Characterizing endogenous delta oscillations in human MEG.

Rhythmic activity in the delta frequency range (0.5-3 Hz) is a prominent feature of brain dynamics. Here, we examined whether spontaneous delta oscillations, as found in invasive recordings in awake animals, can be observed in non-invasive recordings performed in humans with magnetoencephalography (MEG). In humans, delta activity is commonly reported when processing rhythmic sensory inputs, with direct relationships to behaviour. However, rhythmic brain dynamics observed during rhythmic sensory stimulation cannot be interpreted as an endogenous oscillation. To test for endogenous delta oscillations we analysed human MEG data during rest. For comparison, we additionally analysed two conditions in which participants engaged in spontaneous finger tapping and silent counting, arguing that internally rhythmic behaviours could incite an otherwise silent neural oscillator. A novel set of analysis steps allowed us to show narrow spectral peaks in the delta frequency range in rest, and during overt and covert rhythmic activity. Additional analyses in the time domain revealed that only the resting state condition warranted an interpretation of these peaks as endogenously periodic neural dynamics. In sum, this work shows that using advanced signal processing techniques, it is possible to observe endogenous delta oscillations in non-invasive recordings of human brain dynamics.

The Blursday database as a resource to study subjective temporalities during COVID-19.

The COVID-19 pandemic and associated lockdowns triggered worldwide changes in the daily routines of human experience. The Blursday database provides repeated measures of subjective time and related processes from participants in nine countries tested on 14 questionnaires and 15 behavioural tasks during the COVID-19 pandemic. A total of 2,840 participants completed at least one task, and 439 participants completed all tasks in the first session. The database and all data collection tools are accessible to researchers for studying the effects of social isolation on temporal information processing, time perspective, decision-making, sleep, metacognition, attention, memory, self-perception and mindfulness. Blursday includes quantitative statistics such as sleep patterns, personality traits, psychological well-being and lockdown indices. The database provides quantitative insights on the effects of lockdown (stringency and mobility) and subjective confinement on time perception (duration, passage of time and temporal distances). Perceived isolation affects time perception, and we report an inter-individual central tendency effect in retrospective duration estimation.

Implicit Versus Explicit Timing-Separate or Shared Mechanisms?

Time implicitly shapes cognition, but time is also explicitly represented, for instance, in the form of durations. Parsimoniously, the brain could use the same mechanisms for implicit and explicit timing. Yet, the evidence has been equivocal, revealing both joint versus separate signatures of timing. Here, we directly compared implicit and explicit timing using magnetoencephalography, whose temporal resolution allows investigating the different stages of the timing processes. Implicit temporal predictability was induced in an auditory paradigm by a manipulation of the foreperiod. Participants received two consecutive task instructions: discriminate pitch (indirect measure of implicit timing) or duration (direct measure of explicit timing). The results show that the human brain efficiently extracts implicit temporal statistics of sensory environments, to enhance the behavioral and neural responses to auditory stimuli, but that those temporal predictions did not improve explicit timing. In both tasks, attentional orienting in time during predictive foreperiods was indexed by an increase in alpha power over visual and parietal areas. Furthermore, pretarget induced beta power in sensorimotor and parietal areas increased during implicit compared to explicit timing, in line with the suggested role for beta oscillations in temporal prediction. Interestingly, no distinct neural dynamics emerged when participants explicitly paid attention to time, compared to implicit timing. Our work thus indicates that implicit timing shapes the behavioral and sensory response in an automatic way and is reflected in oscillatory neural dynamics, whereas the translation of implicit temporal statistics to explicit durations remains somewhat inconclusive, possibly because of the more abstract nature of this task.

Endogenous modulation of delta phase by expectation-A replication of Stefanics et al., 2010.

The human brain efficiently extracts the temporal statistics of sensory environments and automatically generates expectations about future events. An influential Hypothesis holds that these expectations can find their implementation in neural oscillations, notably in the delta band (.5-3 Hz). Rhythmic fluctuations of cortical excitement are thought to align and match up in phase to the temporal structure of the sensory environment. This alignment is thought to result in the more excitable phase range of neural oscillations to overlap with the predicted onset of sensory events which in turn results in more efficient processing of sensory input, especially so in audition. An unresolved issue concerns whether such phase-aligned rhythmic brain activity is driven exclusively by the exogenous temporal structure of the input, or whether it also reflects phase re-alignment due to endogenous expectations based on stimulus probability and task relevance. In a seminal study, Stefanics et al. (2010) presented stimuli in a rhythmic stream and observed that delta phase consistency across trials was modulated by endogenous target onset expectations: delta phase consistency was higher prior to more probable (strongly expected) compared to less probable (weakly expected) target onsets. The present study replicates Experiment II of the original study, most importantly the modulation of delta phase consistency by endogenous expectations, and underlines a direct relationship between phase locking and behaviour. Our additional analyses locate the sources of the delta phase-alignment to motor, pre-motor, parietal, and temporal areas, and provide evidence for an ongoing delta oscillation, in line with the interpretation of oscillatory phase alignment rather than a transient evoked response. Importantly, this work shows that the phase of delta oscillations can be modulated by top-down control, and hence qualifies as a potential mechanism for the neural implementation of (rhythmic) temporal predictions.

Good scientific practice in EEG and MEG research: Progress and perspectives.

Good scientific practice (GSP) refers to both explicit and implicit rules, recommendations, and guidelines that help scientists to produce work that is of the highest quality at any given time, and to efficiently share that work with the community for further scrutiny or utilization. For experimental research using magneto- and electroencephalography (MEEG), GSP includes specific standards and guidelines for technical competence, which are periodically updated and adapted to new findings. However, GSP also needs to be regularly revisited in a broader light. At the LiveMEEG 2020 conference, a reflection on GSP was fostered that included explicitly documented guidelines and technical advances, but also emphasized intangible GSP: a general awareness of personal, organizational, and societal realities and how they can influence MEEG research. This article provides an extensive report on most of the LiveMEEG contributions and new literature, with the additional aim to synthesize ongoing cultural changes in GSP. It first covers GSP with respect to cognitive biases and logical fallacies, pre-registration as a tool to avoid those and other early pitfalls, and a number of resources to enable collaborative and reproducible research as a general approach to minimize misconceptions. Second, it covers GSP with respect to data acquisition, analysis, reporting, and sharing, including new tools and frameworks to support collaborative work. Finally, GSP is considered in light of ethical implications of MEEG research and the resulting responsibility that scientists have to engage with societal challenges. Considering among other things the benefits of peer review and open access at all stages, the need to coordinate larger international projects, the complexity of MEEG subject matter, and today's prioritization of fairness, privacy, and the environment, we find that current GSP tends to favor collective and cooperative work, for both scientific and for societal reasons.

Target enhancement or distractor suppression? Functionally distinct alpha oscillations form the basis of attention.

Recent advances in attention research have been propelled by the debate on target enhancement versus distractor suppression. A predominant neural correlate of attention is the modulation of alpha oscillatory power (~10 Hz), which signifies shifts of attention in time, space and between sensory modalities. However, the underspecified functional role of alpha oscillations limits the progress of tracking down the neurocognitive basis of attention. In this short opinion article, we review and critically examine a synthesis of three conceptual and methodological aspects that are indispensable for a mechanistic understanding of the role of alpha oscillations for attention. (a) Precise mapping of the anatomical source and the temporal response profile of neural signals reveals distinct alpha oscillatory processes that implement facilitatory versus suppressive components of attention. (b) A testable framework enables unanimous association of alpha modulation with either target enhancement or different forms of distractor suppression (active vs. automatic). (c) Linking anatomically specified alpha oscillations to behavior reveals the causal nature of alpha oscillations for attention. The three reviewed aspects substantially enrich study design, data analysis and interpretation of results to achieve the goal of understanding how anatomically specified and functionally relevant neural oscillations contribute to the implementation of facilitatory versus suppressive components of attention.

Implicit temporal predictability enhances pitch discrimination sensitivity and biases the phase of delta oscillations in auditory cortex.

Can human listeners use implicit temporal contingencies in auditory input to form temporal predictions, and if so, how are these predictions represented endogenously? To assess this question, we implicitly manipulated temporal predictability in an auditory pitch discrimination task: unbeknownst to participants, the pitch of the standard tone could either be deterministically predictive of the temporal onset of the target tone, or convey no predictive information. Predictive and non-predictive conditions were presented interleaved in one stream, and separated by variable inter-stimulus intervals such that there was no dominant stimulus rhythm throughout. Even though participants were unaware of the implicit temporal contingencies, pitch discrimination sensitivity (the slope of the psychometric function) increased when the onset of the target tone was predictable in time (N = 49, 28 female, 21 male). Concurrently recorded EEG data (N = 24) revealed that standard tones that conveyed temporal predictions evoked a more negative N1 component than non-predictive standards. We observed no significant differences in oscillatory power or phase coherence between conditions during the foreperiod. Importantly, the phase angle of delta oscillations (1-3 Hz) in auditory areas in the post-standard and pre-target time windows predicted behavioral pitch discrimination sensitivity. This suggests that temporal predictions are encoded in delta oscillatory phase during the foreperiod interval. In sum, we show that auditory perception benefits from implicit temporal contingencies, and provide evidence for a role of slow neural oscillations in the endogenous representation of temporal predictions, in absence of exogenously driven entrainment to rhythmic input.

Late cortical tracking of ignored speech facilitates neural selectivity in acoustically challenging conditions.

Listening requires selective neural processing of the incoming sound mixture, which in humans is borne out by a surprisingly clean representation of attended-only speech in auditory cortex. How this neural selectivity is achieved even at negative signal-to-noise ratios (SNR) remains unclear. We show that, under such conditions, a late cortical representation (i.e., neural tracking) of the ignored acoustic signal is key to successful separation of attended and distracting talkers (i.e., neural selectivity). We recorded and modeled the electroencephalographic response of 18 participants who attended to one of two simultaneously presented stories, while the SNR between the two talkers varied dynamically between +6 and -6 dB. The neural tracking showed an increasing early-to-late attention-biased selectivity. Importantly, acoustically dominant (i.e., louder) ignored talkers were tracked neurally by late involvement of fronto-parietal regions, which contributed to enhanced neural selectivity. This neural selectivity, by way of representing the ignored talker, poses a mechanistic neural account of attention under real-life acoustic conditions.

Tracking Temporal Hazard in the Human Electroencephalogram Using a Forward Encoding Model.

Human observers automatically extract temporal contingencies from the environment and predict the onset of future events. Temporal predictions are modeled by the hazard function, which describes the instantaneous probability for an event to occur given it has not occurred yet. Here, we tackle the question of whether and how the human brain tracks continuous temporal hazard on a moment-to-moment basis, and how flexibly it adjusts to strictly implicit variations in the hazard function. We applied an encoding-model approach to human electroencephalographic data recorded during a pitch-discrimination task, in which we implicitly manipulated temporal predictability of the target tones by varying the interval between cue and target tone (i.e. the foreperiod). Critically, temporal predictability either was driven solely by the passage of time (resulting in a monotonic hazard function) or was modulated to increase at intermediate foreperiods (resulting in a modulated hazard function with a peak at the intermediate foreperiod). Forward-encoding models trained to predict the recorded EEG signal from different temporal hazard functions were able to distinguish between experimental conditions, showing that implicit variations of temporal hazard bear tractable signatures in the human electroencephalogram. Notably, this tracking signal was reconstructed best from the supplementary motor area, underlining this area's link to cognitive processing of time. Our results underline the relevance of temporal hazard to cognitive processing and show that the predictive accuracy of the encoding-model approach can be utilized to track abstract time-resolved stimuli.

Implicit variations of temporal predictability: Shaping the neural oscillatory and behavioural response.

Being able to predict when an event will occur (temporal predictability) can help us prepare and time our responses. We here sought to delineate the neural and behavioural corollaries of highly implicit, probabilistic temporal predictability in an auditory foreperiod paradigm. To this end, we measured electroencephalography (EEG) and response times in two independent experiments (total N =46). Unbeknownst to participants, we induced a probabilistic variation of cue-target delays (i.e., foreperiods) in a pitch-discrimination task on a noise-embedded tone: The smaller the standard deviation of the underlying foreperiod distribution, the more predictable the time of target occurrence should be. Both experiments showed that more predictive foreperiods sped up listeners' responses. Crucially, neural signatures of temporal predictability emerged when comparing EEG activity between conditions of varying temporal predictability. First, cue-related P2 evoked responses were less pronounced for cues that implicitly signalled temporal predictability of target occurrence. Second, in both experiments, fronto-central delta (1-4Hz) phase coherence was found relatively reduced during predictive foreperiods. Concomitantly, in Experiment II, the most predictive condition yielded a central alpha (7-12Hz) power increase just before the most likely time point of tone onset, likely reflecting improved temporal orienting of attention. In sum, neural oscillations in anticipation of, and response times to a target show that humans are susceptible to even strictly implicit, probabilistic temporal regularities.

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.

How long depends on how fast--perceived flicker dilates subjective duration.

How do humans perceive the passage of time and the duration of events without a dedicated sensory system for timing? Previous studies have demonstrated that when a stimulus changes over time, its duration is subjectively dilated, indicating that duration judgments are based on the number of changes within an interval. In this study, we tested predictions derived from three different accounts describing the relation between a changing stimulus and its subjective duration as either based on (1) the objective rate of changes of the stimulus, (2) the perceived saliency of the changes, or (3) the neural energy expended in processing the stimulus. We used visual stimuli flickering at different frequencies (4-166 Hz) to study how the number of changes affects subjective duration. To this end, we assessed the subjective duration of these stimuli and measured participants' behavioral flicker fusion threshold (the highest frequency perceived as flicker), as well as their threshold for a frequency-specific neural response to the flicker using EEG. We found that only consciously perceived flicker dilated perceived duration, such that a 2 s long stimulus flickering at 4 Hz was perceived as lasting as long as a 2.7 s steady stimulus. This effect was most pronounced at the slowest flicker frequencies, at which participants reported the most consistent flicker perception. Flicker frequencies higher than the flicker fusion threshold did not affect perceived duration at all, even if they evoked a significant frequency-specific neural response. In sum, our findings indicate that time perception in the peri-second range is driven by the subjective saliency of the stimulus' temporal features rather than the objective rate of stimulus changes or the neural response to the changes.

Attentional selection dilates perceived duration.

How do observers judge the passage of time at a short time-scale? Humans are not equipped with a dedicated sensory system for perceiving durations in the same way as they are equipped with systems for perceiving light and sound. Thus, subjective duration depends on the sensory and cognitive processes triggered by sensory input, eg visual or auditory stimuli. Previous studies have demonstrated that the dynamics of this sensory input (eg the rate of stimulus presentation) affect duration judgments. However, it is yet unclear whether automatic or attentive processing of such dynamics accounts for their effect on subjective duration. Automatic and attentive stimulus processing can be distinguished when stimuli are presented in a rapid serial visual presentation (RSVP) paradigm. The second of two targets embedded in an RSVP stream often fails to attract participants' attention and escapes conscious detection, in spite of being automatically processed at a perceptual level. In the present study, we presented RSVP streams and combined a target detection task with a prospective duration judgment task. We demonstrate in three experiments that the number of subjectively perceived target stimuli (and not the number of objectively presented targets) determines subjective duration of the entire RSVP sequence. Target stimuli which escape attentional selection did not affect perceived duration. This finding indicates that attentive rather than automatic processing of stimulus dynamics leads to the subjective time dilation of dynamic stimuli.