EEGManyPipelines: A Large-scale, Grassroots Multi-analyst Study of Electroencephalography Analysis Practices in the Wild.

The ongoing reproducibility crisis in psychology and cognitive neuroscience has sparked increasing calls to re-evaluate and reshape scientific culture and practices. Heeding those calls, we have recently launched the EEGManyPipelines project as a means to assess the robustness of EEG research in naturalistic conditions and experiment with an alternative model of conducting scientific research. One hundred sixty-eight analyst teams, encompassing 396 individual researchers from 37 countries, independently analyzed the same unpublished, representative EEG data set to test the same set of predefined hypotheses and then provided their analysis pipelines and reported outcomes. Here, we lay out how large-scale scientific projects can be set up in a grassroots, community-driven manner without a central organizing laboratory. We explain our recruitment strategy, our guidance for analysts, the eventual outputs of this project, and how it might have a lasting impact on the field.

Spontaneous Alpha-Band Oscillations Bias Subjective Contrast Perception.

Perceptual decisions depend both on the features of the incoming stimulus and on the ongoing brain activity at the moment the stimulus is received. Specifically, trial-to-trial fluctuations in cortical excitability have been linked to fluctuations in the amplitude of prestimulus α oscillations (∼8-13 Hz), which are in turn are associated with fluctuations in subjects' tendency to report the detection of a stimulus. It is currently unknown whether α oscillations bias postperceptual decision-making, or even bias subjective perception itself. To answer this question, we used a contrast discrimination task in which both male and female human subjects reported which of two gratings (one in each hemifield) was perceived as having a stronger contrast. Our EEG analysis showed that subjective contrast was reduced for the stimulus in the hemifield represented in the hemisphere with relatively stronger prestimulus α amplitude, reflecting reduced cortical excitability. Furthermore, the strength of this spontaneous hemispheric lateralization was strongly correlated with the magnitude of individual subjects' biases, suggesting that the spontaneous patterns of α lateralization play a role in explaining the intersubject variability in contrast perception. These results indicate that spontaneous fluctuations in cortical excitability, indicated by patterns of prestimulus α amplitude, affect perceptual decisions by altering the phenomenological perception of the visual world.Significance Statement:Our moment-to-moment perception of the world is shaped by the features of the environment surrounding us, as much as by the constantly evolving states that characterize our brain activity. Previous research showed how the ongoing electrical activity of the brain can influence whether a stimulus has accessed conscious perception. However, evidence is currently missing on whether these electrical brain states can be associated to the subjective experience of a sensory input. Here we show that local changes in patterns of electrical brain activity preceding visual stimulation can bias our phenomenological perception. Importantly, we show that the strength of these variations can help explain the great interindividual variability in how we perceive the visual environment surrounding us.

Covid-19 affects taste independent of taste-smell confusions: results from a combined chemosensory home test and online survey from a large global cohort.

People often confuse smell loss with taste loss, so it is unclear how much gustatory function is reduced in patients self-reporting taste loss. Our pre-registered cross-sectional study design included an online survey in 12 languages with instructions for self-administering chemosensory tests with 10 household items. Between June 2020 and March 2021, 10,953 individuals participated. Of these, 5,225 self-reported a respiratory illness and were grouped based on their reported COVID test results: COVID-positive (COVID+, N = 3,356), COVID-negative (COVID-, N = 602), and COVID unknown for those waiting for a test result (COVID?, N = 1,267). The participants who reported no respiratory illness were grouped by symptoms: sudden smell/taste changes (STC, N = 4,445), other symptoms excluding smell or taste changes (OthS, N = 832), and no symptoms (NoS, N = 416). Taste, smell, and oral irritation intensities and self-assessed abilities were rated on visual analog scales. Compared to the NoS group, COVID+ was associated with a 21% reduction in taste (95% confidence interval (CI): 15-28%), 47% in smell (95% CI: 37-56%), and 17% in oral irritation (95% CI: 10-25%) intensity. There were medium to strong correlations between perceived intensities and self-reported abilities (r = 0.84 for smell, r = 0.68 for taste, and r = 0.37 for oral irritation). Our study demonstrates that COVID-19-positive individuals report taste dysfunction when self-tested with stimuli that have little to none olfactory components. Assessing the smell and taste intensity of household items is a promising, cost-effective screening tool that complements self-reports and may help to disentangle taste loss from smell loss. However, it does not replace standardized validated psychophysical tests.

Timing and kinematics of horizontal within-blink saccades measured by EOG.

Eyeblinks are the brief closures of the lid. They are accompanied by a cocontraction of the eye muscles that temporarily pulls the whole eyeball back into its socket. When blinks occur together with execution of saccadic gaze shifts, they interfere with the saccadic premotor circuit, causing these within-blink saccades to be slower than normal and also time-locked to blinks. To analyze the trajectory of within-blink saccades, subtraction of the entangled blink-related eye movement is required. Here we propose a combination of principal component analysis (PCA) and a regression model to subtract the blink-related component of the eye movement based on the respective blink metrics. We used electrooculography (EOG) to measure eye and lid movements of 12 participants who performed saccades with and without blinks. We found that within-blink saccades are slower than without-blink saccades and are tightly coupled in time to blink onset. Surprisingly, in some participants we observed large dynamic overshoots of up to 15° for saccades of only 5° amplitude. The finding of dynamic overshoots was independently confirmed by dynamic MRI for two of the participants and challenges the current view that within-blink saccades are programmed as slow, but straight, saccades. We hypothesize that the dynamic overshoots could be attributed to inhibition of omnipause neurons during blinks, the simultaneous cocontraction of extraocular muscles, or a combination of both.NEW & NOTEWORTHY This study observed that people make large dynamic overshoots when making a saccadic eye movement within a blink but their eyes are back on target by the time the eyelids are open. We used electrooculography (EOG) to measure eye movements even when the lid is down and introduced a novel procedure to subtract blink-related EOG components. These findings challenge the current view that within-blink saccades are programmed as slow but straight saccades.

Distinct contributions of alpha and theta rhythms to perceptual and attentional sampling.

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.

Testing the effect of tACS over parietal cortex in modulating endogenous alpha rhythm and temporal integration windows in visual perception.

Neural oscillations in the alpha band (8-12 Hz) have been proposed as a key mechanism for the temporal resolution of visual perception. Higher alpha frequencies have been related to improved segregation of visual events over time, whereas lower alpha frequencies have been related to improved temporal integration. Similarly, also the phase of ongoing alpha has been shown to correlate with temporal integration/segregation. To test a causal relationship between alpha oscillations and perception, we here employed multi-channel transcranial alternating current stimulation (mc-tACS) over the right parietal cortex, whereas participants performed a visual temporal integration/segregation task that used identical stimuli with different instructions. Before and after mc-tACS we recorded the resting-state electroencephalogram (EEG) to extract the individual alpha frequency (IAF) and delivered electrical stimulation at slightly slower and faster frequencies (IAF±2 Hz). We hypothesized that this would not only drive endogenous alpha rhythms, but also affect temporal integration and segregation in an opposite way. However, the mc-tACS protocol used here did not consistently increase or decrease the IAF after the stimulation and did not affect temporal integration/segregation accuracy as expected. Although we found some preliminary evidence for an influence of tACS phase on temporal integration accuracy, the ongoing phase of mc-tACS oscillations did not reliably modulate temporal integration/segregation accuracy in a sinusoidal way as would have been predicted by an effective entrainment of brain oscillations. These findings may guide future studies using different stimulation montages for investigating the role of cortical alpha oscillations for human vision.

The extent and specificity of visual exploration determines the formation of recollected memories in complex scenes.

Our visual memories of complex scenes often appear as robust, detailed records of the past. Several studies have demonstrated that active exploration with eye movements improves recognition memory for scenes, but it is unclear whether this improvement is due to stronger feelings of familiarity or more detailed recollection. We related the extent and specificity of fixation patterns at encoding and retrieval to different recognition decisions in an incidental memory paradigm. After incidental encoding of 240 real-world scene photographs, participants (N = 44) answered a surprise memory test by reporting whether an image was new, remembered (indicating recollection), or just known to be old (indicating familiarity). To assess the specificity of their visual memories, we devised a novel report procedure in which participants selected the scene region that they specifically recollected, that appeared most familiar, or that was particularly new to them. At encoding, when considering the entire scene,subsequently recollected compared to familiar or forgotten scenes showed a larger number of fixations that were more broadly distributed, suggesting that more extensive visual exploration determines stronger and more detailed memories. However, when considering only the memory-relevant image areas, fixations were more dense and more clustered for subsequently recollected compared to subsequently familiar scenes. At retrieval, the extent of visual exploration was more restricted for recollected compared to new or forgotten scenes, with a smaller number of fixations. Importantly, fixation density and clustering was greater in memory-relevant areas for recollected versus familiar or falsely recognized images. Our findings suggest that more extensive visual exploration across the entire scene, with a subset of more focal and dense fixations in specific image areas, leads to increased potential for recollecting specific image aspects.

Failed Suppression of Salient Stimuli Precedes Behavioral Errors.

Our visual system is constantly confronted with more information than it can process. To deal with the limited capacity, attention allows us to enhance relevant information and suppress irrelevant information. Particularly, the suppression of salient irrelevant stimuli has shown to be important as it prevents attention to be captured and thus attentional resources to be wasted. This study aimed at directly connecting failures to suppress distraction with a neural marker of suppression, the distractor positivity (Pd). We measured participants' EEG signal while they performed a visual search task in which they had to report a digit inside a shape target while ignoring distractors, one of which could be a salient color singleton. Reports of target digits served as a behavioral index of enhancement, and reports of color distractor digits served as a behavioral index of failed suppression, each measured against reports of neutral distractor digits serving as a baseline. Participants reported the target identity more often than any distractor identity. The singleton identity was reported least often, suggesting suppression of the singleton below baseline. Suppression of salient stimuli was absent in the beginning and then increased throughout the experiment. When the singleton identity was reported, the Pd was observed in a later time window, suggesting that behavioral errors were preceded by failed suppression. Our results provide evidence for the signal suppression hypothesis that states salient items have to be actively suppressed to avoid attentional capture. Our results also provide direct evidence that the Pd is reflecting such active suppression.

Attention explores space periodically at the theta frequency.

Voluntary attention is at the core of a wide variety of cognitive functions. Attention can be oriented to and sustained at a location or reoriented in space to allow processing at other locations-critical in an ever-changing environment. Numerous studies have investigated attentional orienting in time and space, but little is known about the spatiotemporal dynamics of attentional reorienting. Here we explicitly manipulated attentional reorienting using a cuing procedure in a two-alternative forced-choice orientation-discrimination task. We interrogated attentional distribution by flashing two probe stimuli with various delays between the precue and target stimuli. Then we used the probabilities that both probes and neither probe were correctly reported to solve a second-degree equation, which estimates the report probability at each probe location. We demonstrated that attention reorients periodically at ∼4 Hz (theta) between the two stimulus locations. We further characterized the processing dynamics at each stimulus location, and demonstrated that attention samples each location periodically at ∼11 Hz (alpha). Finally, simulations support our findings and show that this method is sufficiently powered, making it a valuable tool for studying the spatiotemporal dynamics of attention.

Spontaneous Neural Oscillations Bias Perception by Modulating Baseline Excitability.

The brain exhibits organized fluctuations of neural activity, even in the absence of tasks or sensory input. A prominent type of such spontaneous activity is the alpha rhythm, which influences perception and interacts with other ongoing neural activity. It is currently hypothesized that states of decreased prestimulus α oscillations indicate enhanced neural excitability, resulting in improved perceptual acuity. Nevertheless, it remains debated how changes in excitability manifest at the behavioral level in perceptual tasks. We addressed this issue by comparing two alternative models describing the effect of spontaneous α power on signal detection. The first model assumes that decreased α power increases baseline excitability, amplifying the response to both signal and noise, predicting a liberal detection criterion with no effect on sensitivity. The second model predicts that decreased α power increases the trial-by-trial precision of the sensory response, resulting in improved sensitivity. We tested these models in two EEG experiments in humans where we analyzed the effects of prestimulus α power on visual detection and discrimination using a signal detection framework. Both experiments provide strong evidence that decreased α power reflects a more liberal detection criterion, rather than improved sensitivity, consistent with the baseline model. In other words, when the task requires detecting stimulus presence versus absence, reduced α oscillations make observers more likely to report the stimulus regardless of actual stimulus presence. Contrary to previous interpretations, these results suggest that states of decreased α oscillations increase the global baseline excitability of sensory systems without affecting perceptual acuity. SIGNIFICANCE STATEMENT: Spontaneous fluctuations of brain activity explain why a faint sensory stimulus is sometimes perceived and sometimes not. The prevailing view is that heightened neural excitability, indexed by decreased α oscillations, promotes better perceptual performance. Here, we provide evidence that heightened neural excitability instead reflects a state of biased perception, during which a person is more likely to see a stimulus, whether or not it is actually present. Therefore, we propose that changes in neural excitability leave the precision of sensory processing unaffected. These results establish the link between spontaneous brain activity and the variability in human perception.

The role of alpha oscillations in distractor inhibition during memory retention.

Only small amounts of visual information, as determined by the capacity of working memory, can be held in an active and accessible state. Thus, it is important to select and maintain information that is relevant while ignoring irrelevant information. However, the underlying neural mechanism of these processes has yet to be identified. One potential candidate are alpha oscillations (8-14 Hz), which have been shown to inhibit stimulus processing in perceptual tasks. During memory maintenance, alpha power increases with set size suggesting that alpha oscillations are involved either in memory maintenance or in the inhibition of task-irrelevant information to protect relevant information from interference. The need for such a protection should increase with the amount of distracting information, but most previous studies did not show any distractors. Therefore, we directly tested whether alpha oscillations are involved in inhibition of distractors during memory maintenance. Participants memorized the orientation of one or two target lines embedded among irrelevant distractors. Distractors were either strong or weak and were present during the retention interval after which participants reported the orientation of probed targets. Computational modeling showed that performance decreased with increasing set size and stronger distraction. Alpha power in the retention interval generally increased with set size, replicating previous studies. However, here stronger distractors reduced alpha power. This finding is in clear contradistinction to previous suggestions, as alpha power decrease indicates higher neuronal excitability. Thus, our data do not support the suggested role of alpha oscillations in inhibition of distraction in working memory.

Early visual processing allows for selective behavior, shifts of attention, and conscious visual experience in spite of masking.

Object-substitution masking (OSM) occurs when a briefly displayed target in a search array is surrounded by a mask, which remains onscreen after the target has disappeared. It has been suggested that OSM results from a specific interference with reentrant visual processing, while the initial feedforward processing is left intact. Here, we tested the prediction that the fastest saccadic responses towards a masked target, supposedly triggered before the onset of reentrant processing, are not impaired by OSM. Indeed, saccades faster than 350ms "escaped" the influence of the mask. Notably, participants' judgements of subjective awareness indicated that stimulus processing during this early stage is not entirely devoid of conscious awareness. Furthermore, the N2pc event-related potential component indicated shifts of spatial attention towards the masked targets on trials with correct fast saccades, suggesting that both target detection and spatial attention can be based on the computations accomplished during the initial feedforward sweep.

Semantic Relations between Visual Objects Can Be Unconsciously Processed but Not Reported under Change Blindness.

Change blindness-the failure to detect changes in visual scenes-has often been interpreted as a result of impoverished visual information encoding or as a failure to compare the prechange and postchange scene. In the present electroencephalography study, we investigated whether semantic features of prechange and postchange information are processed unconsciously, even when observers are unaware that a change has occurred. We presented scenes composed of natural objects in which one object changed from one presentation to the next. Object changes were either semantically related (e.g., rail car changed to rail) or unrelated (e.g., rail car changed to sausage). Observers were first asked to detect whether any change had occurred and then to judge the semantic relation of the two objects involved in the change. We found a semantic mismatch ERP effect, that is, a more negative-going ERP for semantically unrelated compared to related changes, originating from a cortical network including the left middle temporal gyrus and occipital cortex and resembling the N400 effect, albeit at longer latencies. Importantly, this semantic mismatch effect persisted even when observers were unaware of the change and the semantic relationship of prechange and postchange object. This finding implies that change blindness does not preclude the encoding of the prechange and postchange objects' identities and possibly even the comparison of their semantic content. Thus, change blindness cannot be interpreted as resulting from impoverished or volatile visual representations or as a failure to process the prechange and postchange object. Instead, change detection appears to be limited at a later, postperceptual stage.

Flicker-Induced Time Dilation Does Not Modulate EEG Correlates of Temporal Encoding.

In this study, we used EEG to investigate how visual stimulus dynamics (i.e. flicker) affect the mechanisms of duration perception. Previous studies have demonstrated that flickering visual stimuli are judged longer than equally long non-flickering stimuli. We tested whether this effect of flicker on duration judgments is mediated by changes in temporal encoding during the time interval. Here, temporal encoding refers to the perception of the unfolding of time throughout the temporal interval, also termed the "clock stage" in information processing models of interval timing. We hypothesized that if flicker mediates duration perception by affecting temporal encoding, then the dilation-effect should be reflected by neural correlates of temporal encoding. We presented flickering and steady stimuli in a duration bisection task and found that flicker dilated perceived duration. The EEG analysis allowed us to isolate a putative neural correlate of temporal encoding: a modulation of the amplitude of the contingent negative variation (CNV) by stimuli classified as "long" compared to physically identical stimuli classified as "short". However, flicker did not affect the CNV amplitude, suggesting that flicker does not dilate perceived duration by affecting temporal encoding. Possibly, flicker might affect only later stages of temporal processing such as interval comparison or decision making.

The state of a central inhibition system predicts access to visual targets: An ERP study on distractor-induced blindness (DIB).

Distractor-induced blindness (DIB) is elicited in a temporal selection task based on rapid serial visual presentation: Following the onset of a central cue (color change of fixation), a target in a global stream has to be detected (orientation change of tilted bars). Distractors are occasional target-like events presented prior to the onset of the cue. Depending on the number of distractor episodes, a frontal ERP negativity (FN) is evoked, and the detection rate of the forthcoming target is reduced. Here, we provide a concise review of the DIBs' functional characteristics, and examine the relationship between FN activation on target processing and detectability. To this end, ERP responses collected in a set of participants (n=14) were analysed separately with respect to detection performance (hits vs. misses). A gradual increase in the amplitude of the FN with increasing number of distractor events was only observed for forthcoming misses, but not for hits. The FN activation can be linked to a target-related ERP component: The amplitude of a late centro-parietal positivity (LPC) was significantly diminished for misses. In sum, the data support the notion that the DIB reflects the controlled activation of a central inhibition system. Depending on its state of activation, conscious access to stimuli defined by distinctive visual features is restricted.

Prestimulus neural oscillations inhibit visual perception via modulation of response gain.

The ongoing state of the brain radically affects how it processes sensory information. How does this ongoing brain activity interact with the processing of external stimuli? Spontaneous oscillations in the alpha range are thought to inhibit sensory processing, but little is known about the psychophysical mechanisms of this inhibition. We recorded ongoing brain activity with EEG while human observers performed a visual detection task with stimuli of different contrast intensities. To move beyond qualitative description, we formally compared psychometric functions obtained under different levels of ongoing alpha power and evaluated the inhibitory effect of ongoing alpha oscillations in terms of contrast or response gain models. This procedure opens the way to understanding the actual functional mechanisms by which ongoing brain activity affects visual performance. We found that strong prestimulus occipital alpha oscillations-but not more anterior mu oscillations-reduce performance most strongly for stimuli of the highest intensities tested. This inhibitory effect is best explained by a divisive reduction of response gain. Ongoing occipital alpha oscillations thus reflect changes in the visual system's input/output transformation that are independent of the sensory input to the system. They selectively scale the system's response, rather than change its sensitivity to sensory information.

The influence of spontaneous brain oscillations on apparent motion perception.

A good example of inferential processes in perception is long-range apparent motion (AM), the illusory percept of visual motion that occurs when two spatially distinct stationary visual objects are presented in alternating sequence. The AM illusion is strongest at presentation frequencies around 3 Hz. At lower presentation frequencies, the percept varies from trial to trial between AM and sequential alternation, while at higher frequencies perception varies between AM and two simultaneously flickering objects. Previous studies have demonstrated that prestimulus alpha oscillations explain trial-to-trial variability in detection performance for visual stimuli presented at threshold. In the present study, we investigated whether fluctuations of prestimulus alpha oscillations can also account for variations in AM perception. Prestimulus alpha power was stronger when observers reported AM perception in subsequent trials with low presentation frequencies, while at high presentation frequencies there were no significant differences in alpha power preceding AM and veridical flicker perception. Moreover, when observers perceived AM the prestimulus functional connectivity between frontal and occipital channels was increased in the alpha band, as revealed by the imaginary part of coherency, which is insensitive to artefacts from volume conduction. Dynamic causal modelling of steady-state responses revealed that the most likely direction of this fronto-occipital connectivity was from frontal to occipital sources. These results point to a role of ongoing alpha oscillations in the inferential process that gives rise to the perception of AM and suggest that fronto-occipital interactions bias perception towards internally generated predictions.

Imagining triadic interactions simultaneously activates mirror and mentalizing systems.

Coordinated triadic interactions, involving oneself, another person, and an external object, are considered a uniquely human skill. However, the exact mechanisms underlying the ability to engage in such social interactions remain hitherto unknown. We used functional neuroimaging to investigate the neural signature of triadic interactions. For this purpose, participants viewed pictures of objects in a 3T functional magnetic resonance imaging scanner and were asked whether they could imagine this object in a social interaction with another person. We also aimed to dissociate this process from, as well as to find commonalities with, purely self-referential or other-referential processing. In all trial-types, we found activations in core mentalizing brain areas (medial prefrontal cortex, posterior cingulate cortex, precuneus and temporoparietal junction). Furthermore, triadic engagements, but not self-referential or other-referential processing, were associated with activations in classical mirror neuron areas (inferior frontal gyrus and inferior parietal lobe). Finally, mentalizing networks showed a strong functional connectivity with mirror neuron areas exclusively during triadic engagements. These results suggest that the imagined interaction of two agents is processed in a more complex neural social cognitive network than purely self- or other-referential considerations.

Invisible visual stimuli elicit increases in alpha-band power.

The cerebral cortex responds to stimuli of a wide range of intensities. Previous studies have demonstrated that undetectably weak somatosensory stimuli cause a functional deactivation or inhibition in somatosensory cortex. In the present study, we tested whether invisible visual stimuli lead to similar responses, indicated by an increase in EEG alpha-band power-an index of cortical excitability. We presented subliminal and supraliminal visual stimuli after estimating each participant's detection threshold. Stimuli consisted of peripherally presented small circular patches that differed in their contrast to a background consisting of a random white noise pattern. We demonstrate that subliminal and supraliminal stimuli each elicit specific neuronal response patterns. Supraliminal stimuli evoked an early, strongly phase-locked lower-frequency response representing the evoked potential and induced a decrease in alpha-band power from 400 ms on. By contrast, subliminal visual stimuli induced an increase of non-phase-locked power around 300 ms that was maximal within the alpha-band. This response might be due to an inhibitory mechanism, which reduces spurious visual activation that is unlikely to result from external stimuli.