022a, Double-blinding subjects to the experiment's purpose is an important standard in neurofeedback studies. However, it is difficult to provide evidence that humans are entirely unaware of certain information. This study used insights from consciousness studies and neurophenomenology to develop a contingency awareness questionnaire for neurofeedback. We assessed whether participants had an awareness of experimental purposes to manipulate their attention and multisensory perception. A subset of subjects (5 out of 20) gained a degree of awareness of experimental purposes as evidenced by their correct guess about the purposes of the experiment to affect their attention and multisensory perceptions specific to their double-blinded group assignment. The results warrant replication before they are applied to clinical neurofeedback studies, given the considerable time taken to perform the questionnaire (∼25 min). We discuss the strengths and limitations of our contingency awareness questionnaire and the growing appeal of the double-blinded standard in clinical neurofeedback studies. |
Nan Li; Olaf Dimigen; Werner Sommer; Suiping Wang Parafoveal words can modulate sentence meaning: Electrophysiological evidence from an RSVP-with-flanker task Journal Article In: Psychophysiology, vol. 59, pp. 1–18, 2022. @article{Li2022d, During natural reading, readers can take up some visual information from not-yet-fixated words to the right of the current fixation and it is well-established that this parafoveal preview facilitates the subsequent foveal processing of the word. However, the extraction and integration of word meaning from parafoveal words and their possible influence on the semantic content of the sentence are controversial. In the current study, we recorded event-related potentials (ERPs) in the RSVP-with-flankers paradigm to test whether and how updates of sentential meaning, based only on parafoveal information, may influence the subsequent foveal processing. In Chinese sentences, the congruency of parafoveal and foveal target words with the sentence was orthogonally manipulated. In contrast to previous research, we also controlled for potentially confounding effects of parafoveal-to-foveal repetition priming (identity preview effects) on the N400. Crucially, we found that the classic effect of foveal congruency on the N400 component only appeared when the word in preview had been congruent with sentence meaning; in contrast, there was no N400 as a function of foveal incongruency when the preview word had also been incongruent. These results indicate that sentence meaning rapidly adapts to parafoveal preview, altering the semantic context for the subsequently fixated word. We also show that correct parafoveal preview generally attenuates the N400 once a word is fixated, regardless of congruency. Taken together, our findings underline the highly generative and adaptive framework of language comprehension. |
Baiwei Liu; Anna C. Nobre; Freek Ede Functional but not obligatory link between microsaccades and neural modulation by covert spatial attention Journal Article In: Nature Communications, vol. 13, pp. 1–10, 2022. @article{Liu2022, Covert spatial attention is associated with spatial modulation of neural activity as well as with directional biases in fixational eye movements known as microsaccades. We studied how these two ‘fingerprints' of attention are interrelated in humans. We investigated spatial modulation of 8-12 Hz EEG alpha activity and microsaccades when attention is directed internally within the spatial layout of visual working memory. Consistent with a common origin, spatial modulations of alpha activity and microsaccades co-vary: alpha lateralisation is stronger in trials with microsaccades toward versus away from the memorised location of the to-be-attended item and occurs earlier in trials with earlier microsaccades toward this item. Critically, however, trials without attention-driven microsaccades nevertheless show clear spatial modulation of alpha activity – comparable to trials with attention-driven microsaccades. Thus, directional biases in microsaccades correlate with neural signatures of spatial attention, but they are not necessary for neural modulation by spatial attention to be manifest. |
Sasu Mäkelä; Jan Kujala; Riitta Salmelin Removing ocular artifacts from magnetoencephalographic data on naturalistic reading of continuous texts Journal Article In: Frontiers in Neuroscience, vol. 16, pp. 1–18, 2022. @article{Maekelae2022, Naturalistic reading paradigms and stimuli consisting of long continuous texts are essential for characterizing the cortical basis of reading. Due to the highly dynamic nature of the reading process, electrophysiological brain imaging methods with high spatial and temporal resolution, such as magnetoencephalography (MEG), are ideal for tracking them. However, as electrophysiological recordings are sensitive to electromagnetic artifacts, data recorded during naturalistic reading is confounded by ocular artifacts. In this study, we evaluate two different pipelines for removing ocular artifacts from MEG data collected during continuous, naturalistic reading, with the focus on saccades and blinks. Both pipeline alternatives are based on blind source separation methods but differ fundamentally in their approach. The first alternative is a multi-part process, in which saccades are first extracted by applying Second-Order Blind Identification (SOBI) and, subsequently, FastICA is used to extract blinks. The other alternative uses a single powerful method, Adaptive Mixture ICA (AMICA), to remove all artifact types at once. The pipelines were tested, and their effects compared on MEG data recorded from 13 subjects in a naturalistic reading task where the subjects read texts with the length of multiple pages. Both pipelines performed well, extracting the artifacts in a single component per artifact type in most subjects. Signal power was reduced across the whole cortex in all studied frequency bands from 1 to 90 Hz, but especially in the frontal cortex and temporal pole. The results were largely similar for the two pipelines, with the exception that SOBI-FastICA reduced signal in the right frontal cortex in all studied frequency bands more than AMICA. However, there was considerable interindividual variation in the effects of the pipelines. As a holistic conclusion, we choose to recommend AMICA for removing artifacts from MEG data on naturalistic reading but note that the SOBI-FastICA pipeline has also various favorable characteristics. |
Jacqueline Katharina Meier; Bernhard P. Staresina; Lars Schwabe Stress diminishes outcome but enhances response representations during instrumental learning Journal Article In: eLife, vol. 11, pp. 1–25, 2022. @article{Meier2022, Stress may shift behavioural control from a goal-directed system that encodes action-outcome relationships to a habitual system that learns stimulus-response associations. Although this shift to habits is highly relevant for stress-related psychopathologies, limitations of existing behavioural paradigms hinder research from answering the fundamental question of whether the stress-induced bias to habits is due to reduced outcome processing or enhanced response processing at the time of stimulus presentation, or both. Here, we used EEG-based multivariate pattern analysis to decode neural outcome representations crucial for goal-directed control, as well as response representations during instrumental learning. We show that stress reduced outcome representations but enhanced response representations. Both were directly associated with a behavioural index of habitual responding. Furthermore, changes in outcome and response representations were uncorrelated, suggesting that these may reflect distinct processes. Our findings indicate that habitual behaviour under stress may be the result of both enhanced stimulus-response processing and diminished outcome processing. |
Xinzhen Pei; Guiying Xu; Yunhui Zhou; Luna Tao; Xiaozhu Cui; Zhenyu Wang; Bingru Xu; An-Li Wang; Xi Zhao; Haijun Dong; Yan An; Yang Cao; Ruxue Li; Honglin Hu; Yuguo Yu A simultaneous electroencephalography and eye-tracking dataset in elite athletes during alertness and concentration tasks Journal Article In: Scientific Data, vol. 9, pp. 1–15, 2022. @article{Pei2022, The dataset of simultaneous 64-channel electroencephalography (EEG) and high-speed eye-tracking (ET) recordings was collected from 31 professional athletes and 43 college students during alertness behavior task (ABT) and concentration cognitive task (CCT). The CCT experiment lasting 1–2 hours included five sessions for groups of the Shooting, Archery and Modern Pentathlon elite athletes and the controls. Concentration targets included shooting target and combination target with or without 24 different directions of visual distractors and 2 types of music distractors. Meditation and Schulte Grid trainings were done as interventions. Analysis of the dataset aimed to extract effective biological markers of eye movement and EEG that can assess the concentration level of talented athletes compared with same-aged controls. Moreover, this dataset is useful for the research of related visual brain-computer interfaces. |
Thomas Pfeffer; Christian Keitel; Daniel S. Kluger; Anne Keitel; Alena Russmann; Gregor Thut; Tobias H. Donner; Joachim Gross Coupling of pupil-and neuronal population dynamics reveals diverse influences of arousal on cortical processing Journal Article In: eLife, vol. 11, pp. 1–28, 2022. @article{Pfeffer2022, Fluctuations in arousal, controlled by subcortical neuromodulatory systems, continuously shape cortical state, with profound consequences for information processing. Yet, how arousal signals influence cortical population activity in detail has so far only been characterized for a few selected brain regions. Traditional accounts conceptualize arousal as a homogeneous modulator of neural population activity across the cerebral cortex. Recent insights, however, point to a higher specificity of arousal effects on different components of neural activity and across cortical regions. Here, we provide a comprehensive account of the relationships between fluctuations in arousal and neuronal population activity across the human brain. Exploiting the established link between pupil size and central arousal systems, we performed concurrent magnetoencephalographic (MEG) and pupillographic recordings in a large number of participants, pooled across three laboratories. We found a cascade of effects relative to the peak timing of spontaneous pupil dilations: Decreases in low-frequency (2–8 Hz) activity in temporal and lateral frontal cortex, followed by increased highfrequency (>64 Hz) activity in mid-frontal regions, followed by monotonic and inverted U relationships with intermediate frequency-range activity (8–32 Hz) in occipito-parietal regions. Pupil-linked arousal also coincided with widespread changes in the structure of the aperiodic component of cortical population activity, indicative of changes in the excitation-inhibition balance in underlying microcircuits. Our results provide a novel basis for studying the arousal modulation of cognitive computations in cortical circuits. |
Mattia Pietrelli; Jason Samaha; Bradley R. Postle Spectral distribution dynamics across different attentional priority states Journal Article In: Journal of Neuroscience, vol. 42, no. 19, pp. 4026–4041, 2022. @article{Pietrelli2022, Anticipatory covert spatial attention improves performance on tests of visual detection and discrimination, and shifts are accompanied by decreases and increases of a band power at electroencephalography (EEG) electrodes corresponding to the attended and unattended location, respectively. Although the increase at the unattended location is often interpreted as an active mechanism (e.g., inhibiting processing at the unattended location), most experiments cannot rule out the alternative possibility that it is a secondary consequence of selection elsewhere. To adjudicate between these accounts, we designed a Posner- style visual cueing task in which male and female human participants made orientation judgments of targets appearing at one of four locations: up, down, right, or left. Critically, trials were blocked such that within a block the locations along one meridian alternated in status between attended and unattended, and targets never appeared at the other two, making them irrelevant. Analyses of the concurrently measured EEG signal were conducted on “traditional” narrowband a (8–14 Hz), as well as on two components resulting from the decomposition of this signal: “periodic” a;and the slope of the aperiodic 1/f-like component. Although data from right-left blocks replicated the familiar pattern of lateralized asymmetry in narrowband a power, with neither a signal couldwe findevidence for any difference inthe time course at unattended versus irrelevant locations, an outcome consistent with the secondary-consequence interpretation of attention- related dynamics in the a band. Additionally, 1/f slope was shallower at attended and unattended locations, relative to irrelevant, suggesting a tonic adjustment of physiological state. |
Michael Plöchl; Ian Fiebelkorn; Sabine Kastner; Jonas Obleser Attentional sampling of visual and auditory objects is captured by theta-modulated neural activity Journal Article In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3067–3082, 2022. @article{Ploechl2022, Recent evidence suggests that visual attention alternately samples two behaviourally relevant objects at approximately 4 Hz, rhythmically shifting between the objects. Whether similar attentional rhythms exist in other sensory modalities, however, is not yet clear. We therefore adapted and extended an established paradigm to investigate visual and potential auditory attentional rhythms, as well as possible interactions, on both a behavioural (detection performance |
Frida A. B. Printzlau; Nicholas E. Myers; Sanjay G. Manohar; Mark G. Stokes Neural reinstatement tracks spread of attention between object features in working memory Journal Article In: Journal of Cognitive Neuroscience, vol. 34, no. 9, pp. 1681–1701, 2022. @article{Printzlau2022, Attention can be allocated in working memory ( WM) to select and privilege relevant content. It is unclear whether attention selects individual features or whole objects in WM. Here, we used behavioral measures, eye-tracking, and EEG to test the hypothesis that attention spreads between an object's features in WM. Twenty-six participants completed a WM task that asked them to recall the angle of one of two oriented, colored bars after a delay while EEG and eye-tracking data were collected. During the delay, an orthogonal “incidental task” cued the color of one item for a match/mismatch judgment. On congruent trials (50%), the cued item was probed for subsequent orientation recall; on incongruent trials (50%), the other memory item was probed. As predicted, selecting the color of an object in WM brought other features of the cued object into an attended state as revealed by EEG decoding, oscillatory α-power, gaze bias, and improved orientation recall performance. Together, the results show that attentional selection spreads between an object's features in WM, consistent with object-based attentional selection. Analyses of neural processing at recall revealed that the selected object was automatically compared with the probe, whether it was the target for recall or not. This provides a potential mechanism for the observed benefits of nonpredictive cueing in WM, where a selected item is prioritized for subsequent decision-making. |
Estelle Raffin; Adrien Witon; Roberto F. Salamanca-Giron; Krystel R. Huxlin; Friedhelm C. Hummel Functional segregation within the dorsal frontoparietal network: A multimodal dynamic causal modeling study Journal Article In: Cerebral Cortex, vol. 32, no. 15, pp. 3187–3205, 2022. @article{Raffin2022, Discrimination and integration of motion direction requires the interplay of multiple brain areas. Theoretical accounts of perception suggest that stimulus-related (i.e., exogenous) and decision-related (i.e., endogenous) factors affect distributed neuronal processing at different levels of the visual hierarchy. To test these predictions, we measured brain activity of healthy participants during a motion discrimination task, using electroencephalography (EEG) and functional magnetic resonance imaging (fMRI). We independently modeled the impact of exogenous factors (task demand) and endogenous factors (perceptual decision-making) on the activity of the motion discrimination network and applied Dynamic Causal Modeling (DCM) to both modalities. DCM for event-related potentials (DCM-ERP) revealed that task demand impacted the reciprocal connections between the primary visual cortex (V1) and medial temporal areas (V5). With practice, higher visual areas were increasingly involved, as revealed by DCM-fMRI. Perceptual decision-making modulated higher levels (e.g., V5-to-Frontal Eye Fields, FEF), in a manner predictive of performance. Our data suggest that lower levels of the visual network support early, feature-based selection of responses, especially when learning strategies have not been implemented. In contrast, perceptual decision-making operates at higher levels of the visual hierarchy by integrating sensory information with the internal state of the subject. |
Kati Roesmann; Ida Wessing; Sophia Kraß; Elisabeth J. Leehr; Tim Klucken; Thomas Straube; Markus Junghöfer Developmental aspects of fear generalization – A MEG study on neurocognitive correlates in adolescents versus adults Journal Article In: Developmental Cognitive Neuroscience, vol. 58, pp. 1–12, 2022. @article{Roesmann2022, Background: Fear generalization is pivotal for the survival-promoting avoidance of potential danger, but, if too pronounced, it promotes pathological anxiety. Similar to adult patients with anxiety disorders, healthy children tend to show overgeneralized fear responses. Objective: This study aims to investigate neuro-developmental aspects of fear generalization in adolescence – a critical age for the development of anxiety disorders. Methods: We compared healthy adolescents (14–17 years) with healthy adults (19–34 years) regarding their fear responses towards tilted Gabor gratings (conditioned stimuli, CS; and slightly differently titled generalization stimuli, GS). In the conditioning phase, CS were paired (CS+) or remained unpaired (CS-) with an aversive stimulus (unconditioned stimuli, US). In the test phase, behavioral, peripheral and neural responses to CS and GS were captured by fear- and UCS expectancy ratings, a perceptual discrimination task, pupil dilation and source estimations of event-related magnetic fields. Results: Closely resembling adults, adolescents showed robust generalization gradients of fear ratings, pupil dilation, and estimated neural source activity. However, in the UCS expectancy ratings, adolescents revealed shallower generalization gradients indicating overgeneralization. Moreover, adolescents showed stronger visual cortical activity after as compared to before conditioning to all stimuli. Conclusion: Various aspects of fear learning and generalization appear to be mature in healthy adolescents. Yet, cognitive aspects might show a slower course of development. |
Amirsaman Sajad; Steven P. Errington; Jeffrey D. Schall Functional architecture of executive control and associated event-related potentials in macaques Journal Article In: Nature Communications, vol. 13, no. 1, pp. 1–19, 2022. @article{Sajad2022, The medial frontal cortex (MFC) enables executive control by monitoring relevant information and using it to adapt behavior. In macaques performing a saccade countermanding (stop-signal) task, we simultaneously recorded electrical potentials over MFC and neural spiking across all layers of the supplementary eye field (SEF). We report the laminar organization of neurons enabling executive control by monitoring the conflict between incompatible responses, the timing of events, and sustaining goal maintenance. These neurons were a mix of narrow-spiking and broad-spiking found in all layers, but those predicting the duration of control and sustaining the task goal until the release of operant control were more commonly narrow-spiking neurons confined to layers 2 and 3 (L2/3). We complement these results with evidence for a monkey homolog of the N2/P3 event-related potential (ERP) complex associated with response inhibition. N2 polarization varied with error-likelihood and P3 polarization varied with the duration of expected control. The amplitude of the N2 and P3 were predicted by the spike rate of different classes of neurons located in L2/3 but not L5/6. These findings reveal features of the cortical microcircuitry supporting executive control and producing associated ERPs. |
René Michel; Laura Dugué; Niko A. Busch Distinct contributions of alpha and theta rhythms to perceptual and attentional sampling Journal Article In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3025–3039, 2022. @article{Michel2022, Accumulating evidence suggests that visual perception operates in an oscillatory fashion at an alpha frequency (around 10 Hz). Moreover, visual attention also seems to operate rhythmically, albeit at a theta frequency (around 5 Hz). Both rhythms are often associated to "perceptual snapshots" taken at the favorable phases of these rhythms. However, less is known about the unfavorable phases: do they constitute "blind gaps," requiring the observer to guess, or is information sampled with reduced precision insufficient for the task demands? As simple detection or discrimination tasks cannot distinguish these options, we applied a continuous report task by asking for the exact orientation of a Landolt ring's gap to estimate separate model parameters for precision and the amount of guessing. We embedded this task in a well-established psychophysical protocol by densely sampling such reports across 20 cue-target stimulus onset asynchronies in a Posner-like cueing paradigm manipulating involuntary spatial attention. Testing the resulting time courses of the guessing and precision parameters for rhythmicities using a fast Fourier transform, we found an alpha rhythm (9.6 Hz) in precision for invalidly cued trials and a theta rhythm (4.8 Hz) in the guess rate across validity conditions. These results suggest distinct roles of the perceptual alpha and the attentional theta rhythm. We speculate that both rhythms result in environmental sampling characterized by fluctuating spatial resolution, speaking against a strict succession of blind gaps and perceptual snapshots. |
Sara Milligan; Martín Antúnez; Horacio A. Barber; Elizabeth R. Schotter Are eye movements and EEG on the same page?: A coregistration study on parafoveal preview and lexical frequency Journal Article In: Journal of Experimental Psychology: General, pp. 1–23, 2022. @article{Milligan2022, Readers extract visual and linguistic information not only from fixated words but also upcoming parafoveal words to introduce new input efficiently into the language processing pipeline. The lexical frequency of upcoming words and similarity with subsequent foveal information both influence the amount of time people spend once they fixate the word foveally. However, it is unclear from eye movements alone the extent to which parafoveal word processing, and the integration of that word with foveally obtained information, continues after saccade plans have been initiated. To investigate the underlying neural processes involved in word recognition after saccade planning, we coregistered electroencephalogram (EEG) and eye movements during a gaze-contingent display change paradigm. We orthogonally manipulated the frequency of the parafoveal and foveal words and measured fixation related potentials (FRPs) upon foveal fixation. Eye movements showed primarily an effect of preview frequency, suggesting that saccade planning is based on the familiarity of the parafoveal input. FRPs, on the other hand, demonstrated a disruption in downstream processing when parafoveal and foveal input differed, but only when the parafoveal word was high frequency. These findings demonstrate that lexical processing continues after the eyes have moved away from a word and that eye movements and FRPs provide distinct but complementary accounts about oculomotor behavior and neural processing that cannot be obtained from either method in isolation. Furthermore, these findings put constraints on models of reading by suggesting that lexical processes that occur before an eye movement program is initiated are qualitatively different from those that occur afterward. |
Michael T. Miuccio; Gregory J. Zelinsky; Joseph Schmidt Are all real-world objects created equal? Estimating the “set-size” of the search target in visual working memory Journal Article In: Psychophysiology, vol. 59, pp. 1–13, 2022. @article{Miuccio2022, Are all real-world objects created equal? Visual search difficulty increases with the number of targets and as target-related visual working memory (VWM) load increases. Our goal was to investigate the load imposed by individual real-world objects held in VWM in the context of search. Measures of visual clutter attempt to quantify real-world set-size in the context of scenes. We applied one of these measures, the number of proto-objects, to individual real-world objects and used contralateral delay activity (CDA) to measure the resulting VWM load. The current study presented a real-world object as a target cue, followed by a delay where CDA was measured. This was followed by a four-object search array. We compared CDA and later search performance from target cues containing a high or low number of proto-objects. High proto-object target cues resulted in greater CDA, longer search RTs, target dwell times, and reduced search guidance, relative to low proto-object targets. These findings demonstrate that targets with more proto-objects result in a higher VWM load and reduced search performance. This shows that the number of proto-objects contained within individual objects produce set-size like effects in VWM and suggests proto-objects may be a viable unit of measure of real-world VWM load. Importantly, this demonstrates that not all real-world objects are created equal. |
Dinavahi V. P. S. Murty; Keerthana Manikandan; Wupadrasta Santosh Kumar; Ranjini Garani Ramesh; Simran Purokayastha; Bhargavi Nagendra; M. L. Abhishek; Aditi Balakrishnan; Mahendra Javali; Naren Prahalada Rao; Supratim Ray Stimulus-induced gamma rhythms are weaker in human elderly with mild cognitive impairment and Alzheimer's disease Journal Article In: Bio-protocol, vol. 12, no. 7, pp. 1–13, 2022. @article{Murty2022, Stimulus-induced narrow-band gamma oscillations (20–70 Hz) are induced in the visual areas of the brain when particular visual stimuli, such as bars, gratings, or full-screen hue, are shown to the subject. Such oscillations are modulated by higher cognitive functions, like attention, and working memory, and have been shown to be abnormal in certain neuropsychiatric disorders, such as schizophrenia, autism, and Alzheimer's disease. However, although electroencephalogram (EEG) remains one of the most non-invasive, inexpensive, and accessible methods to record brain signals, some studies have failed to observe discernable gamma oscillations in human EEG. In this manuscript, we have described in detail a protocol to elicit robust gamma oscillations in human EEG. We believe that our protocol could help in developing non-invasive gamma-based biomarkers in human EEG, for the early detection of neuropsychiatric disorders. |
Adam J. Naples; Jennifer H. Foss-Feig; Julie M. Wolf; Vinod H. Srihari; James C. McPartland Predictability modulates neural response to eye contact in ASD Journal Article In: Molecular Autism, vol. 13, no. 42, pp. 1–12, 2022. @article{Naples2022, Background: Deficits in establishing and maintaining eye-contact are early and persistent vulnerabilities of autism spectrum disorder (ASD), and the neural bases of these deficits remain elusive. A promising hypothesis is that social features of autism may reflect difficulties in making predictions about the social world under conditions of uncertainty. However, no research in ASD has examined how predictability impacts the neural processing of eye-contact in naturalistic interpersonal interactions. Method: We used eye tracking to facilitate an interactive social simulation wherein onscreen faces would establish eye-contact when the participant looked at them. In Experiment One, receipt of eye-contact was unpredictable; in Experiment Two, receipt of eye-contact was predictable. Neural response to eye-contact was measured via the N170 and P300 event-related potentials (ERPs). Experiment One included 23 ASD and 46 typically developing (TD) adult participants. Experiment Two included 25 ASD and 43 TD adult participants. Results: When receipt of eye-contact was unpredictable, individuals with ASD showed increased N170 and increased, but non-specific, P300 responses. The magnitude of the N170 responses correlated with measures of sensory and anxiety symptomology, such that increased response to eye-contact was associated with increased symptomology. However, when receipt of eye-contact was predictable, individuals with ASD, relative to controls, exhibited slower N170s and no differences in the amplitude of N170 or P300. Limitations: Our ASD sample was composed of adults with IQ > 70 and included only four autistic women. Thus, further research is needed to evaluate how these results generalize across the spectrum of age, sex, and cognitive ability. Additionally, as analyses were exploratory, some findings failed to survive false-discovery rate adjustment. Conclusions: Neural response to eye-contact in ASD ranged from attenuated to hypersensitive depending on the predictability of the social context. These findings suggest that the vulnerabilities in eye-contact during social interactions in ASD may arise from differences in anticipation and expectation of eye-contact in addition to the perception of gaze alone. |
Ádám Nárai; Zsuzsanna Nemecz; Zoltán Vidnyánszky; Béla Weiss Lateralization of orthographic processing in fixed-gaze and natural reading conditions Journal Article In: Cortex, vol. 157, pp. 99–116, 2022. @article{Narai2022, Lateralized processing of orthographic information is a hallmark of proficient reading. However, how this finding obtained for fixed-gaze processing of orthographic stimuli translates to ecologically valid reading conditions remained to be clarified. To address this shortcoming, here we assessed the lateralization of early orthographic processing in fixed-gaze and natural reading conditions using concurrent eye-tracking and EEG data recorded from young adults without reading difficulties. Sensor-space analyses confirmed the well-known left-lateralized negative-going deflection of fixed-gaze EEG activity throughout the period of early orthographic processing. At the same time, fixation-related EEG activity exhibited left-lateralized followed by right-lateralized processing of text stimuli during natural reading. A strong positive relationship was found between the early leftward lateralization in fixed-gaze and natural reading conditions. Using source-space analyses, early left-lateralized brain activity was obtained in lateraloccipital and posterior ventral occipito-temporal cortices reflecting letter-level processing in both conditions. In addition, in the same time interval, left-lateralized source activity was found also in premotor and parietal brain regions during natural reading. While brain activity remained left-lateralized in later stages representing word-level processing in posterior and middle ventral temporal regions in the fixed-gaze condition, fixation-related source activity became stronger in the right hemisphere in medial and more anterior ventral temporal brain regions indicating higher-level processing of orthographic information. Although our results show a strong positive relationship between the lateralization of letter-level processing in the two reading modes and suggest lateralized brain activity as a general marker for processing of orthographic information, they also clearly indicate the need for reading research in ecologically valid conditions to identify the neural basis of visuospatial attentional, oculomotor and higher-level processes specific to natural reading. |
Nir Ofir; Ayelet N. Landau Neural signatures of evidence accumulation in temporal decisions Journal Article In: Current Biology, vol. 32, no. 18, pp. 4093–4100, 2022. @article{Ofir2022, Cognitive models of interval timing can be formulated as an accumulation-to-bound process.1–5 However, the physiological manifestation of such processes has not yet been identified. We used electroencephalography (EEG) to measure the neural responses of participants while they performed a temporal bisection task in which they were requested to categorize the duration of visual stimuli as short or long.6 We found that the stimulus-offset and response-locked activity depends on both stimulus duration and the participants' decision. To relate this activity to the underlying cognitive processes, we used a drift-diffusion model.7 The model includes a noisy accumulator starting with the stimulus onset and a decision threshold. According to the model, a stimulus duration will be categorized as “long” if the accumulator reaches the threshold during stimulus presentation. Otherwise, it will be categorized as “short.” We found that at the offset of stimulus presentation, an EEG response marks the distance of the accumulator from the threshold. Therefore, this model offers an accurate description of our behavioral data as well as the EEG response using the same two model parameters. We then replicated this finding in an identical experiment conducted in the tactile domain. We also extended this finding to two different temporal ranges (sub- and supra-second). Taken together, the work provides a new way to study the cognitive processes underlying temporal decisions, using a combination of behavior, EEG, and modeling. |
Sergio Osorio; Martín Irani; Javiera Herrada; Francisco Aboitiz Neural responses to sensory novelty with and without conscious access Journal Article In: NeuroImage, vol. 262, pp. 1–12, 2022. @article{Osorio2022, Detection of novel stimuli that violate statistical regularities in the sensory scene is of paramount importance for the survival of biological organisms. Event-related potentials, phasic increases in pupil size, and evoked changes in oscillatory power have been proposed as markers of sensory novelty detection. However, how conscious access to novelty modulates these different brain responses is not well understood. Here, we studied the neural responses to sensory novelty in the auditory modality with and without conscious access. We identified individual thresholds for conscious auditory discrimination and presented to our participants sequences of tones, where the last stimulus could be another standard, a subthreshold target or a suprathreshold target. Participants were instructed to report whether the last tone of each sequence was the same or different from those preceding it. Results indicate that attentional orientation to behaviorally relevant stimuli and overt decision-making mechanisms, indexed by the P3 event-related response and reaction times, best predict whether a novel stimulus will be consciously accessed. Theta power and pupil size do not predict conscious access to novelty, but instead reflect information maintenance and unexpected sensory uncertainty. These results highlight the interplay between bottom-up and top-down mechanisms and how the brain weights neural responses to novelty and uncertainty during perception and goal-directed behavior. |
Andreas Alexandersen; Gábor Csifcsák; Josephine Groot; Matthias Mittner In: Neuroimage: Reports, vol. 2, no. 3, pp. 1–19, 2022. @article{Alexandersen2022, Mind wandering (MW) is a mental phenomenon humans experience daily. Yet, we lack a complete understanding of the neural basis of this pervasive mental state. Over the past decade there has been an increase in publications using transcranial direct current stimulation (tDCS) to modulate the propensity to mind wander, but findings are diverse, and a satisfactory conclusion is missing. Recently, Boayue et al. (2020) reported successful reduction of mind wandering using high-definition tDCS (HD-tDCS) over the dorsolateral prefrontal cortex, providing preliminary evidence for the efficacy of HD-tDCS in interfering with mind wandering. The current study is a high-powered, pre-registered direct replication attempt of the effect found by Boayue et al. (2020). In addition, we investigated whether the effects of HD-tDCS on mind wandering would be prolonged and assessed the underlying processes of mind wandering using electroencephalography (EEG) and pupillometry during a finger-tapping random sequence generation task that requires the use of executive resources. We failed to find any evidence of the original effect of reduced MW during and after stimulation. When combining our data with the data from Boayue et al. (2020), the original effect of reduced MW caused by HD-tDCS disappeared. In addition, we observed increased occipital alpha power as task duration increased and increased midfrontal theta power preceding response patterns signaling high executive function use. Finally, tonic and phasic pupil size decreased as task duration increased yet, phasic responses were increased, while tonic responses were reduced preceding reports of MW. Additionally phasic pupil size also showed a tendency to be increased during periods of high executive function use. Importantly, none of the EEG or pupil measures were modulated by HD-tDCS. We conclude that HD-tDCS over the dorsolateral prefrontal cortex does not affect MW propensity and its neural signatures. Furthermore, we recommend that previously reported effects of tDCS on mind wandering and other cognitive functions should only be accepted after a successful pre-registered replication. |
Kristijan Armeni; Umut Güçlü; Marcel Gerven; Jan-Mathijs Schoffelen A 10-hour within-participant magnetoencephalography narrative dataset to test models of language comprehension Journal Article In: Scientific Data, vol. 9, pp. 1–18, 2022. @article{Armeni2022, Recently, cognitive neuroscientists have increasingly studied the brain responses to narratives. At the same time, we are witnessing exciting developments in natural language processing where large-scale neural network models can be used to instantiate cognitive hypotheses in narrative processing. Yet, they learn from text alone and we lack ways of incorporating biological constraints during training. To mitigate this gap, we provide a narrative comprehension magnetoencephalography (MEG) data resource that can be used to train neural network models directly on brain data. We recorded from 3 participants, 10 separate recording hour-long sessions each, while they listened to audiobooks in English. After story listening, participants answered short questions about their experience. To minimize head movement, the participants wore MEG-compatible head casts, which immobilized their head position during recording. We report a basic evoked-response analysis showing that the responses accurately localize to primary auditory areas. The responses are robust and conserved across 10 sessions for every participant. We also provide usage notes and briefly outline possible future uses of the resource. |
Alessandro Benedetto; Hao Tam Ho; Maria Concetta Morrone The readiness potential correlates with action-linked modulation of visual accuracy Journal Article In: Eneuro, pp. 1–17, 2022. @article{Benedetto2022, Visual accuracy is consistently shown to be modulated around the time of the action execution. The neural underpinning of this motor-induced modulation of visual perception is still unclear. Here, we investigate with EEG whether it is related to the readiness potential, an event-related potential linked to motor preparation. Across 18 human participants, the magnitude of visual modulation following a voluntary button press was found to correlate with the readiness potential amplitude measured during visual discrimination. Participants' amplitude of the readiness potential in a purely motor-task was also found to correlate with the extent of the motor-induced modulation of visual perception in the visuomotor task. These results provide strong evidence that perceptual changes close to action execution are associated with motor preparation processes and that this mechanism is independent of task contingencies. Further, our findings suggest that the readiness potential provides a fingerprint of individual visuomotor interaction. |
Sven Braeutigam; Jessica Clare Scaife; Tipu Aziz; Rebecca J. Park A longitudinal magnetoencephalographic study of the effects of deep brain stimulation on neuronal dynamics in severe anorexia nervosa Journal Article In: Frontiers in Behavioral Neuroscience, vol. 16, pp. 1–12, 2022. @article{Braeutigam2022, Anorexia Nervosa (AN) is a debilitating psychiatric disorder characterized by the relentless pursuit of thinness, leading to severe emaciation. Magnetoencephalography (MEG)was used to record the neuronal response in seven patients with treatment-resistant AN while completing a disorder-relevant food wanting task. The patients underwent a 15-month protocol, where MEG scans were conducted pre-operatively, post-operatively prior to deep brain stimulation (DBS) switch on, twice during a blind on/off month and at protocol end. Electrodes were implanted bilaterally into the nucleus accumbens with stimulation at the anterior limb of the internal capsule using rechargeable implantable pulse generators. Three patients met criteria as responders at 12 months of stimulation, showing reductions of eating disorder psychopathology of over 35%. An increase in alpha power, as well as evoked power at latencies typically associated with visual processing, working memory, and contextual integration was observed in ON compared to OFF sessions across all seven patients. Moreover, an increase in evoked power at P600-like latencies as well as an increase in γ-band phase-locking over anterior-to-posterior regions were observed for high- compared to low-calorie food image only in ON sessions. These findings indicate that DBS modulates neuronal process in regions far outside the stimulation target site and at latencies possibly reflecting task specific processing, thereby providing further evidence that deep brain stimulation can play a role in the treatment of otherwise intractable psychiatric disorders. |
Marion Brickwedde; Yulia Bezsudnova; Anna Kowalczyk; Ole Jensen; Alexander Zhigalov Application of rapid invisible frequency tagging for brain computer interfaces Journal Article In: Journal of Neuroscience Methods, vol. 382, pp. 1–9, 2022. @article{Brickwedde2022, Background: Brain-computer interfaces (BCI) based on steady-state visual evoked potentials (SSVEPs/SSVEFs) are among the most commonly used BCI systems. They require participants to covertly attend to visual objects flickering at specified frequencies. The attended location is decoded online by analysing the power of neuronal responses at the flicker frequency. New method: We implemented a novel rapid invisible frequency-tagging technique, utilizing a state-of-the-art projector with refresh rates of up to 1440 Hz. We flickered the luminance of visual objects at 56 and 60 Hz, which was invisible to participants but produced strong neuronal responses measurable with magnetoencephalography (MEG). The direction of covert attention, decoded from frequency-tagging responses, was used to control an online BCI PONG game. Results: Our results show that seven out of eight participants were able to play the pong game controlled by the frequency-tagging signal, with average accuracies exceeding 60 %. Importantly, participants were able to modulate the power of the frequency-tagging response within a 1-second interval, while only seven occipital sensors were required to reliably decode the neuronal response. Comparison with existing methods: In contrast to existing SSVEP-based BCI systems, rapid frequency-tagging does not produce a visible flicker. This extends the time-period participants can use it without fatigue, by avoiding distracting visual input. Furthermore, higher frequencies increase the temporal resolution of decoding, resulting in higher communication rates. Conclusion: Using rapid invisible frequency-tagging opens new avenues for fundamental research and practical applications. In combination with novel optically pumped magnetometers (OPMs), it could facilitate the development of high-speed and mobile next-generation BCI systems. |
Maximilian Bruchmann; Sebastian Schindler; Mandana Dinyarian; Thomas Straube The role of phase and orientation for ERP modulations of spectrum-manipulated fearful and neutral faces Journal Article In: Psychophysiology, vol. 59, no. 3, pp. 1–13, 2022. @article{Bruchmann2022, Prioritized processing of fearful compared to neutral faces has been proposed to result from evolutionary adaptation of the contrast sensitivity function (CSF) to the features of emotionally relevant faces and/or vice versa. However, it is unknown whether a stimulus merely has to feature the amplitude spectrum of a fearful face to be prioritized or whether the relevant spatial frequencies have to occur with specific phases and orientations. Prioritized processing is indexed by specific increases of Event-Related Potentials (ERPs) of the EEG and occurs throughout different early processing stages, indexed by emotion-related modulations of the P1, N170, and EPN. In this pre-registered study, we manipulated phase and amplitude properties of the Fourier spectra of neutral and fearful faces to test the effect of phase coherence (PC, face vs. scramble) and orientation coherence (OC, original vs. rotational average) and their interactions with differential emotion processing. We found that differential emotion processing was not present at the level of P1 but strongly affected N170 and EPN. In both cases, intact phase coherence was required for enhanced processing of fearful faces. OC did not interact with emotion. While faces produced the typical N170 effect, we observed a reversed effect for scrambles. Additional exploratory independent component analysis (ICA) suggests that this reversal could signal a mismatch between an early "perceptual hypothesis" and feedback of configural information. In line with our expectations, fearful-neutral differences for the N170 and EPN depend on configural information, i.e., recognizable faces. |
Sebastian Schindler; Niko Busch; Maximilian Bruchmann; Maren-Isabel Wolf; Thomas Straube Early ERP functions are indexed by lateralized effects to peripherally presented emotional faces and scrambles Journal Article In: Psychophysiology, vol. 59, no. 2, pp. 1–16, 2022. @article{Schindler2022b, A large body of research suggests that early event-related potentials (ERPs), such as the P1 and N1, are potentiated by attention and represent stimulus amplification. However, recent accounts suggest that the P1 is associated with inhibiting the irrelevant visual field evidenced by a pronounced ipsilateral P1 during sustained attention to peripherally presented stimuli. The current EEG study further investigated this issue to reveal how lateralized ERP findings are modulated by face and emotional information. Therefore, participants were asked to fixate the center of the screen and pay sustained attention either to the right or left visual field, where angry or neutral faces or their Fourier phase-scrambled versions were presented. We found a bilateral P1 to all stimuli with relatively increased, but delayed, ipsilateral P1 amplitudes to faces but not to scrambles. Explorative independent component analyses dissociated an earlier lateralized larger contralateral P1 from a later bilateral P1. By contrast, the N170 showed a contralateral enhancement to all stimuli, which was most pronounced for neutral faces attended in the left hemifield. Finally, increased contralateral alpha power was found for both attended hemifields but was not significantly related to poststimulus ERPs. These results provide evidence against a general inhibitory role of the P1 but suggest stimulus-specific relative enhancements of the ipsilateral P1 for the irrelevant visual hemifield. The lateralized N170, however, is associated with stimulus amplification as a function of facial features. |
Sebastian Schindler; Theresa Sofie Richter; Maximilian Bruchmann; Niko A. Busch; Thomas Straube Effects of task load, spatial attention, and trait anxiety on neuronal responses to fearful and neutral faces Journal Article In: Psychophysiology, vol. 59, pp. 1–14, 2022. @article{Schindler2022, There is an ongoing debate on how different components of the event-related potential (ERP) to threat-related facial expressions are modulated by attentional conditions and interindividual differences in trait anxiety. In the current study (N = 80), we examined ERPs to centrally presented, task-irrelevant fearful and neutral faces, while participants had to solve a face-unrelated visual task, which differed in difficulty and spatial position. Critically, we used a fixation-controlled experimental design and ensured the spatial attention manipulation by spectral analysis of the EEG. Besides the factors emotion, spatial attention, and perceptual load, we also investigated correlations between trait anxiety and ERPs. While P1 emotion effects were insignificant, the N170 was increased to fearful faces regardless of load and spatial attention conditions. During the EPN time window, a significantly increased negativity for fearful faces was observed only during low load and spatial attention to the face. We found no significant relationship between ERPs and trait anxiety, questioning the hypothesis of a general hypersensitivity toward fearful expressions in anxious individuals. These results show a high resistance of the N170 amplitude increase for fearful faces to spatial attention and task load manipulations. By contrast, the EPN modulation by fearful faces index a resource-dependent stage of the ERP, requiring both spatial attention at the location of faces and low load of the face-irrelevant task. |
Elena Selezneva; Nicole Wetzel The impact of probabilistic cues on sound-related pupil dilation and ERP responses in 7–9-year-old children Journal Article In: Auditory Perception & Cognition, vol. 5, no. 1-2, pp. 86–106, 2022. @article{Selezneva2022, Control of involuntary orienting of attention toward new but task-irrelevant events is essential to successfully perform a task. We investigated top-down control of involuntary orienting of attention caused by task-irrelevant novel sounds embedded in a sequence of repeated standard sounds in 7–9-year-old children (N = 30) and in an adult control group (N = 30). The type of sound was announced by visual cues, which were correct in 80% of the trials. We co-registered sound-related pupil dilation responses (PDR), the attention-related component P3a in the EEG and performance. Task-irrelevant novel sounds evoked increased amplitudes of the PDR and the P3a and prolonged reaction times in both age groups. In children only, invalidly cued novel sounds evoked larger PDR amplitudes than validly cued novel sounds, while this cue effect was not observed for standard sounds. In both age groups, P3a amplitudes in the centro-parietal region were reduced to the correctly cued compared to the incorrectly cued novel sounds, indicating top-down control of orienting of attention. The reaction time prolongation to both validly and invalidly cued novel sounds were similar in both age groups. These findings demonstrate that children are capable of reducing the orienting of attention and evaluation triggered by task-irrelevant sounds by using probabilistic cues. Children's pupil results indicate a high sensitivity of pupil dynamics to cue-related top-down influences on novel sound processing, emphasizing the utility of pupillometry in developmental research. |
Benjamin J. Stauch; Alina Peter; Isabelle Ehrlich; Zora Nolte; Pascal Fries Human visual gamma for color stimuli Journal Article In: eLife, vol. 11, pp. 1–18, 2022. @article{Stauch2022, Strong gamma-band oscillations in primate early visual cortex can be induced by homogeneous color surfaces (Peter et al., 2019; Shirhatti and Ray, 2018). Compared to other hues, particularly strong gamma oscillations have been reported for red stimuli. However, precortical color processing and the resultant strength of input to V1 have often not been fully controlled for. Therefore, stronger responses to red might be due to differences in V1 input strength. We presented stimuli that had equal luminance and cone contrast levels in a color coordinate system based on responses of the lateral geniculate nucleus, the main input source for area V1. With these stimuli, we recorded magnetoencephalography in 30 human participants. We found gamma oscillations in early visual cortex which, contrary to previous reports, did not differ between red and green stimuli of equal L-M cone contrast. Notably, blue stimuli with contrast exclusively on the S-cone axis induced very weak gamma responses, as well as smaller event-related fields and poorer change-detection performance. The strength of human color gamma responses for stimuli on the L-M axis could be well explained by L-M cone contrast and did not show a clear red bias when L-M cone contrast was properly equalized. |
Anna Lena Stroh; Konstantin Grin; Frank Rösler; Davide Bottari; José Ossandón; Bruno Rossion; Brigitte Röder Developmental experiences alter the temporal processing characteristics of the visual cortex: Evidence from deaf and hearing native signers Journal Article In: European Journal of Neuroscience, vol. 55, no. 6, pp. 1629–1644, 2022. @article{Stroh2022, To date, the extent to which early experience shapes the functional characteristics of neural circuits is still a matter of debate. In the present study, we tested whether congenital deafness and/or the acquisition of a sign language alter the temporal processing characteristics of the visual system. Moreover, we investigated whether, assuming cross-modal plasticity in deaf individuals, the temporal processing characteristics of possibly reorganised auditory areas resemble those of the visual cortex. Steady-state visual evoked potentials (SSVEPs) were recorded in congenitally deaf native signers, hearing native signers, and hearing nonsigners. The luminance of the visual stimuli was periodically modulated at 12, 21, and 40 Hz. For hearing nonsigners, the optimal driving rate was 12 Hz. By contrast, for the group of hearing signers, the optimal driving rate was 12 and 21 Hz, whereas for the group of deaf signers, the optimal driving rate was 21 Hz. We did not observe evidence for cross-modal recruitment of auditory cortex in the group of deaf signers. These results suggest a higher preferred neural processing rate as a consequence of the acquisition of a sign language. |
Lina Teichmann; Denise Moerel; Anina N. Rich; Chris I. Baker The nature of neural object representations during dynamic occlusion Journal Article In: Cortex, vol. 153, pp. 66–86, 2022. @article{Teichmann2022, Objects disappearing briefly from sight due to occlusion is an inevitable occurrence in everyday life. Yet we generally have a strong experience that occluded objects continue to exist, despite the fact that they objectively disappear. This indicates that neural object representations must be maintained during dynamic occlusion. However, it is unclear what the nature of such representation is and in particular whether it is perception-like or more abstract, for example, reflecting limited features such as position or movement direction only. In this study, we address this question by examining how different object features such as object shape, luminance, and position are represented in the brain when a moving object is dynamically occluded. We apply multivariate decoding methods to Magnetoencephalography (MEG) data to track how object representations unfold over time. Our methods allow us to contrast the representations of multiple object features during occlusion and enable us to compare the neural responses evoked by visible and occluded objects. The results show that object position information is represented during occlusion to a limited extent while object identity features are not maintained through the period of occlusion. Together, this suggests that the nature of object representations during dynamic occlusion is different from visual representations during perception. |
William Thyer; Kirsten C. S. Adam; Gisella K. Diaz; Itzel N. Velázquez Sánchez; Edward K. Vogel; Edward Awh Storage in visual working memory recruits a content-independent pointer system Journal Article In: Psychological Science, vol. 33, no. 10, pp. 1680–1694, 2022. @article{Thyer2022, Past work has shown that storage in working memory elicits stimulus-specific neural activity that tracks the stored content. Here, we present evidence for a distinct class of load-sensitive neural activity that indexes items without representing their contents per se. We recorded electroencephalogram (EEG) activity while adult human subjects stored varying numbers of items in visual working memory. Multivariate analysis of the scalp topography of EEG voltage enabled precise tracking of the number of individuated items stored and robustly predicted individual differences in working memory capacity. Critically, this signature of working memory load generalized across variations in both the type and number of visual features stored about each item, suggesting that it tracked the number of individuated memory representations and not the content of those memories. We hypothesize that these findings reflect the operation of a capacity-limited pointer system that supports on-line storage and attentive tracking. |
Anne E. Urai; Tobias H. Donner Persistent activity in human parietal cortex mediates perceptual choice repetition bias Journal Article In: Nature Communications, vol. 13, no. 1, pp. 1–15, 2022. @article{Urai2022, Humans and other animals tend to repeat or alternate their previous choices, even when judging sensory stimuli presented in a random sequence. It is unclear if and how sensory, associative, and motor cortical circuits produce these idiosyncratic behavioral biases. Here, we combined behavioral modeling of a visual perceptual decision with magnetoencephalographic (MEG) analyses of neural dynamics, across multiple regions of the human cerebral cortex. We identified distinct history-dependent neural signals in motor and posterior parietal cortex. Gamma-band activity in parietal cortex tracked previous choices in a sustained fashion, and biased evidence accumulation toward choice repetition; sustained beta-band activity in motor cortex inversely reflected the previous motor action, and biased the accumulation starting point toward alternation. The parietal, not motor, signal mediated the impact of previous on current choice and reflected individual differences in choice repetition. In sum, parietal cortical signals seem to play a key role in shaping choice sequences. |
Mats W. J. Es; Tom R. Marshall; Eelke Spaak; Ole Jensen; Jan-Mathijs Schoffelen Phasic modulation of visual representations during sustained attention Journal Article In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3191–3208, 2022. @article{Es2022, Sustained attention has long been thought to benefit perception in a continuous fashion, but recent evidence suggests that it affects perception in a discrete, rhythmic way. Periodic fluctuations in behavioral performance over time, and modulations of behavioral performance by the phase of spontaneous oscillatory brain activity point to an attentional sampling rate in the theta or alpha frequency range. We investigated whether such discrete sampling by attention is reflected in periodic fluctuations in the decodability of visual stimulus orientation from magnetoencephalographic (MEG) brain signals. In this exploratory study, human subjects attended one of the two grating stimuli, while MEG was being recorded. We assessed the strength of the visual representation of the attended stimulus using a support vector machine (SVM) to decode the orientation of the grating (clockwise vs. counterclockwise) from the MEG signal. We tested whether decoder performance depended on the theta/alpha phase of local brain activity. While the phase of ongoing activity in the visual cortex did not modulate decoding performance, theta/alpha phase of activity in the frontal eye fields and parietal cortex, contralateral to the attended stimulus did modulate decoding performance. These findings suggest that phasic modulations of visual stimulus representations in the brain are caused by frequency-specific top-down activity in the frontoparietal attention network, though the behavioral relevance of these effects could not be established. |
Prateek Dhamija; Allison Wong; Asaf Gilboa Early auditory event related potentials distinguish higher-order from first-order aversive conditioning Journal Article In: Frontiers in Behavioral Neuroscience, vol. 16, pp. 1–14, 2022. @article{Dhamija2022, Stimuli in reality rarely co-occur with primary reward or punishment to allow direct associative learning of value. Instead, value is thought to be inferred through complex higher-order associations. Rodent research has demonstrated that the formation and maintenance of first-order and higher-order associations are supported by distinct neural substrates. In this study, we explored whether this pattern of findings held true for humans. Participants underwent first-order and subsequent higher-order conditioning using an aversive burst of white noise or neutral tone as the unconditioned stimuli. Four distinct tones, initially neutral, served as first-order and higher-order conditioned stimuli. Autonomic and neural responses were indexed by pupillometry and evoked response potentials (ERPs) respectively. Conditioned aversive values of first-order and higher-order stimuli led to increased autonomic responses, as indexed by pupil dilation. Distinct temporo-spatial auditory evoked response potentials were elicited by first-order and high-order conditioned stimuli. Conditioned first-order responses peaked around 260 ms and source estimation suggested a primary medial prefrontal and amygdala source. Conversely, conditioned higher-order responses peaked around 120 ms with an estimated source in the medial temporal lobe. Interestingly, pupillometry responses to first-order conditioned stimuli were diminished after higher order training, possibly signifying concomitant incidental extinction, while responses to higher-order stimuli remained. This suggests that once formed, higher order associations are at least partially independent of first order conditioned representations. This experiment demonstrates that first-order and higher-order conditioned associations have distinct neural signatures, and like rodents, the medial temporal lobe may be specifically involved with higher-order conditioning. |
Marwa El Zein; Ray J. Dolan; Bahador Bahrami Shared responsibility decreases the sense of agency in the human brain Journal Article In: Journal of Cognitive Neuroscience, vol. 34, pp. 2065–2081, 2022. @article{ElZein2022, Sharing responsibility in social decision-making helps individuals use the flexibility of the collective context to benefit them-selves by claiming credit for good outcomes or avoiding the blame for bad outcomes. Using magnetoencephalography, we examined the neuronal basis of the impact that social context has on this flexible sense of responsibility. Participants performed a gambling task in various social contexts and reported feeling less responsibility when playing as a member of a team. A reduced magnetoencephalography outcome processing effect was observed as a function of decreasing responsibility at 200 msec post outcome onset and was centered over parietal, central, and frontal brain regions. Before outcome revelation in socially made decisions, an attenuated motor preparation signature at 500 msec after stimulus onset was found. A boost in reported responsibility for positive outcomes in social contexts was associated with increased activity in regions related to social and reward processing. Together, these results show that sharing responsibility with others reduces agency, influencing pre-outcome motor preparation and post-outcome processing, and provides opportunities to flexibly claim credit for positive outcomes. |
Hesham A. Elshafei; Corinne Orlemann; Saskia Haegens The impact of eye closure on anticipatory α activity in a tactile discrimination task Journal Article In: eNeuro, vol. 9, no. 1, pp. 1–11, 2022. @article{Elshafei2022, One of the very first observations made regarding α oscillations (8–14 Hz), is that they increase in power over posterior areas when awake participants close their eyes. Recent work, especially in the context of (spatial) attention, suggests that α activity reflects a mechanism of functional inhibition. However, it remains unclear how eye closure impacts anticipatory α modulation observed in attention paradigms, and how this affects subsequent behavioral performance. Here, we recorded magnetoencephalography (MEG) in 33 human participants performing a tactile discrimination task with their eyes open versus closed. We replicated the hallmarks of previous somatosensory spatial attention studies: α lateralization across the somatosensory cortices as well as α increase over posterior (visual) regions. Furthermore, we found that eye closure leads to (1) reduced task performance; (2) widespread increase in α power; and (3) reduced anticipatory visual α modulation (4) with no effect on somatosensory α lateralization. Regardless of whether participants had their eyes open or closed, increased visual α power and somatosensory α lateralization improved their performance. Thus, we provide evidence that eye closure does not alter the impact of anticipatory α modulations on behavioral performance. We propose there is an optimal visual α level for somatosensory task performance, which can be achieved through a combination of eye closure and top-down anticipatory attention. |
João Estiveira; Camila Dias; Diana Costa; João Castelhano; Miguel Castelo-Branco; Teresa Sousa An action-independent role for midfrontal theta activity prior to error commission Journal Article In: Frontiers in Human Neuroscience, vol. 16, pp. 1–17, 2022. @article{Estiveira2022, Error-related electroencephalographic (EEG) signals have been widely studied concerning the human cognitive capability of differentiating between erroneous and correct actions. Midfrontal error-related negativity (ERN) and theta band oscillations are believed to underlie post-action error monitoring. However, it remains elusive how early monitoring activity is trackable and what are the pre-response brain mechanisms related to performance monitoring. Moreover, it is still unclear how task-specific parameters, such as cognitive demand or motor control, influence these processes. Here, we aimed to test pre- and post-error EEG patterns for different types of motor responses and investigate the neuronal mechanisms leading to erroneous actions. We implemented a go/no-go paradigm based on keypresses and saccades. Participants received an initial instruction about the direction of response to be given based on a facial cue and a subsequent one about the type of action to be performed based on an object cue. The paradigm was tested in 20 healthy volunteers combining EEG and eye tracking. We found significant differences in reaction time, number, and type of errors between the two actions. Saccadic responses reflected a higher number of premature responses and errors compared to the keypress ones. Nevertheless, both led to similar EEG patterns, supporting previous evidence for increased ERN amplitude and midfrontal theta power during error commission. Moreover, we found pre-error decreased theta activity independent of the type of action. Source analysis suggested different origin for such pre- and post-error neuronal patterns, matching the anterior insular cortex and the anterior cingulate cortex, respectively. This opposite pattern supports previous evidence of midfrontal theta not only as a neuronal marker of error commission but also as a predictor of action performance. Midfrontal theta, mostly associated with alert mechanisms triggering behavioral adjustments, also seems to reflect pre-response attentional mechanisms independently of the action to be performed. Our findings also add to the discussion regarding how salience network nodes interact during performance monitoring by suggesting that pre- and post-error patterns have different neuronal sources within this network. |
John J. Foxe; Emily J. Knight; Evan J. Myers; Cody Zhewei Cao; Sophie Molholm; Edward G. Freedman The strength of feedback processing is associated with resistance to visual backward masking during Illusory Contour processing in adult humans Journal Article In: NeuroImage, vol. 259, pp. 1–10, 2022. @article{Foxe2022, Re-entrant feedback processing is a key mechanism of visual object-recognition, especially under compromised viewing conditions where only sparse information is available and object features must be interpolated. Illusory Contour stimuli are commonly used in conjunction with Visual Evoked Potentials (VEP) to study these filling-in processes, with characteristic modulation of the VEP in the ∼100-150 ms timeframe associated with this re-entrant processing. Substantial inter-individual variability in timing and amplitude of feedback-related VEP modulation is observed, raising the question whether this variability might underlie inter-individual differences in the ability to form strong perceptual gestalts. Backward masking paradig ms have been used to study inter-individual variance in the ability to form robust object perceptions before processing of the mask interferes with object-recognition. Some individuals recognize objects when the time between target object and mask is extremely short, whereas others struggle to do so even at longer target-to-mask intervals. We asked whether timing and amplitude of feedback-related VEP modulations were associated with individual differences in resistance to backward masking. Participants (N=40) showed substantial performance variability in detecting Illusory Contours at intermediate target-to-mask intervals (67 ms and 117 ms), allowing us to use kmeans clustering to divide the population into four performance groups (poor, low-average, high-average, superior). There was a clear relationship between the amplitude (but not the timing) of feedback-related VEP modulation and Illusory Contour detection during backward masking. We conclude that individual differences in the strength of feedback processing in neurotypical humans lead to differences in the ability to quickly establish perceptual awareness of incomplete visual objects. |
Anna L. Gert; Benedikt V. Ehinger; Silja Timm; Tim C. Kietzmann; Peter König WildLab: A naturalistic free viewing experiment reveals previously unknown electroencephalography signatures of face processing Journal Article In: European Journal of Neuroscience, vol. 56, pp. 6022–6038, 2022. @article{Gert2022, Neural mechanisms of face perception are predominantly studied in well-controlled experimental settings that involve random stimulus sequences and fixed eye positions. While powerful, the employed paradigms are far from what constitutes natural vision. Here, we demonstrate the feasibility of ecologically more valid experimental paradigms using natural viewing behavior, by combining a free viewing paradigm on natural scenes, free of photographer bias, with advanced data processing techniques that correct for overlap effects and co-varying nonlinear dependencies of multiple eye movement parameters. We validate this approach by replicating classic N170 effects in neural responses, triggered by fixation onsets (fERPs). Importantly, our more natural stimulus paradigm yielded smaller variability between subjects than the classic setup. Moving beyond classic temporal and spatial effect locations, our experiment furthermore revealed previously unknown signatures of face processing: This includes category-specific modulation of the event-related potential (ERP)'s amplitude even before fixation onset, as well as adaptation effects across subsequent fixations depending on their history. |
Erin Goddard; Thomas A. Carlson; Alexandra Woolgar Spatial and feature-selective attention have distinct, interacting effects on population-level tuning Journal Article In: Journal of Cognitive Neuroscience, vol. 34, no. 2, pp. 290–312, 2022. @article{Goddard2022, Attention can be deployed in different ways: When searching for a taxi in New York City, we can decide where to attend (e.g., to the street) and what to attend to (e.g., yellow cars). Although we use the same word to describe both processes, nonhuman primate data suggest that these produce distinct effects on neural tuning. This has been challenging to assess in humans, but here we used an opportunity afforded by multivariate decoding of MEG data. We found that attending to an object at a particular location and attending to a particular object feature produced effects that interacted multiplicatively. The two types of attention induced distinct patterns of enhancement in occipital cortex, with feature-selective attention producing relatively more enhancement of small feature differences and spatial attention producing relatively larger effects for larger feature differences. An information flow analysis further showed that stimulus representations in occipital cortex were Granger-caused by coding in frontal cortices earlier in time and that the timing of this feedback matched the onset of attention effects. The data suggest that spatial and feature-selective attention rely on distinct neural mechanisms that arise from frontal-occipital information exchange, interacting multiplicatively to selectively enhance task-relevant information. |
Mariana M. Gusso; Kate L. Christison-Lagay; David Zuckerman; Ganesh Chandrasekaran; Sharif I. Kronemer; Julia Z. Ding; Noah C. Freedman; Percy Nohama; Hal Blumenfeld More than a feeling: Scalp EEG and eye signals in conscious tactile perception Journal Article In: Consciousness and Cognition, vol. 105, pp. 1–14, 2022. @article{Gusso2022, Understanding the neural basis of consciousness is a fundamental goal of neuroscience, and sensory perception is often used as a proxy for consciousness in empirical studies. However, most studies rely on reported perception of visual stimuli. Here we present behavior, high density scalp EEG and eye metric recordings collected simultaneously during a novel tactile threshold perception task. We found significant N80, N140 and P300 event related potentials in perceived trials and in perceived versus not perceived trials. Significance was limited to a P100 and P300 in not perceived trials. We also found an increase in pupil diameter and blink rate and a decrease in microsaccade rate following perceived relative to not perceived tactile stimuli. These findings support the use of eye metrics as a measure of physiological arousal associated with conscious perception. Eye metrics may also represent a novel path toward the creation of tactile no-report tasks in the future. |
Tjerk P. Gutteling; Lonieke Sillekens; Nilli Lavie; Ole Jensen Alpha oscillations reflect suppression of distractors with increased perceptual load Journal Article In: Progress in Neurobiology, vol. 214, pp. 1–13, 2022. @article{Gutteling2022, Attention serves an essential role in cognition and behavior allowing us to focus on behaviorally-relevant objects while ignoring distraction. Perceptual load theory states that attentional resources are allocated according to the requirements of the task, i.e., its ‘load'. The theory predicts that the resources left to process irrelevant, possibly distracting stimuli, are reduced when the perceptual load is high. However, it remains unclear how this allocation of attentional resources specifically relates to neural excitability and suppression mechanisms. In this magnetoencephalography (MEG) study, we show that brain oscillations in the alpha band (8–13 Hz) implemented the suppression of distracting objects when the perceptual load was high. In parallel, high load increased the neuronal excitability for target objects, as reflected by rapid invisible frequency tagging. We suggest that the allocation of resources in tasks with high perceptual load is implemented by a gain increase for targets, complemented by distractor suppression reflected by alpha-band oscillations closing the ‘gate' for interference. |
Qing He; Xin Yue Yang; Baoqi Gong; Keyan Bi; Fang Fang Boosting visual perceptual learning by transcranial alternating current stimulation over the visual cortex at alpha frequency Journal Article In: Brain Stimulation, vol. 15, no. 3, pp. 546–553, 2022. @article{He2022b, Background: Transcranial alternating current stimulation (tACS) has been widely used to alter ongoing brain rhythms in a frequency-specific manner to modulate relevant cognitive functions, including visual functions. Therefore, it is a useful tool for exploring the causal role of neural oscillations in cognition. Visual functions can be improved substantially by training, which is called visual perceptual learning (VPL). However, whether and how tACS can modulate VPL is still unclear. Objective: This work aims to explore how tACS modulates VPL and the role of neural oscillations in VPL. Methods: A between-subjects design was adopted. Subjects were assigned to six groups and undertook five daily training sessions to execute an orientation discrimination task. During training, five groups received occipital tACS stimulation at 6, 10, 20, 40, and sham 10 Hz respectively, and one group was stimulated at the sensorimotor regions by 10 Hz tACS. Results: Compared with the sham stimulation, occipital tACS at 10 Hz, but not at other frequencies, accelerated perceptual learning and increased the performance improvement. However, these modulatory effects were absent when 10 Hz tACS was delivered to the sensorimotor areas. Moreover, the tACS-induced performance improvement lasted at least two months after the end of training. Conclusion: TACS can facilitate orientation discrimination learning in a frequency- and location-specific manner. Our findings provide strong evidence for a pivotal role of alpha oscillations in boosting VPL and shed new light on the design of effective neuromodulation protocols that can facilitate rehabilitation for patients with neuro-ophthalmological disorders. |
Klaartje T. H. Heinen; J. Leon Kenemans; Stefan Stigchel Recruitment of a long-term memory supporting neural network during repeated maintenance of a multi-item abstract visual image in working memory Journal Article In: Scientific Reports, vol. 12, pp. 1–17, 2022. @article{Heinen2022, Humans can flexibly transfer information between different memory systems. Information in visual working memory (VWM) can for instance be stored in long-term memory (LTM). Conversely, information can be retrieved from LTM and temporarily held in WM when needed. It has previously been suggested that a neural transition from parietal- to midfrontal activity during repeated visual search reflects transfer of information from WM to LTM. Whether this neural transition indeed reflects consolidation and is also observed when memorizing a rich visual scene (rather than responding to a single target), is not known. To investigate this, we employed an EEG paradigm, in which abstract six-item colour-arrays were repeatedly memorized and explicitly visualized, or merely attended to. Importantly, we tested the functional significance of a potential neural shift for longer-term consolidation in a subsequent recognition task. Our results show a gradually enhanced- and sustained modulation of the midfrontal P170 component and a decline in parietal CDA, during repeated WM maintenance. Improved recollection/visualization of memoranda upon WM-cueing, was associated with contralateral parietal- and right temporal activity. Importantly, only colour-arrays previously held in WM, induced a greater midfrontal P170-response, together with left temporal- and late centro-parietal activity, upon re-exposure. These findings provide evidence for recruitment of an LTM-supporting neural network which facilitates visual WM maintenance. |
Katherine L. Hermann; Shridhar R. Singh; Isabelle A. Rosenthal; Dimitrios Pantazis; Bevil R. Conway Temporal dynamics of the neural representation of hue and luminance polarity Journal Article In: Nature Communications, vol. 13, pp. 1–19, 2022. @article{Hermann2022, Hue and luminance contrast are basic visual features. Here we use multivariate analyses of magnetoencephalography data to investigate the timing of the neural computations that extract them, and whether they depend on common neural circuits. We show that hue and luminance-contrast polarity can be decoded from MEG data and, with lower accuracy, both features can be decoded across changes in the other feature. These results are consistent with the existence of both common and separable neural mechanisms. The decoding time course is earlier and more temporally precise for luminance polarity than hue, a result that does not depend on task, suggesting that luminance contrast is an updating signal that separates visual events. Meanwhile, cross-temporal generalization is slightly greater for representations of hue compared to luminance polarity, providing a neural correlate of the preeminence of hue in perceptual grouping and memory. Finally, decoding of luminance polarity varies depending on the hues used to obtain training and testing data. The pattern of results is consistent with observations that luminance contrast is mediated by both L-M and S cone sub-cortical mechanisms. |
Beatriz Herrera; Jacob A. Westerberg; Michelle S. Schall; Alexander Maier; Geoffrey F. Woodman; Jeffrey D. Schall; Jorge J. Riera Resolving the mesoscopic missing link: Biophysical modeling of EEG from cortical columns in primates Journal Article In: NeuroImage, vol. 263, pp. 1–14, 2022. @article{Herrera2022, Event-related potentials (ERP) are among the most widely measured indices for studying human cognition. While their timing and magnitude provide valuable insights, their usefulness is limited by our understanding of their neural generators at the circuit level. Inverse source localization offers insights into such generators, but their solutions are not unique. To address this problem, scientists have assumed the source space generating such signals comprises a set of discrete equivalent current dipoles, representing the activity of small cortical regions. Based on this notion, theoretical studies have employed forward modeling of scalp potentials to understand how changes in circuit-level dynamics translate into macroscopic ERPs. However, experimental validation is lacking because it requires in vivo measurements of intracranial brain sources. Laminar local field potentials (LFP) offer a mechanism for estimating intracranial current sources. Yet, a theoretical link between LFPs and intracranial brain sources is missing. Here, we present a forward modeling approach for estimating mesoscopic intracranial brain sources from LFPs and predict their contribution to macroscopic ERPs. We evaluate the accuracy of this LFP-based representation of brain sources utilizing synthetic laminar neurophysiological measurements and then demonstrate the power of the approach in vivo to clarify the source of a representative cognitive ERP component. To that end, LFP was measured across the cortical layers of visual area V4 in macaque monkeys performing an attention demanding task. We show that area V4 generates dipoles through layer-specific transsynaptic currents that biophysically recapitulate the ERP component through the detailed forward modeling. The constraints imposed on EEG production by this method also revealed an important dissociation between computational and biophysical contributors. As such, this approach represents an important bridge between laminar microcircuitry, through the mesoscopic activity of cortical columns to the patterns of EEG we measure at the scalp. |
Zhenhong Hu; Immanuel B. H. Samuel; Sreenivasan Meyyappan; Ke Bo; Chandni Rana; Mingzhou Ding Aftereffects of frontoparietal theta tACS on verbal working memory: Behavioral and neurophysiological analysis Journal Article In: IBRO Neuroscience Reports, vol. 13, pp. 469–477, 2022. @article{Hu2022a, Verbal working memory is supported by a left-lateralized frontoparietal theta oscillatory (4–8 Hz) network. We tested whether stimulating the left frontoparietal network at theta frequency during verbal working memory can produce observable after-stimulation effects in behavior and neurophysiology. Weak theta-band alternating electric currents were delivered via two 4 × 1 HD electrode arrays centered at F3 and P3. Three stimulation configurations, including in-phase, anti-phase, or sham, were tested on three different days in a cross-over (within-subject) design. On each test day, the subject underwent three experimental sessions: pre-, duringand post-stimulation sessions. In all sessions, the subject performed a Sternberg verbal working memory task with three levels of memory load (load 2, 4 and 6), imposing three levels of cognitive demand. Analyzing behavioral and EEG data from the post-stimulation session, we report two main observations. First, in-phase stimulation improved task performance in subjects with higher working memory capacity (WMC) under higher memory load (load 6). Second, in-phase stimulation enhanced frontoparietal theta synchrony during working memory retention in subjects with higher WMC under higher memory loads (load 4 and load 6), and the enhanced frontoparietal theta synchronization is mainly driven by enhanced frontal→parietal theta Granger causality. These observations suggest that (1) in-phase theta transcranial alternating current stimulation (tACS) during verbal working memory can result in observable behavioral and neurophysiological consequences post stimulation, (2) the short-term plasticity effects are state- and individual-dependent, and (3) enhanced executive control underlies improved behavioral performance ### Competing Interest Statement The authors have declared no competing interest. |
Stefano Ioannucci; Guillermo Borragán; Alexandre Zénon Passive visual stimulation induces fatigue under conditions of high arousal elicited by auditory tasks Journal Article In: Journal of Experimental Psychology: General, vol. 151, no. 12, pp. 3097–3113, 2022. @article{Ioannucci2022, Theories of cognitive fatigue disagree on whether performance decrement is caused by motivational or functional alterations. Here, drawing inspiration from the habituation and visual adaptation literature, we tested the assumption that keeping neural networks active for an extensive period of time entails consequences at the subjective and objective level—the defining characteristics of fatigue—when confounds such as motivation, boredom, and level of skill are controlled. In Experiment 1, we revealed that passive visual stimulation affected the performance of a subsequent task that was carried out in the same portion of visual space. While under conditions of low cognitive load and arousal, participants improved their performance in the stimulated quadrant; the reverse was observed under high arousal conditions. This latter performance decrement correlated also with the reported subjective level of fatigue and occurred while neural responses to the saturating stimulus remained constant, as assessed through steady-state EEG. In subsequent experiments, we replicated and further characterized this performance deterioration effect, revealing its specificity to the stimulated eye and stimulus orientation. Across the three experiments, the decrease in performance was correlated with pupil-linked arousal, suggesting its mediating effect in this phenomenon. In sum, we show that repeated stimulation of neural networks under high-arousal conditions leads to their altered functional performance, a mechanism which may play a role in the development of global cognitive fatigue |
Gelu Ionescu; Aline Frey; Nathalie Guyader; Emmanuelle Kristensen; Anton Andreev; Anne Guérin-Dugué Synchronization of acquisition devices in neuroimaging: An application using co-registration of eye movements and electroencephalography Journal Article In: Behavior Research Methods, vol. 54, pp. 2545–2564, 2022. @article{Ionescu2022, Interest in applications for the simultaneous acquisition of data from different devices is growing. In neuroscience for example, co-registration complements and overcomes some of the shortcomings of individual methods. However, precise synchronization of the different data streams involved is required before joint data analysis. Our article presents and evaluates a synchronization method which maximizes the alignment of information across time. Synchronization through common triggers is widely used in all existing methods, because it is very simple and effective. However, this solution has been found to fail in certain practical situations, namely for the spurious detection of triggers and/or when the timestamps of triggers sampled by each acquisition device are not jointly distributed linearly for the entire duration of an experiment. We propose two additional mechanisms, the "Longest Common Subsequence" algorithm and a piecewise linear regression, in order to overcome the limitations of the classical method of synchronizing common triggers. The proposed synchronization method was evaluated using both real and artificial data. Co-registrations of electroencephalographic signals (EEG) and eye move- ments were used for real data. We compared the effectiveness of our method to another open source method implemented using EYE-EEG toolbox. Overall, we show that our method, implemented in C++ as a DOS application, is very fast, robust and fully automatic. |
Shanice E. W. Janssens; Sanne Ten Oever; Alexander T. Sack; Tom A. Graaf “Broadband alpha transcranial alternating current stimulation”: Exploring a new biologically calibrated brain stimulation protocol Journal Article In: NeuroImage, vol. 253, pp. 1–12, 2022. @article{Janssens2022, Transcranial alternating current stimulation (tACS) can be used to study causal contributions of oscillatory brain mechanisms to cognition and behavior. For instance, individual alpha frequency (IAF) tACS was reported to enhance alpha power and impact visuospatial attention performance. Unfortunately, such results have been inconsistent and difficult to replicate. In tACS, stimulation generally involves one frequency, sometimes individually calibrated to a peak value observed in an M/EEG power spectrum. Yet, the ‘peak' actually observed in such power spectra often contains a broader range of frequencies, raising the question whether a biologically calibrated tACS protocol containing this fuller range of alpha-band frequencies might be more effective. Here, we introduce ‘Broadband-alpha-tACS', a complex individually calibrated electrical stimulation protocol. We band-pass filtered left posterior resting-state EEG data around the IAF (± 2 Hz), and converted that time series into an electrical waveform for tACS stimulation of that same left posterior parietal cortex location. In other words, we stimulated a brain region with a ‘replay' of its own alpha-band frequency content, based on spontaneous activity. Within-subjects (N = 24), we compared to a sham tACS session the effects of broadband-alpha tACS, power-matched spectral inverse (‘alpha-removed') control tACS, and individual alpha frequency (IAF) tACS, on EEG alpha power and performance in an endogenous attention task previously reported to be affected by alpha tACS. Broadband-alpha-tACS significantly modulated attention task performance (i.e., reduced the rightward visuospatial attention bias in trials without distractors, and reduced attention benefits). Alpha-removed tACS also reduced the rightward visuospatial attention bias. IAF-tACS did not significantly modulate attention task performance compared to sham tACS, but also did not statistically significantly differ from broadband-alpha-tACS. This new broadband-alpha-tACS approach seems promising, but should be further explored and validated in future studies. |
Jianrong Jia; Ying Fan; Huan Luo Alpha-band phase modulates bottom-up feature processing Journal Article In: Cerebral Cortex, vol. 32, no. 6, pp. 1–9, 2022. @article{Jia2022a, Recent studies reveal that attention operates in a rhythmic manner, that is, sampling each location or feature alternatively over time. However, most evidence derives from top-down tasks, and it remains elusive whether bottom-up processing also entails dynamic coordination. Here, we developed a novel feature processing paradigm and combined time-resolved behavioral measurements and electroencephalogram (EEG) recordings to address the question. Specifically, a salient color in a multicolor display serves as a noninformative cue to capture attention and presumably reset the oscillations of feature processing. We then measured the behavioral performance of a probe stimulus associated with either high- or low-salient color at varied temporal lags after the cue. First, the behavioral results (i.e., reaction time) display an alpha-band (~8 Hz) profile with a consistent phase lag between high- and low-salient conditions. Second, simultaneous EEG recordings show that behavioral performance is modulated by the phase of alpha-band neural oscillation at the onset of the probes. Finally, high- and low-salient probes are associated with distinct preferred phases of alpha-band neural oscillations. Taken together, our behavioral and neural results convergingly support a central function of alpha-band rhythms in feature processing, that is, features with varied saliency levels are processed at different phases of alpha neural oscillations. |
Lu Jiang; Xiaoyang Li; Weihua Pei; Xiaorong Gao; Yijun Wang A hybrid brain-computer interface based on visual evoked potential and pupillary response Journal Article In: Frontiers in Human Neuroscience, vol. 16, pp. 1–16, 2022. @article{Jiang2022a, Brain-computer interface (BCI) based on steady-state visual evoked potential (SSVEP) has been widely studied due to the high information transfer rate (ITR), little user training, and wide subject applicability. However, there are also disadvantages such as visual discomfort and “BCI illiteracy.” To address these problems, this study proposes to use low-frequency stimulations (12 classes, 0.8–2.12 Hz with an interval of 0.12 Hz), which can simultaneously elicit visual evoked potential (VEP) and pupillary response (PR) to construct a hybrid BCI (h-BCI) system. Classification accuracy was calculated using supervised and unsupervised methods, respectively, and the hybrid accuracy was obtained using a decision fusion method to combine the information of VEP and PR. Online experimental results from 10 subjects showed that the averaged accuracy was 94.90 ± 2.34% (data length 1.5 s) for the supervised method and 91.88 ± 3.68% (data length 4 s) for the unsupervised method, which correspond to the ITR of 64.35 ± 3.07 bits/min (bpm) and 33.19 ± 2.38 bpm, respectively. Notably, the hybrid method achieved higher accuracy and ITR than that of VEP and PR for most subjects, especially for the short data length. Together with the subjects' feedback on user experience, these results indicate that the proposed h-BCI with the low-frequency stimulation paradigm is more comfortable and favorable than the traditional SSVEP-BCI paradigm using the alpha frequency range. |
Richard Johnston; Adam C. Snyder; Rachel S. Schibler; Matthew A. Smith EEG signals index a global signature of arousal embedded in neuronal population recordings Journal Article In: eNeuro, vol. 9, no. 3, pp. 1–16, 2022. @article{Johnston2022a, Electroencephalography (EEG) has long been used to index brain states, from early studies describing activity in the presence and absence of visual stimulation to modern work employing complex perceptual tasks. These studies have shed light on brain-wide signals but often lack explanatory power at the single neuron level. Similarly, single neuron recordings can suffer from an inability to measure brain-wide signals accessible using EEG. Here, we combined these techniques while monkeys performed a change detection task and discovered a novel link between spontaneous EEG activity and a neural signal embedded in the spiking responses of neuronal populations. This “slow drift” was associated with fluctuations in the subjects' arousal levels over time: decreases in prestimulus a power were accompanied by increases in pupil size and decreases in microsaccade rate. These re- sults show that brain-wide EEG signals can be used to index modes of activity present in single neuron recordings, that in turn reflect global changes in brain state that influence perception and behavior. |
Oren Kadosh; Yoram Bonneh Face familiarity revealed by fixational eye movements and fixation-related potentials in free viewing Journal Article In: Scientific Reports, vol. 12, no. 1, pp. 1–13, 2022. @article{Kadosh2022a, Event-related potentials (ERPs) and the oculomotor inhibition (OMI) in response to visual transients are known to be sensitive to stimulus properties, attention, and expectation. We have recently found that the OMI is also sensitive to face familiarity. In natural vision, stimulation of the visual cortex is generated primarily by saccades, and it has been recently suggested that fixation-related potentials (FRPs) share similar components with the ERPs. Here, we investigated whether FRPs and microsaccade inhibition (OMI) in free viewing are sensitive to face familiarity. Observers freely watched a slideshow of seven unfamiliar and one familiar facial images presented randomly for 4-s periods, with multiple images per identity. We measured the occipital fixation-related N1 relative to the P1 magnitude as well as the associated fixation-triggered OMI. We found that the average N1-P1 was significantly smaller and the OMI was shorter for the familiar face, compared with any of the seven unfamiliar faces. Moreover, the P1 was suppressed across saccades for the familiar but not for the unfamiliar faces. Our results highlight the sensitivity of the occipital FRPs to stimulus properties such as face familiarity and advance our understanding of the integration process across successive saccades in natural vision. |
Sharif I. Kronemer; Mark Aksen; Julia Z. Ding; Jun Hwan Ryu; Qilong Xin; Zhaoxiong Ding; Jacob S. Prince; Hunki Kwon; Aya Khalaf; Sarit Forman; David S. Jin; Kevin Wang; Kaylie Chen; Claire Hu; Akshar Agarwal; Erik Saberski; Syed Mohammad Adil Wafa; Owen P. Morgan; Jia Wu; Kate L. Christison-Lagay; Nicholas Hasulak; Martha Morrell; Alexandra Urban; R. Todd Constable; Michael Pitts; R. Mark Richardson; Michael J. Crowley; Hal Blumenfeld Human visual consciousness involves large scale cortical and subcortical networks independent of task report and eye movement activity Journal Article In: Nature Communications, vol. 13, pp. 1–17, 2022. @article{Kronemer2022, The full neural circuits of conscious perception remain unknown. Using a visual perception task, we directly recorded a subcortical thalamic awareness potential (TAP). We also developed a unique paradigm to classify perceived versus not perceived stimuli using eye measurements to remove confounding signals related to reporting on conscious experiences. Using fMRI, we discovered three major brain networks driving conscious visual perception independent of report: first, increases in signal detection regions in visual, fusiform cortex, and frontal eye fields; and in arousal/salience networks involving midbrain, thalamus, nucleus accumbens, anterior cingulate, and anterior insula; second, increases in frontoparietal attention and executive control networks and in the cerebellum; finally, decreases in the default mode network. These results were largely maintained after excluding eye movement-based fMRI changes. Our findings provide evidence that the neurophysiology of consciousness is complex even without overt report, involving multiple cortical and subcortical networks overlapping in space and time. |
Louisa Kulke; Lena Brümmer; Arezoo Pooresmaeili; Annekathrin Schacht Visual competition attenuates emotion effects during overt attention shifts Journal Article In: Psychophysiology, vol. 59, pp. 1–14, 2022. @article{Kulke2022, Numerous different objects are simultaneously visible in a person's visual field, competing for attention. This competition has been shown to affect eye-movements and early neural responses toward stimuli, while the role of a stimulus' emotional meaning for mechanisms of overt attention shifts under competition is unclear. The current study combined EEG and eye-tracking to investigate effects of competition and emotional content on overt shifts of attention to human face stimuli. Competition prolonged the latency of the P1 component and of saccades, while faces showing emotional expressions elicited an early posterior negativity (EPN). Remarkably, the emotion-related modulation of the EPN was attenuated when two stimuli were competing for attention compared to non-competition. In contrast, no interaction effects of emotional expression and competition were observed on other event-related potentials. This finding indicates that competition can decelerate attention shifts in general and also diminish the emotion-driven attention capture, measured through the smaller effects of emotional expression on EPN amplitude. Reduction of the brain's responsiveness to emotional content in the presence of distractors contradicts models that postulate fully automatic processing of emotions. |
Wupadrasta Santosh Kumar; Keerthana Manikandan; Dinavahi V. P. S. Murty; Ranjini Garani Ramesh; Simran Purokayastha; Mahendra Javali; Naren Prahalada Rao; Supratim Ray Stimulus-induced narrowband gamma oscillations are test–retest reliable in human EEG Journal Article In: Cerebral Cortex Communications, vol. 3, no. 1, pp. 1–15, 2022. @article{Kumar2022a, Visual stimulus-induced gamma oscillations in electroencephalogram (EEG) recordings have been recently shown to be compromised in subjects with preclinical Alzheimer's Disease (AD), suggesting that gamma could be an inexpensive biomarker for AD diagnosis provided its characteristics remain consistent across multiple recordings. Previous magnetoencephalography studies in young subjects have reported consistent gamma power over recordings separated by a few weeks to months. Here, we assessed the consistency of stimulus-induced slow (20–35 Hz) and fast gamma (36–66 Hz) oscillations in subjects (n = 40) (age: 50–88 years) in EEG recordings separated by a year, and tested the consistency in the magnitude of gamma power, its temporal evolution and spectral profile. Gamma had distinct spectral/temporal characteristics across subjects, which remained consistent across recordings (average intraclass correlation of ~0.7). Alpha (8–12 Hz) and steady-state-visually evoked-potentials were also reliable. We further tested how EEG features can be used to identify 2 recordings as belonging to the same versus different subjects and found high classifier performance (AUC of ~0.89), with temporal evolution of slow gamma and spectral profile being most informative. These results suggest that EEG gamma oscillations are reliable across sessions separated over long durations and can also be a potential tool for subject identification. |
Dominik Welke; Edward A. Vessel Naturalistic viewing conditions can increase task engagement and aesthetic preference but have only minimal impact on EEG quality Journal Article In: NeuroImage, vol. 256, pp. 1–19, 2022. @article{Welke2022, Free gaze and moving images are typically avoided in EEG experiments due to the expected generation of artifacts and noise. Yet for a growing number of research questions, loosening these rigorous restrictions would be beneficial. Among these is research on visual aesthetic experiences, which often involve open-ended exploration of highly variable stimuli. Here we systematically compare the effect of conservative vs. more liberal experimental settings on various measures of behavior, brain activity and physiology in an aesthetic rating task. Our primary aim was to assess EEG signal quality. 43 participants either maintained fixation or were allowed to gaze freely, and viewed either static images or dynamic (video) stimuli consisting of dance performances or nature scenes. A passive auditory background task (auditory steady-state response; ASSR) was added as a proxy measure for overall EEG recording quality. We recorded EEG, ECG and eye tracking data, and participants rated their aesthetic preference and state of boredom on each trial. Whereas both behavioral ratings and gaze behavior were affected by task and stimulus manipulations, EEG SNR was barely affected and generally robust across all conditions, despite only minimal preprocessing and no trial rejection. In particular, we show that using video stimuli does not necessarily result in lower EEG quality and can, on the contrary, significantly reduce eye movements while increasing both the participants' aesthetic response and general task engagement. We see these as encouraging results indicating that — at least in the lab — more liberal experimental conditions can be adopted without significant loss of signal quality. |
Wenyuan Yu; Wenhui Sun; Nai Ding Asymmetrical cross-modal influence on neural encoding of auditory and visual features in natural scenes Journal Article In: NeuroImage, vol. 255, pp. 1–10, 2022. @article{Yu2022b, Natural scenes contain multi-modal information, which is integrated to form a coherent perception. Previous studies have demonstrated that cross-modal information can modulate neural encoding of low-level sensory features. These studies, however, mostly focus on the processing of single sensory events or rhythmic sensory sequences. Here, we investigate how the neural encoding of basic auditory and visual features is modulated by cross-modal information when the participants watch movie clips primarily composed of non-rhythmic events. We presented audiovisual congruent and audiovisual incongruent movie clips, and since attention can modulate cross-modal interactions, we separately analyzed high- and low-arousal movie clips. We recorded neural responses using electroencephalography (EEG), and employed the temporal response function (TRF) to quantify the neural encoding of auditory and visual features. The neural encoding of sound envelope is enhanced in the audiovisual congruent condition than the incongruent condition, but this effect is only significant for high-arousal movie clips. In contrast, audiovisual congruency does not significantly modulate the neural encoding of visual features, e.g., luminance or visual motion. In summary, our findings demonstrate asymmetrical cross-modal interactions during the processing of natural scenes that lack rhythmicity: Congruent visual information enhances low-level auditory processing, while congruent auditory information does not significantly modulate low-level visual processing. |
Marie Zelenina; MacIej Kosilo; Janir Da Cruz; Marília Antunes; Patrícia Figueiredo; Mitul A. Mehta; Diana Prata Temporal dynamics of intranasal oxytocin in human brain electrophysiology Journal Article In: Cerebral Cortex, vol. 32, no. 14, pp. 3110–3126, 2022. @article{Zelenina2022, Oxytocin (OT) is a key modulator of human social cognition, popular in behavioral neuroscience. To adequately design and interpret intranasal OT (IN-OT) research, it is crucial to know for how long it affects human brain function once administered. However, this has been mostly deduced from peripheral body f luids studies, or uncommonly used dosages. We aimed to characterize IN-OT's effects on human brain function using resting-state EEG microstates across a typical experimental session duration. Nineteen healthy males participated in a double-blind, placebo-controlled, within-subject, cross-over design of 24 IU of IN-OT in 12-min windows 15 min-to1 h 42min after administration. We observed IN-OT effects on all microstates, across the observation span. During eyes-closed, IN-OT increased duration and contribution of A and contribution and occurrence of D, decreased duration and contribution of B and C; and increased transition probability C-to-B and C-to-D. In eyes-open, it increased A-to-C and A-to-D. As microstates A and D have been related to phonological auditory and attentional networks, respectively, we posit IN-OT may tune the brain for reception of external stimuli, particularly of social nature - tentatively supporting current neurocognitive hypotheses of OT. Moreover, we contrast our overall results against a comprehensive literature review of IN-OT time-course effects in the brain, highlighting comparability issues. |
Maren-Isabel Wolf; Maximilian Bruchmann; Gilles Pourtois; Sebastian Schindler; Thomas Straube In: Cerebral Cortex, vol. 32, no. 10, pp. 2112–2128, 2022. @article{Wolf2022a, Until today, there is an ongoing discussion if attention processes interact with the information processing stream already at the level of the C1, the earliest visual electrophysiological response of the cortex. We used two highly powered experiments (each N = 52) and examined the effects of task relevance, spatial attention, and attentional load on individual C1 amplitudes for the upper or lower visual hemifield. Bayesian models revealed evidence for the absence of load effects but substantial modulations by task-relevance and spatial attention. When the C1-eliciting stimulus was a task-irrelevant, interfering distracter, we observed increased C1 amplitudes for spatially unattended stimuli. For spatially attended stimuli, different effects of task-relevance for the two experiments were found. Follow-up exploratory single-trial analyses revealed that subtle but systematic deviations from the eye-gaze position at stimulus onset between conditions substantially influenced the effects of attention and task relevance on C1 amplitudes, especially for the upper visual field. For the subsequent P1 component, attentional modulations were clearly expressed and remained unaffected by these deviations. Collectively, these results suggest that spatial attention, unlike load or task relevance, can exert dissociable top-down modulatory effects at the C1 and P1 levels. |
Seth B. Winward; James Siklos-Whillans; Roxane J. Itier In: Neuroimage: Reports, vol. 2, no. 4, pp. 1–17, 2022. @article{Winward2022, Recent ERP research using a gaze-contingent paradigm suggests the face-sensitive N170 component is modulated by the presence of a face outline, the number of parafoveal facial features, and the type of feature in parafovea (Parkington and Itier, 2019). The present study re-analyzed these data using robust mass univariate statistics available through the LIMO toolbox, allowing the examination of the ERP signal across all electrodes and time points. We replicated the finding that the presence of a face outline significantly reduced ERP latencies and amplitudes, suggesting it is an important cue to the prototypical face template. However, we found that this effect began around 114 ms, and was maximal during the P1-N170 and N170-P2 intervals. The number of features present in parafovea also impacted the entire waveform, with systematic reductions in amplitude and latency as the number of features increased. This effect was maximal around 120 ms during the P1-N170 interval and around 170 ms between the N170 and P2. The ERP response was also modulated by feature type; contrary to previous findings this effect was maximal around 200 ms and the P2 peak. Although we provide partial repli- cation of the previous results on the N170, the effects were more temporally distributed in the present analysis. These effects were generally maximal before and after the N170 and were the weakest at the N170 peak itself. This re-analysis demonstrates that classical ERP analysis can obscure important aspects of face processing beyond the N170 peak, and that tools like mass univariate statistics are needed to shed light on the whole time-course of face processing. |
TianHong Zhang; YingYu Yang; LiHua Xu; XiaoChen Tang; YeGang Hu; Xin Xiong; YanYan Wei; HuiRu Ru Cui; YingYing Tang; HaiChun Liu; Tao Chen; Zhi Liu; Li Hui; ChunBo Li; XiaoLi Guo; JiJun Wang Inefficient integration during multiple facial processing in pre-morbid and early phases of psychosis Journal Article In: The World Journal of Biological Psychiatry, vol. 23, no. 5, pp. 1–13, 2022. @article{Zhang2022j, Objectives: We used eye-tracking to evaluate multiple facial context processing and event-related potential (ERP) to evaluate multiple facial recognition in individuals at clinical high risk (CHR) for psychosis. Methods: In total, 173 subjects (83 CHRs and 90 healthy controls [HCs]) were included and their emotion perception performances were accessed. A total of 40 CHRs and 40 well-matched HCs completed an eye-tracking task where they viewed pictures depicting a person in the foreground, presented as context-free, context-compatible, and context-incompatible. During the two-year follow-up, 26 CHRs developed psychosis, including 17 individuals who developed first-episode schizophrenia (FES). Eighteen well-matched HCs were made to complete the face number detection ERP task with image stimuli of one, two, or three faces. Results: Compared to the HC group, the CHR group showed reduced visual attention to contextual processing when viewing multiple faces. With the increasing complexity of contextual faces, the differences in eye-tracking characteristics also increased. In the ERP task, the N170 amplitude decreased with a higher face number in FES patients, while it increased with a higher face number in HCs. Conclusions: Individuals in the very early phase of psychosis showed facial processing deficits with supporting evidence of different scan paths during context processing and disruption of N170 during multiple facial recognition. |
Jing Zhu; Shiqing Wei; Xiannian Xie; Changlin Yang; Yizhou Li; Xiaowei Li; Bin Hu Content-based multiple evidence fusion on EEG and eye movements for mild depression recognition Journal Article In: Computer Methods and Programs in Biomedicine, vol. 226, pp. 1–11, 2022. @article{Zhu2022a, Background and objective: Depression is a serious neurological disorder that has become a major health problem worldwide. The detection of mild depression is important for the diagnosis of depression in early stages. This research seeks to find a more accurate fusion model which can be used for mild depression detection using Electroencephalography and eye movement data. Methods: This study proposes a content-based multiple evidence fusion (CBMEF) method, which fuses EEG and eye movement data at decision level. The method mainly includes two modules, the classification performance matrix module and the dual-weight fusion module. The classification performance matrices of different modalities are estimated by Bayesian rule based on confusion matrix and Mahalanobis distance, and the matrices were used to correct the classification results. Then the relative conflict degree of each modality is calculated, and different weights are assigned to the above modalities at the decision fusion layer according to this conflict degree. Results: The experimental results show that the proposed method outperforms other fusion methods as well as the single modality results. The highest accuracies achieved 91.12%, and sensitivity, specificity and precision were 89.20%, 93.03%, 92.76%. Conclusions: The promising results showed the potential of the proposed approach for the detection of mild depression. The idea of introducing the classification performance matrix and the dual-weight model to multimodal biosignals fusion casts a new light on the researches of depression recognition. |
Jing Zhu; Changlin Yang; Xiannian Xie; Shiqing Wei; Yizhou Li; Xiaowei Li; Bin Hu Mutual Information Based Fusion Model (MIBFM): Mild depression recognition using EEG and pupil area signals Journal Article In: Journal of LATEX Class Files, vol. 3045, pp. 1–14, 2022. @article{Zhu2022b, The detection of mild depression is conducive to the early intervention and treatment of depression. This study explored the fusion of electroencephalography (EEG) and pupil area signals to build an effective and convenient mild depression recognition model. We proposed Mutual Information Based Fusion Model (MIBFM), which innovatively used pupil area signals to select EEG electrodes based on mutual information. Then we extracted features from EEG and pupil area signals in different bands, and fused bimodal features using the denoising autoencoder. Experimental results showed that MIBFM could obtain the highest accuracy of 87.03%. And MIBFM exhibited better performance than other existing methods. Our findings validate the effectiveness of the use of pupil area as signals, which makes eye movement signals can be easily obtained using high resolution camera, and the EEG electrode selection scheme based on mutual information is also proved to be an applicable solution for data dimension reduction and multimodal complementary information screening. This study casts a new light for mild depression recognition using multimodal data of EEG and pupil area signals, and provides a theoretical basis for the development of portable and universal application systems. |
Stephen Whitmarsh; Christophe Gitton; Veikko Jousmäki; Jérôme Sackur; Catherine Tallon-Baudry Neuronal correlates of the subjective experience of attention Journal Article In: European Journal of Neuroscience, vol. 55, no. 11-12, pp. 3465–3482, 2022. @article{Whitmarsh2022, The effect of top–down attention on stimulus-evoked responses and alpha oscillations and the association between arousal and pupil diameter are well established. However, the relationship between these indices, and their contribution to the subjective experience of attention, remains largely unknown. Participants performed a sustained (10–30 s) attention task in which rare (10%) targets were detected within continuous tactile stimulation (16 Hz). Trials were followed by attention ratings on an 8-point visual scale. Attention ratings correlated negatively with contralateral somatosensory alpha power and positively with pupil diameter. The effect of pupil diameter on attention ratings extended into the following trial, reflecting a sustained aspect of attention related to vigilance. The effect of alpha power did not carry over to the next trial and furthermore mediated the association between pupil diameter and attention ratings. Variations in steady-state amplitude reflected stimulus processing under the influence of alpha oscillations but were only weakly related to subjective ratings of attention. Together, our results show that both alpha power and pupil diameter are reflected in the subjective experience of attention, albeit on different time spans, while continuous stimulus processing might not contribute to the experience of attention. |
Wen Wen; Zhibang Huang; Yin Hou; Sheng Li Tracking neural markers of template formation and implementation in attentional inhibition under different distractor consistency Journal Article In: Journal of Neuroscience, vol. 42, no. 24, pp. 4927–4936, 2022. @article{Wen2022, Performing visual search tasks requires optimal attention deployment to promote targets and inhibit distractors. Rejection templates based on the distractor's feature can be built to constrain the search process. We measured electroencephalography (EEG) of human participants of both sexes when they performed a visual search task in conditions where the distractor cues were constant within a block (fixed-cueing) or changed on a trial-by-trial basis (varied-cueing). In the fixed-cueing condition, sustained decoding of the cued colors could be achieved during the retention interval and the participants with higher decoding accuracy showed larger suppression benefits of the distractor cueing in the search period. In the varied-cueing condition, the cued color could only be transiently decoded after its onset and the higher decoding accuracy was observed from the participants who demonstrated lower suppression benefit. The differential neural representations of the to-be-ignored color in the two cueing conditions as well as their reverse associations with behavioral performance implied that rejection templates were formed in the fixed-cueing condition but not in the varied-cueing condition. Additionally, we observed stronger posterior alpha lateralization and mid-frontal theta/beta power during the retention interval of the varied-cueing condition, indicating the cognitive costs in template formation caused by the trialwise change of distractor colors. Taken together, our findings revealed the neural markers associated with the critical roles of distractor consistency in linking template formation to successful inhibition. |
2021 |
Aurélien Weiss; Valérian Chambon; Junseok K. Lee; Jan Drugowitsch; Valentin Wyart Interacting with volatile environments stabilizes hidden-state inference and its brain signatures Journal Article In: Nature Communications, vol. 12, pp. 2228, 2021. @article{Weiss2021, Making accurate decisions in uncertain environments requires identifying the generative cause of sensory cues, but also the expected outcomes of possible actions. Although both cognitive processes can be formalized as Bayesian inference, they are commonly studied using different experimental frameworks, making their formal comparison difficult. Here, by framing a reversal learning task either as cue-based or outcome-based inference, we found that humans perceive the same volatile environment as more stable when inferring its hidden state by interaction with uncertain outcomes than by observation of equally uncertain cues. Multivariate patterns of magnetoencephalographic (MEG) activity reflected this behavioral difference in the neural interaction between inferred beliefs and incoming evidence, an effect originating from associative regions in the temporal lobe. Together, these findings indicate that the degree of control over the sampling of volatile environments shapes human learning and decision-making under uncertainty. |
Bo Yao; Jason R. Taylor; Briony Banks; Sonja A. Kotz Reading direct speech quotes increases theta phase-locking: Evidence for cortical tracking of inner speech? Journal Article In: NeuroImage, vol. 239, pp. 118313, 2021. @article{Yao2021a, Growing evidence shows that theta-band (4–7 Hz) activity in the auditory cortex phase-locks to rhythms of overt speech. Does theta activity also encode the rhythmic dynamics of inner speech? Previous research established that silent reading of direct speech quotes (e.g., Mary said: “This dress is lovely!”) elicits more vivid inner speech than indirect speech quotes (e.g., Mary said that the dress was lovely). As we cannot directly track the phase alignment between theta activity and inner speech over time, we used EEG to measure the brain's phase-locked responses to the onset of speech quote reading. We found that direct (vs. indirect) quote reading was associated with increased theta phase synchrony over trials at 250–500 ms post-reading onset, with sources of the evoked activity estimated in the speech processing network. An eye-tracking control experiment confirmed that increased theta phase synchrony in direct quote reading was not driven by eye movement patterns, and more likely reflects synchronous phase resetting at the onset of inner speech. These findings suggest a functional role of theta phase modulation in reading-induced inner speech. |
Roberto F. Salanamca-Giron; Estelle Raffin; Sarah B. Zandvliet; Martin Seeber; Christoph M. Michel; Paul Sauseng; Krystel R. Huxlin; Friedhelm C. Hummel Enhancing visual motion discrimination by desynchronizing bifocal oscillatory activity Journal Article In: NeuroImage, vol. 240, pp. 118299, 2021. @article{SalanamcaGiron2021, Visual motion discrimination involves reciprocal interactions in the alpha band between the primary visual cortex (V1) and mediotemporal areas (V5/MT). We investigated whether modulating alpha phase synchronization using individualized multisite transcranial alternating current stimulation (tACS) over V5 and V1 regions would improve motion discrimination. We tested 3 groups of healthy subjects with the following conditions: (1) individualized In-Phase V1alpha-V5alpha tACS (0° lag), (2) individualized Anti-Phase V1alpha-V5alpha tACS (180° lag) and (3) sham tACS. Motion discrimination and EEG activity were recorded before, during and after tACS. Performance significantly improved in the Anti-Phase group compared to the In-Phase group 10 and 30 min after stimulation. This result was explained by decreases in bottom-up alpha-V1 gamma-V5 phase-amplitude coupling. One possible explanation of these results is that Anti-Phase V1alpha-V5alpha tACS might impose an optimal phase lag between stimulation sites due to the inherent speed of wave propagation, hereby supporting optimized neuronal communication. |
Giulia C. Salgari; Geoffrey F. Potts; Joseph Schmidt; Chi C. Chan; Christopher C. Spencer; Jeffrey S. Bedwell Event-related potentials to rare visual targets and negative symptom severity in a transdiagnostic psychiatric sample Journal Article In: Clinical Neurophysiology, vol. 132, no. 7, pp. 1526–1536, 2021. @article{Salgari2021, Objectives: Negative psychiatric symptoms are often resistant to treatments, regardless of the disorder in which they appear. One model for a cause of negative symptoms is impairment in higher-order cognition. The current study examined how particular bottom-up and top-down mechanisms of selective attention relate to severity of negative symptoms across a transdiagnostic psychiatric sample. Methods: The sample consisted of 130 participants: 25 schizophrenia-spectrum disorders, 26 bipolar disorders, 18 unipolar depression, and 61 nonpsychiatric controls. The relationships between attentional event-related potentials following rare visual targets (i.e., N1, N2b, P2a, and P3b) and severity of the negative symptom domains of anhedonia, avolition, and blunted affect were evaluated using frequentist and Bayesian analyses. Results: P3b and N2b mean amplitudes were inversely related to the Positive and Negative Syndrome Scale-Negative Symptom Factor severity score across the entire sample. Subsequent regression analyses showed a significant negative transdiagnostic relationship between P3b amplitude and blunted affect severity. Conclusions: Results indicate that negative symptoms, and particularly blunted affect, may have a stronger association with deficits in top-down mechanisms of selective attention. Significance: This suggests that people with greater severity of blunted affect, independent of diagnosis, do not allocate sufficient cognitive resources when engaging in activities requiring selective attention. |
Sebastian Schindler; Clara Tirloni; Maximilian Bruchmann; Thomas Straube Face and emotional expression processing under continuous perceptual load tasks: An ERP study Journal Article In: Biological Psychology, vol. 161, pp. 108056, 2021. @article{Schindler2021, High perceptual load is thought to impair already the early stages of visual processing of task-irrelevant visual stimuli. However, recent studies showed no effects of perceptual load on early ERPs in response to task-irrelevant emotional faces. In this preregistered EEG study (N = 40), we investigated the effects of continuous perceptual load on ERPs to fearful and neutral task-irrelevant faces and their phase-scrambled versions. Perceptual load did not modulate face or emotion effects for the P1 or N170. In contrast, larger face-scramble and fearful-neutral differentiation were found during low as compared to high load for the Early Posterior Negativity (EPN). Further, face-independent P1, but face-dependent N170 emotional modulations were observed. Taken together, our findings show that P1 and N170 face and emotional modulations are highly resistant to load manipulations, indicating a high degree of automaticity during this processing stage, whereas the EPN might represent a bottleneck in visual information processing. |
Constanze Schmitt; Jakob C. B. Schwenk; Adrian Schütz; Jan Churan; André Kaminiarz; Frank Bremmer Preattentive processing of visually guided self-motion in humans and monkeys Journal Article In: Progress in Neurobiology, vol. 205, pp. 102117, 2021. @article{Schmitt2021, The visually-based control of self-motion is a challenging task, requiring – if needed – immediate adjustments to keep on track. Accordingly, it would appear advantageous if the processing of self-motion direction (heading) was predictive, thereby accelerating the encoding of unexpected changes, and un-impaired by attentional load. We tested this hypothesis by recording EEG in humans and macaque monkeys with similar experimental protocols. Subjects viewed a random dot pattern simulating self-motion across a ground plane in an oddball EEG paradigm. Standard and deviant trials differed only in their simulated heading direction (forward-left vs. forward-right). Event-related potentials (ERPs) were compared in order to test for the occurrence of a visual mismatch negativity (vMMN), a component that reflects preattentive and likely also predictive processing of sensory stimuli. Analysis of the ERPs revealed signatures of a prediction mismatch for deviant stimuli in both humans and monkeys. In humans, a MMN was observed starting 110 ms after self-motion onset. In monkeys, peak response amplitudes following deviant stimuli were enhanced compared to the standard already 100 ms after self-motion onset. We consider our results strong evidence for a preattentive processing of visual self-motion information in humans and monkeys, allowing for ultrafast adjustments of their heading direction. |
Omer Sharon; Firas Fahoum; Yuval Nir Transcutaneous vagus nerve stimulation in humans induces pupil dilation and attenuates alpha oscillations Journal Article In: Journal of Neuroscience, vol. 41, no. 2, pp. 320–330, 2021. @article{Sharon2021, Vagus nerve stimulation (VNS) is widely used to treat drug-resistant epilepsy and depression. While the precise mechanisms mediating its long-term therapeutic effects are not fully resolved, they likely involve locus coeruleus (LC) stimulation via the nucleus of the solitary tract, which receives afferent vagal inputs. In rats, VNS elevates LC firing and forebrain noradrenaline levels, whereas LC lesions suppress VNS therapeutic efficacy. Noninvasive transcutaneous VNS (tVNS) uses electrical stimulation that targets the auricular branch of the vagus nerve at the cymba conchae of the ear. However, the extent to which tVNS mimics VNS remains unclear. Here, we investigated the short-term effects of tVNS in healthy human male volunteers (n = 24), using high-density EEG and pupillometry during visual fixation at rest. We compared short (3.4 s) trials of tVNS to sham electrical stimulation at the earlobe (far from the vagus nerve branch) to control for somatosensory stimulation. Although tVNS and sham stimulation did not differ in subjective intensity ratings, tVNS led to robust pupil dilation (peaking 4-5 s after trial onset) that was significantly higher than following sham stimulation. We further quantified, using parallel factor analysis, how tVNS modulates idle occipital alpha (8-13Hz) activity identified in each participant. We found greater attenuation of alpha oscillations by tVNS than by sham stimulation. This demonstrates that tVNS reliably induces pupillary and EEG markers of arousal beyond the effects of somatosensory stimulation, thus supporting the hypothesis that tVNS elevates noradrenaline and other arousal-promoting neuromodulatory signaling, and mimics invasive VNS. |
Jack W. Silcox; Brennan R. Payne The costs (and benefits) of effortful listening on context processing: A simultaneous electrophysiology, pupillometry, and behavioral study Journal Article In: Cortex, vol. 142, pp. 296–316, 2021. @article{Silcox2021, There is an apparent disparity between the fields of cognitive audiology and cognitive electrophysiology as to how linguistic context is used when listening to perceptually challenging speech. To gain a clearer picture of how listening effort impacts context use, we conducted a pre-registered study to simultaneously examine electrophysiological, pupillometric, and behavioral responses when listening to sentences varying in contextual constraint and acoustic challenge in the same sample. Participants (N = 44) listened to sentences that were highly constraining and completed with expected or unexpected sentence-final words (“The prisoners were planning their escape/party”) or were low-constraint sentences with unexpected sentence-final words (“All day she thought about the party”). Sentences were presented either in quiet or with +3 dB SNR background noise. Pupillometry and EEG were simultaneously recorded and subsequent sentence recognition and word recall were measured. While the N400 expectancy effect was diminished by noise, suggesting impaired real-time context use, we simultaneously observed a beneficial effect of constraint on subsequent recognition memory for degraded speech. Importantly, analyses of trial-to-trial coupling between pupil dilation and N400 amplitude showed that when participants' showed increased listening effort (i.e., greater pupil dilation), there was a subsequent recovery of the N400 effect, but at the same time, higher effort was related to poorer subsequent sentence recognition and word recall. Collectively, these findings suggest divergent effects of acoustic challenge and listening effort on context use: while noise impairs the rapid use of context to facilitate lexical semantic processing in general, this negative effect is attenuated when listeners show increased effort in response to noise. However, this effort-induced reliance on context for online word processing comes at the cost of poorer subsequent memory. |
Rodolfo Solís-Vivanco; Ole Jensen; Mathilde Bonnefond New insights on the ventral attention network: Active suppression and involuntary recruitment during a bimodal task Journal Article In: Human Brain Mapping, vol. 42, no. 6, pp. 1699–1713, 2021. @article{SolisVivanco2021, Detection of unexpected, yet relevant events is essential in daily life. fMRI studies have revealed the involvement of the ventral attention network (VAN), including the temporo-parietal junction (TPJ), in such process. In this MEG study with 34 participants (17 women), we used a bimodal (visual/auditory) attention task to determine the neuronal dynamics associated with suppression of the activity of the VAN during top-down attention and its recruitment when information from the unattended sensory modality is involuntarily integrated. We observed an anticipatory power increase of alpha/beta oscillations (12–20 Hz, previously associated with functional inhibition) in the VAN following a cue indicating the modality to attend. Stronger VAN power increases were associated with better task performance, suggesting that the VAN suppression prevents shifting attention to distractors. Moreover, the TPJ was synchronized with the frontal eye field in that frequency band, indicating that the dorsal attention network (DAN) might participate in such suppression. Furthermore, we found a 12–20 Hz power decrease and enhanced synchronization, in both the VAN and DAN, when information between sensory modalities was congruent, suggesting an involvement of these networks when attention is involuntarily enhanced due to multisensory integration. Our results show that effective multimodal attentional allocation includes the modulation of the VAN and DAN through upper-alpha/beta oscillations. Altogether these results indicate that the suppressing role of alpha/beta oscillations might operate beyond sensory regions. |
Jemaine E. Stacey; Mark Crook-Rumsey; Alexander Sumich; Christina J. Howard; Trevor Crawford; Kinneret Livne; Sabrina Lenzoni; Stephen Badham Age differences in resting state EEG and their relation to eye movements and cognitive performance Journal Article In: Neuropsychologia, vol. 157, pp. 107887, 2021. @article{Stacey2021, Prior research has focused on EEG differences across age or EEG differences across cognitive tasks/eye tracking. There are few studies linking age differences in EEG to age differences in behavioural performance which is necessary to establish how neuroactivity corresponds to successful and impaired ageing. Eighty-six healthy participants completed a battery of cognitive tests and eye-tracking measures. Resting state EEG (n = 75, 31 young, 44 older adults) was measured for delta, theta, alpha and beta power as well as for alpha peak frequency. Age deficits in cognition were aligned with the literature, showing working memory and inhibitory deficits along with an older adult advantage in vocabulary. Older adults showed poorer eye movement accuracy and response times, but we did not replicate literature showing a greater age deficit for antisaccades than for prosaccades. We replicated EEG literature showing lower alpha peak frequency in older adults but not literature showing lower alpha power. Older adults also showed higher beta power and less parietal alpha power asymmetry than young adults. Interaction effects showed that better prosaccade performance was related to lower beta power in young adults but not in older adults. Performance at the trail making test part B (measuring task switching and inhibition) was improved for older adults with higher resting state delta power but did not depend on delta power for young adults. It is argued that individuals with higher slow-wave resting EEG may be more resilient to age deficits in tasks that utilise cross-cortical processing. |
Benjamin J. Stauch; Alina Peter; Heike Schuler; Pascal Fries Stimulus-specific plasticity in human visual gamma-band activity and functional connectivity Journal Article In: eLife, vol. 10, pp. e68240, 2021. @article{Stauch2021, Under natural conditions, the visual system often sees a given input repeatedly. This provides an opportunity to optimize processing of the repeated stimuli. Stimulus repetition has been shown to strongly modulate neuronal-gamma band synchronization, yet crucial questions remained open. Here we used magnetoencephalography in 30 human subjects and find that gamma decreases across ≈10 repetitions and then increases across further repetitions, revealing plastic changes of the activated neuronal circuits. Crucially, increases induced by one stimulus did not affect responses to other stimuli, demonstrating stimulus specificity. Changes partially persisted when the inducing stimulus was repeated after 25 minutes of intervening stimuli. They were strongest in early visual cortex and increased interareal feedforward influences. Our results suggest that early visual cortex gamma synchronization enables adaptive neuronal processing of recurring stimuli. These and previously reported changes might be due to an interaction of oscillatory dynamics with established synaptic plasticity mechanisms. |
Chloé Stengel; Marine Vernet; Julià L. Amengual; Antoni Valero-Cabré In: Scientific Reports, vol. 11, pp. 3807, 2021. @article{Stengel2021, Correlational evidence in non-human primates has reported increases of fronto-parietal high-beta (22–30 Hz) synchrony during the top-down allocation of visuo-spatial attention. But may inter-regional synchronization at this specific frequency band provide a causal mechanism by which top-down attentional processes facilitate conscious visual perception? To address this question, we analyzed electroencephalographic (EEG) signals from a group of healthy participants who performed a conscious visual detection task while we delivered brief (4 pulses) rhythmic (30 Hz) or random bursts of Transcranial Magnetic Stimulation (TMS) to the right Frontal Eye Field (FEF) prior to the onset of a lateralized target. We report increases of inter-regional synchronization in the high-beta band (25–35 Hz) between the electrode closest to the stimulated region (the right FEF) and right parietal EEG leads, and increases of local inter-trial coherence within the same frequency band over bilateral parietal EEG contacts, both driven by rhythmic but not random TMS patterns. Such increases were accompained by improvements of conscious visual sensitivity for left visual targets in the rhythmic but not the random TMS condition. These outcomes suggest that high-beta inter-regional synchrony can be modulated non-invasively and that high-beta oscillatory activity across the right dorsal fronto-parietal network may contribute to the facilitation of conscious visual perception. Our work supports future applications of non-invasive brain stimulation to restore impaired visually-guided behaviors by operating on top-down attentional modulatory mechanisms. |
David W. Sutterer; Andrew J. Coia; Vincent Sun; Steven K. Shevell; Edward Awh Decoding chromaticity and luminance from patterns of EEG activity Journal Article In: Psychophysiology, vol. 58, no. 4, pp. e13779, 2021. @article{Sutterer2021, A long-standing question in the field of vision research is whether scalp-recorded EEG activity contains sufficient information to identify stimulus chromaticity. Recent multivariate work suggests that it is possible to decode which chromaticity an observer is viewing from the multielectrode pattern of EEG activity. There is debate, however, about whether the claimed effects of stimulus chromaticity on visual evoked potentials (VEPs) are instead caused by unequal stimulus luminances, which are achromatic differences. Here, we tested whether stimulus chromaticity could be decoded when potential confounds with luminance were minimized by (1) equating chromatic stimuli in luminance using heterochromatic flicker photometry for each observer and (2) independently varying the chromaticity and luminance of target stimuli, enabling us to test whether the pattern for a given chromaticity generalized across wide variations in luminance. We also tested whether luminance variations can be decoded from the topography of voltage across the scalp. In Experiment 1, we presented two chromaticities (appearing red and green) at three luminance levels during separate trials. In Experiment 2, we presented four chromaticities (appearing red, orange, yellow, and green) at two luminance levels. Using a pattern classifier and the multielectrode pattern of EEG activity, we were able to accurately decode the chromaticity and luminance level of each stimulus. Furthermore, we were able to decode stimulus chromaticity when we trained the classifier on chromaticities presented at one luminance level and tested at a different luminance level. Thus, EEG topography contains robust information regarding stimulus chromaticity, despite large variations in stimulus luminance. |
David W. Sutterer; Sean M. Polyn; Geoffrey F. Woodman a-Band activity tracks a two-dimensional spotlight of attention during spatial working memory maintenance Journal Article In: Journal of Neurophysiology, vol. 125, no. 3, pp. 957–971, 2021. @article{Sutterer2021a, Covert spatial attention is thought to facilitate the maintenance of locations in working memory, and EEG a-band activity (8–12Hz) is proposed to track the focus of covert attention. Recent work has shown that multivariate patterns of a-band activity track the polar angle of remembered locations relative to fixation. However, a defining feature of covert spatial attention is that it facilitates processing in a specific region of the visual field, and prior work has not determined whether patterns of a-band activity track the two-dimensional (2-D) coordinates of remembered stimuli within a visual hemifield or are instead maximally sensitive to the polar angle of remembered locations around fixation. Here, we used a lateralized spatial estimation task, in which observers remembered the location of one or two target dots presented to one side of fixation, to test this question. By applying a linear discriminant classifier to the topography of a-band activity, we found that we were able to decode the location of remembered stimuli. Critically, model comparison revealed that the pattern of classifier choices observed across remembered positions was best explained by a model assuming that a-band activity tracks the 2-D coordinates of remembered locations rather than a model assuming that a-band activity tracks the polar angle of remembered locations relative to fixation. These results support the hypothesis that this a-band activity is involved in the spotlight of attention, and arises from mid- to lower-level visual areas involved in maintaining spatial locations in working memory. NEW & NOTEWORTHY A substantial body of work has shown that patterns of EEG a-band activity track the angular coordinates of attended and remembered stimuli around fixation, but whether these patterns track the two-dimensional coordinates of stimuli presented within a visual hemifield remains an open question. Here, we demonstrate that a-band activity tracks the two-dimensional coordinates of remembered stimuli within a hemifield, showing that a-band activity reflects a spotlight of attention focused on locations maintained in working memory. |
Yu Takagi; Laurence Tudor Hunt; Mark W. Woolrich; Timothy E. J. Behrens; Miriam C. Klein-Flügge Adapting non-invasive human recordings along multiple task-axes shows unfolding of spontaneous and over-trained choice Journal Article In: eLife, vol. 10, pp. 1–27, 2021. @article{Takagi2021, Choices rely on a transformation of sensory inputs into motor responses. Using invasive single neuron recordings, the evolution of a choice process has been tracked by projecting population neural responses into state spaces. Here, we develop an approach that allows us to recover similar trajectories on a millisecond timescale in non-invasive human recordings. We selectively suppress activity related to three task-axes, relevant and irrelevant sensory inputs and response direction, in magnetoencephalography data acquired during context-dependent choices. Recordings from premotor cortex show a progression from processing sensory input to processing the response. In contrast to previous macaque recordings, information related to choice-irrelevant features is represented more weakly than choice-relevant sensory information. To test whether this mechanistic difference between species is caused by extensive over-training common in non-human primate studies, we trained humans on >20,000 trials of the task. Choice-irrelevant features were still weaker than relevant features in premotor cortex after over-training. |
Travis N. Talcott; Nicholas Gaspelin Eye movements are not mandatorily preceded by the N2pc component Journal Article In: Psychophysiology, vol. 58, no. 6, pp. e13821, 2021. @article{Talcott2021, Researchers typically distinguish between two mechanisms of attentional selection in vision: overt and covert attention. A commonplace assumption is that overt eye movements are automatically preceded by shifts of covert attention during visual search. Although the N2pc component is a putative index of covert attentional orienting, little is currently known about its relationship with overt eye movements. This is because most previous studies of the N2pc component prohibit overt eye movements. The current study assessed this relationship by concurrently measuring covert attention (via the N2pc) and overt eye movements (via eye tracking). Participants searched displays for a lateralized target stimulus and were allowed to generate overt eye movements during the search. We then assessed whether overt eye movements were preceded by the N2pc component. The results indicated that saccades were preceded by an N2pc component, but only when participants were required to carefully inspect the target stimulus before initiating the eye movement. When participants were allowed to make naturalistic eye movements in service of visual search, there was no evidence of an N2pc component before eye movements. These findings suggest that the N2pc component does not always precede overt eye movements during visual search. Implications for understanding the relationship between covert and overt attention are discussed. |
Wieske Zoest; Christoph Huber-Huber; Matthew D. Weaver; Clayton Hickey Strategic distractor suppression improves selective control in human vision Journal Article In: Journal of Neuroscience, vol. 41, no. 33, pp. 7120–7135, 2021. @article{Zoest2021, Our visual environment is complicated, and our cognitive capacity is limited. As a result, we must strategically ignore some stimuli to prioritize others. Common sense suggests that foreknowledge of distractor characteristics, like location or color, might help us ignore these objects. But empirical studies have provided mixed evidence, often showing that knowing about a distractor before it appears counterintuitively leads to its attentional selection. What has looked like strategic distractor suppression in the past is now commonly explained as a product of prior experience and implicit statistical learning, and the long-standing notion the distractor suppression is reflected in a band oscillatory brain activity has been challenged by results appearing to link a to target resolution. Can we strategically, proactively suppress distractors? And, if so, does this involve a? Here, we use the concurrent recording of human EEG and eye movements in optimized experimental designs to identify behavior and brain activity associated with proactive distractor suppression. Results from three experiments show that knowing about distractors before they appear causes a reduction in electrophysiological indices of covert attentional selection of these objects and a reduction in the overt deployment of the eyes to the location of the objects. This control is established before the distractor appears and is predicted by the power of cue-elicited a activity over the visual cortex. Foreknowledge of distractor characteristics therefore leads to improved selective control, and a oscillations in visual cortex reflect the implementation of this strategic, proactive mechanism. |
Seungji Lee; Doyoung Lee; Hyunjae Gil; Ian Oakley; Yang Seok Cho; Sung-Phil Kim Eye fixation-related potentials during visual search on acquaintance and newly-learned faces Journal Article In: Brain Sciences, vol. 11, no. 2, pp. 1–15, 2021. @article{Lee2021b, Searching familiar faces in the crowd may involve stimulus-driven attention by emotional significance, together with goal-directed attention due to task-relevant needs. The present study investigated the effect of familiarity on attentional processes by exploring eye fixation-related potentials (EFRPs) and eye gazes when humans searched for, among other distracting faces, either an acquaintance's face or a newly-learned face. Task performance and gaze behavior were indistinguishable for identifying either faces. However, from the EFRP analysis, after a P300 component for successful search of target faces, we found greater deflections of right parietal late positive potentials in response to newly-learned faces than acquaintance's faces, indicating more involvement of goaldirected attention in processing newly-learned faces. In addition, we found greater occipital negativity elicited by acquaintance's faces, reflecting emotional responses to significant stimuli. These results may suggest that finding a familiar face in the crowd would involve lower goal-directed attention and elicit more emotional responses. |
Cai S. Longman; Heike Elchlepp; Stephen Monsell; Aureliu Lavric Serial or parallel proactive control of components of task-set? A task-switching investigation with concurrent EEG and eye-tracking Journal Article In: Neuropsychologia, vol. 160, pp. 107984, 2021. @article{Longman2021, Among the issues examined by studies of cognitive control in multitasking is whether processes underlying performance in the different tasks occur serially or in parallel. Here we ask a similar question about processes that pro-actively control task-set. In task-switching experiments, several indices of task-set preparation have been extensively documented, including anticipatory orientation of gaze to the task-relevant location (an unambiguous marker of reorientation of attention), and a positive polarity brain potential over the posterior cortex (whose functional significance is less well understood). We examine whether these markers of preparation occur in parallel or serially, and in what order. On each trial a cue required participants to make a semantic classification of one of three digits presented simultaneously, with the location of each digit consistently associated with one of three classification tasks (e.g., if the task was odd/even, the digit at the top of the display was relevant). The EEG positivity emerged following, and appeared time-locked to, the anticipatory fixation on the task-relevant location, which might suggest serial organisation. However, the fixation-locked positivity was not better defined than the cue-locked positivity; in fact, for the trials with the earliest fixations the positivity was better time-locked to the cue onset. This is more consistent with (re)orientation of spatial attention occurring in parallel with, but slightly before, the reconfiguration of other task-set components indexed by the EEG positivity. |
Sara LoTemplio; Jack Silcox; Kara D. Federmeier; Brennan R. Payne Inter- and intra-individual coupling between pupillary, electrophysiological, and behavioral responses in a visual oddball task Journal Article In: Psychophysiology, vol. 58, no. 4, pp. e13758, 2021. @article{LoTemplio2021, Although the P3b component of the event-related brain potential is one of the most widely studied components, its underlying generators are not currently well understood. Recent theories have suggested that the P3b is triggered by phasic activation of the locus-coeruleus norepinephrine (LC-NE) system, an important control center implicated in facilitating optimal task-relevant behavior. Previous research has reported strong correlations between pupil dilation and LC activity, suggesting that pupil diameter is a useful indicator for ongoing LC-NE activity. Given the strong relationship between LC activity and pupil dilation, if the P3b is driven by phasic LC activity, there should be a robust trial-to-trial relationship with the phasic pupillary dilation response (PDR). However, previous work examining relationships between concurrently recorded pupillary and P3b responses has not supported this. One possibility is that the relationship between the measures might be carried primarily by either inter-individual (i.e., between-participant) or intra-individual (i.e., within-participant) contributions to coupling, and prior work has not systematically delineated these relationships. Doing so in the current study, we do not find evidence for either inter-individual or intra-individual relationships between the PDR and P3b responses. However, baseline pupil dilation did predict the P3b. Interestingly, both the PDR and P3b independently predicted inter-individual and intra-individual variability in decision response time. Implications for the LC-P3b hypothesis are discussed. |
Sarah D. McCrackin; Roxane J. Itier I can see it in your eyes: Perceived gaze direction impacts ERP and behavioural measures of affective theory of mind Journal Article In: Cortex, vol. 143, pp. 205–222, 2021. @article{McCrackin2021, Looking at someone's eyes is thought to be important for affective theory of mind (aTOM), our ability to infer their emotional state. However, it is unknown whether an individual's gaze direction influences our aTOM judgements and what the time course of this influence might be. We presented participants with sentences describing individuals in positive, negative or neutral scenarios, followed by direct or averted gaze neutral face pictures of those individuals. Participants made aTOM judgements about each person's mental state, including their affective valence and arousal, and we investigated whether the face gaze direction impacted those judgements. Participants rated that gazers were feeling more positive when they displayed direct gaze as opposed to averted gaze, and that they were feeling more aroused during negative contexts when gaze was averted as opposed to direct. Event-related potentials associated with face perception and affective processing were examined using mass-univariate analyses to track the time-course of this eye-gaze and affective processing interaction at a neural level. Both positive and negative trials were differentiated from neutral trials at many stages of processing. This included the early N200 and EPN components, believed to reflect automatic emotion areas activation and attentional selection respectively. This also included the later P300 and LPP components, thought to reflect elaborative cognitive appraisal of emotional content. Critically, sentence valence and gaze direction interacted over these later components, which may reflect the incorporation of eye-gaze in the cognitive evaluation of another's emotional state. The results suggest that gaze perception directly impacts aTOM processes, and that altered eye-gaze processing in clinical populations may contribute to associated aTOM impairments. |
Sarah D. McCrackin; Roxane J. Itier Feeling through another's eyes: Perceived gaze direction impacts ERP and behavioural measures of positive and negative affective empathy Journal Article In: NeuroImage, vol. 226, pp. 117605, 2021. @article{McCrackin2021a, Looking at the eyes informs us about the thoughts and emotions of those around us, and impacts our own emotional state. However, it is unknown how perceiving direct and averted gaze impacts our ability to share the gazer's positive and negative emotions, abilities referred to as positive and negative affective empathy. We presented 44 participants with contextual sentences describing positive, negative and neutral events happening to other people (e.g. “Her newborn was saved/killed/fed yesterday afternoon.”). These were designed to elicit positive, negative, or little to no empathy, and were followed by direct or averted gaze images of the individuals described. Participants rated their affective empathy for the individual and their own emotional valence on each trial. Event-related potentials time-locked to face-onset and associated with empathy and emotional processing were recorded to investigate whether they were modulated by gaze direction. Relative to averted gaze, direct gaze was associated with increased positive valence in the positive and neutral conditions and with increased positive empathy ratings. A similar pattern was found at the neural level, using robust mass-univariate statistics. The N100, thought to reflect an automatic activation of emotion areas, was modulated by gaze in the affective empathy conditions, with opposite effect directions in positive and negative conditions. The P200, an ERP component sensitive to positive stimuli, was modulated by gaze direction only in the positive empathy condition. Positive and negative trials were processed similarly at the early N200 processing stage, but later diverged, with only negative trials modulating the EPN, P300 and LPP components. These results suggest that positive and negative affective empathy are associated with distinct time-courses, and that perceived gaze direction uniquely modulates positive empathy, highlighting the importance of studying empathy with face stimuli. |
Amir H. Meghdadi; Barry Giesbrecht; Miguel P. Eckstein EEG signatures of contextual influences on visual search with real scenes Journal Article In: Experimental Brain Research, vol. 239, no. 3, pp. 797–809, 2021. @article{Meghdadi2021, The use of scene context is a powerful way by which biological organisms guide and facilitate visual search. Although many studies have shown enhancements of target-related electroencephalographic activity (EEG) with synthetic cues, there have been fewer studies demonstrating such enhancements during search with scene context and objects in real world scenes. Here, observers covertly searched for a target in images of real scenes while we used EEG to measure the steady state visual evoked response to objects flickering at different frequencies. The target appeared in its typical contextual location or out of context while we controlled for low-level properties of the image including target saliency against the background and retinal eccentricity. A pattern classifier using EEG activity at the relevant modulated frequencies showed target detection accuracy increased when the target was in a contextually appropriate location. A control condition for which observers searched the same images for a different target orthogonal to the contextual manipulation, resulted in no effects of scene context on classifier performance, confirming that image properties cannot explain the contextual modulations of neural activity. Pattern classifier decisions for individual images were also related to the aggregated observer behavioral decisions for individual images. Together, these findings demonstrate target-related neural responses are modulated by scene context during visual search with real world scenes and can be related to behavioral search decisions. |
Michael Christopher Melnychuk; Ian H. Robertson; Emanuele R. G. Plini; Paul M. Dockree In: Brain Sciences, vol. 11, pp. 1324, 2021. @article{Melnychuk2021, Yogic and meditative traditions have long held that the fluctuations of the breath and the mind are intimately related. While respiratory modulation of cortical activity and attentional switching are established, the extent to which electrophysiological markers of attention exhibit synchronization with respiration is unknown. To this end, we examined (1) frontal midline theta-beta ratio (TBR), an indicator of attentional control state known to correlate with mind wandering episodes and functional connectivity of the executive control network; (2) pupil diameter (PD), a known proxy measure of locus coeruleus (LC) noradrenergic activity; and (3) respiration for evidence of phase synchronization and information transfer (multivariate Granger causality) during quiet restful breathing. Our results indicate that both TBR and PD are simultaneously synchronized with the breath, suggesting an underlying oscillation of an attentionally relevant electrophysiological index that is phase-locked to the respiratory cycle which could have the potential to bias the attentional system into switching states. We highlight the LC's pivotal role as a coupling mechanism between respiration and TBR, and elaborate on its dual functions as both a chemosensitive respiratory nucleus and a pacemaker of the attentional system. We further suggest that an appreciation of the dynamics of this weakly coupled oscillatory system could help deepen our understanding of the traditional claim of a relationship between breathing and attention. |
Yali Pan; Steven Frisson; Ole Jensen Neural evidence for lexical parafoveal processing Journal Article In: Nature Communications, vol. 12, pp. 5234, 2021. @article{Pan2021a, In spite of the reduced visual acuity, parafoveal information plays an important role in natural reading. However, competing models on reading disagree on whether words are previewed parafoveally at the lexical level. We find neural evidence for lexical parafoveal processing by combining a rapid invisible frequency tagging (RIFT) approach with magnetoencephalography (MEG) and eye-tracking. In a silent reading task, target words are tagged (flickered) subliminally at 60 Hz. The tagging responses measured when fixating on the pre-target word reflect parafoveal processing of the target word. We observe stronger tagging responses during pre-target fixations when followed by low compared with high lexical frequency targets. Moreover, this lexical parafoveal processing is associated with individual reading speed. Our findings suggest that reading unfolds in the fovea and parafovea simultaneously to support fluent reading. |