Pre-stimulus influences on auditory perception arising from sensory representations and decision processes

The qualities of perception depend not only on the sensory inputs but also on the brain state before stimulus presentation. Although the collective evidence from neuroimaging studies for a relation between prestimulus state and perception is strong, the interpretation in the context of sensory computations or decision processes has remained difficult. In the auditory system, for example, previous studies have reported a wide range of effects in terms of the perceptually relevant frequency bands and state parameters (phase/power). To dissociate influences of state on earlier sensory representations and higher-level decision processes, we collected behavioral and EEG data in human participants performing two auditory discrimination tasks relying on distinct acoustic features. Using single-trial decoding, we quantified the relation between prestimulus activity, relevant sensory evidence, and choice in different task-relevant EEG components. Within auditory networks, we found that phase had no direct influence on choice, whereas power in task-specific frequency bands affected the encoding of sensory evidence. Within later-activated frontoparietal regions, theta and alpha phase had a direct influence on choice, without involving sensory evidence. These results delineate two consistent mechanisms by which prestimulus activity shapes perception. However, the timescales of the relevant neural activity depend on the specific brain regions engaged by the respective task

Analytical methods and experimental approaches for electrophysiological studies of brain oscillations

Brain oscillations are increasingly the subject of electrophysiological studies probing their role in the functioning and dysfunction of the human brain. In recent years this research area has seen rapid and significant changes in the experimental approaches and analysis methods. This article reviews these developments and provides a structured overview of experimental approaches, spectral analysis techniques and methods to establish relationships between brain oscillations and behaviour

Alpha-band rhythms in visual task performance: phase-locking by rhythmic sensory stimulation

Oscillations are an important aspect of neuronal activity. Interestingly, oscillatory patterns are also observed in behaviour, such as in visual performance measures after the presentation of a brief sensory event in the visual or another modality. These oscillations in visual performance cycle at the typical frequencies of brain rhythms, suggesting that perception may be closely linked to brain oscillations. We here investigated this link for a prominent rhythm of the visual system (the alpha-rhythm, 8-12 Hz) by applying rhythmic visual stimulation at alpha-frequency (10.6 Hz), known to lead to a resonance response in visual areas, and testing its effects on subsequent visual target discrimination. Our data show that rhythmic visual stimulation at 10.6 Hz: 1) has specific behavioral consequences, relative to stimulation at control frequencies (3.9 Hz, 7.1 Hz, 14.2 Hz), and 2) leads to alpha-band oscillations in visual performance measures, that 3) correlate in precise frequency across individuals with resting alpha-rhythms recorded over parieto-occipital areas. The most parsimonious explanation for these three findings is entrainment (phase-locking) of ongoing perceptually relevant alpha-band brain oscillations by rhythmic sensory events. These findings are in line with occipital alpha-oscillations underlying periodicity in visual performance, and suggest that rhythmic stimulation at frequencies of intrinsic brain-rhythms can be used to reveal influences of these rhythms on task performance to study their functional roles

Alpha And Gamma Oscillation Amplitudes Synergistically Predict The Perception Of Forthcoming Nociceptive Stimuli

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Neural correlates of conscious visual processing

The objective of the current thesis is to evaluate the role of alpha band activity and neural trial-to-trial variability in conscious visual perception and their relationship to each other. We investigate these measures in electrophysiological recordings of monkeys as well as the electroencephalogram (EEG) of humans using a generalized flash suppression (GFS) paradigm.2021-06-0

Functional Dissociation of Ongoing Oscillatory Brain States

The state of a neural assembly preceding an incoming stimulus is assumed to modulate the processing of subsequently presented stimuli. The nature of this state can differ with respect to the frequency of ongoing oscillatory activity. Oscillatory brain activity of specific frequency range such as alpha (8–12 Hz) and gamma (above 30 Hz) band oscillations are hypothesized to play a functional role in cognitive processing. Therefore, a selective modulation of this prestimulus activity could clarify the functional role of these prestimulus fluctuations. For this purpose, we adopted a novel non-invasive brain-computer-interface (BCI) strategy to selectively increase alpha or gamma band activity in the occipital cortex combined with an adaptive presentation of visual stimuli within specific brain states. During training, oscillatory brain activity was estimated online and fed back to the participants to enable a deliberate modulation of alpha or gamma band oscillations. Results revealed that volunteers selectively increased alpha and gamma frequency oscillations with a high level of specificity regarding frequency range and localization. At testing, alpha or gamma band activity was classified online and at defined levels of activity, visual objects embedded in noise were presented instantly and had to be detected by the volunteer. In experiment I, the effect of two levels of prestimulus gamma band activity on visual processing was examined. During phases of increased gamma band activity significantly more visual objects were detected. In experiment II, the effect was compared against increased levels of alpha band activity. An improvement of visual processing was only observed for enhanced gamma band activity. Both experiments demonstrate the specific functional role of prestimulus gamma band oscillations for perceptual processing. We propose that the BCI method permits the selective modulation of oscillatory activity and the direct assessment of behavioral consequences to test for functional dissociations of different oscillatory brain states

Individual differences in alpha frequency drive crossmodal illusory perception

Perception routinely integrates inputs from different senses. Stimulus temporal proximity critically determines whether or not these inputs are bound together. Despite the temporal window of integration being a widely accepted notion, its neurophysiological substrate remains unclear. Many types of common audio-visual interactions occur within a time window of -100ms [1-5]. For example, in the sound- induced double-flash illusion, when two beeps are presented within -100ms together with one flash, a second illusory flash is often perceived [2]. Due to their intrinsic rhythmic nature, brain oscillations are one candidate mechanism for gating the temporal window of integration. Interestingly, occipital alpha-band oscillations cycle on average every -100ms with peak frequencies ranging between 8-14Hz (i.e. 120-60ms cycle). Moreover, presenting a brief tone can phase-reset such oscillations in visual cortex [6, 7]. Based on these observations, we hypothesized that the duration of each alpha cycle might provide the temporal unit to bind audio-visual events. Here we first recorded EEG while participants performed the sound-induced double-flash illusion task [4] and found positive correlation between individual alpha-frequency (IAF) peak and the size of the temporal window of the illusion. Participants then performed the same task while receiving occipital transcranial alternating current stimulation (tACS), to modulate oscillatory activity [8] either at their IAF or at off-peak alpha-frequencies (IAF±2Hz). Compared to IAF tACS, IAF-2Hz and IAF+2Hz tACS respectively enlarged and shrunk the temporal window of illusion, suggesting that alpha oscillations might represent the temporal unit of visual processing that cyclically gates perception and the neurophysiological substrate promoting audio-visual interactions

Relating alpha power modulations to competing visuospatial attention theories

Visuospatial attention theories often propose hemispheric asymmetries underlying the control of attention. In general support of these theories, previous EEG/MEG studies have shown that spatial attention is associated with hemispheric modulation of posterior alpha power (gating by inhibition). However, since measures of alpha power are typically expressed as lateralization scores, or collapsed across left and right attention shifts, the individual hemispheric contribution to the attentional control mechanism remains unclear. This is, however, the most crucial and decisive aspect in which the currently competing attention theories continue to disagree. To resolve this long-standing conflict, we derived predictions regarding alpha power modulations from Heilman's hemispatial theory and Kinsbourne's interhemispheric competition theory and tested them empirically in an EEG experiment. We used an attention paradigm capable of isolating alpha power modulation in two attentional states, namely attentional bias in a neutral cue condition and spatial orienting following directional cues. Differential alpha modulations were found for both hemispheres across conditions. When anticipating peripheral visual targets without preceding directional cues (neutral condition), posterior alpha power in the left hemisphere was generally lower and more strongly modulated than in the right hemisphere, in line with the interhemispheric competition theory. Intriguingly, however, while alpha power in the right hemisphere was modulated by both, cue-directed leftward and rightward attention shifts, the left hemisphere only showed modulations by rightward shifts of spatial attention, in line with the hemispatial theory. This suggests that the two theories may not be mutually exclusive, but rather apply to different attentional states

Human Cortical Traveling Waves: Dynamical Properties and Correlations with Responses

The spatiotemporal behavior of human EEG oscillations is investigated. Traveling waves in the alpha and theta ranges are found to be common in both prestimulus and poststimulus EEG activity. The dynamical properties of these waves, including their speeds, directions, and durations, are systematically characterized for the first time, and the results show that there are significant changes of prestimulus spontaneous waves in the presence of an external stimulus. Furthermore, the functional relevance of these waves is examined by studying how they are correlated with reaction times on a single trial basis; prestimulus alpha waves traveling in the frontal-to-occipital direction are found to be most correlated to reaction speeds. These findings suggest that propagating waves of brain oscillations might be involved in mediating long-range interactions between widely distributed parts of human cortex

Unbewusste Modulatoren der somatosensorischen Wahrnehmung

It is intriguing that perception of the same stimulus can vary profoundly from trial to trial. For example, it has been shown in many studies that weak, so-called “near-threshold stimuli” are sometimes consciously perceived and sometimes not. In my thesis, I have been investigating factors which underlie this profound perceptual variability in the somatosensory domain. Together with my colleagues, I performed three studies in which we tested three different types of presumed non-conscious modulators of somatosensory perception. In the first – behavioral - study, we investigated how the presence of subliminal noise during a peripheral somatosensory stimulation influences perception. Counter-intuitively, we found that peripheral noise can even improve perception of weak somatosensory stimuli. In our interpretation, this occurs most likely due to “stochastic resonance” effects (Study I: Iliopoulos et al. 2014). In the second – behavioral and EEG - study, we tested the effect of different forms of pulsed subliminal stimulation (single pulses versus pulse trains) on brain rhythms and somatosensory perception. Following-up on previous results of our group, we tested the hypothesis that subliminal pulsed stimulation impairs perception of subsequent stimuli via centrally enhanced Mu rhythm. Interestingly, the main result of this study was that trains of subliminal stimuli indeed inhibited subsequent somatosensory detection, however, - in contrast to our previous findings for single pulses – trains were associated with decreased Mu rhythm. We conclude that central rhythms most likely play a role in mediating the perceptual modulation of peripheral subliminal stimuli, however, the relationship is more complex than previously assumed (Study II: Iliopoulos et al. 2020). In the third study, we examined the influence of interoceptive signaling, especially from the heart, on somatosensory perception. The hypothesis was that the cardiac phase (systole versus diastole) and the so-called heart-evoked potential (HEP) would modulate somatosensory perception. Indeed, our study showed that somatosensory perception was better during diastole than during systole and detection performance declined as the amplitude of the HEP increased. Our interpretation of the former effect assumes that all events which occur simultaneously with the “pulse” are assumed by the brain to be pulse-synchronous peripheral noise and therefore suppressed. Our interpretation of the latter effect (HEP) assumes that HEP is a marker of the relative balance between interoception and exteroception (Study III: Al et al. 2020). In conclusion, in the studies which form the basis for my thesis, we have shown that somatosensory perception is modulated by peripheral effects (modes of peripheral stimulation, peripheral noise), central effects (Mu rhythm) and interoceptive signals from the heart. The precise interplay between these modulators is an exciting research topic for future studies.Interessanterweise kann die Wahrnehmung desselben Reizes von Augenblick zu Augenblick so stark variieren, dass dieser manchmal bewusst wahrgenommen wird und manchmal nicht. In meiner Dissertation habe ich Faktoren untersucht, die dieser Wahrnehmungsvariabilität im somatosensorischen (SS) System zugrunde liegen. Mit meinen Kollegen habe ich drei Studien durchgeführt, in denen wir verschiedene mutmaßlich unbewusste Modulatoren der SS-Wahrnehmung untersuchten. In der ersten Studie untersuchten wir, wie die Wahrnehmung peripherer SS-Reize durch unterschwelliges Rauschen beeinflusst wird. Wir konnten zeigen, dass peripheres Rauschen die Wahrnehmung schwacher Reize verbessert. Dies ist ein Hinweis auf das Vorliegen von "stochastischen Resonanzeffekten" (Studie I: Iliopoulos et al. 2014). In der zweiten Studie, die neben behavioralen Messungen auch elektroencephalographische (EEG) Messungen umfasste, testeten wir die Auswirkung verschiedener Formen gepulster unterschwelliger elektrischer Fingerstimulationen (Einzelpulse gegen Pulsserien) auf die Wahrnehmung und auf Hirn-rhythmen. Ausgehend von früheren Ergebnissen unserer Arbeitsgruppe überprüften wir, ob repetitive subliminale Stimulationen die Wahrnehmung nachfolgender Reize über einen zentral verstärkten Mu-Rhythmus beeinträchtigen. Das Ergebnis dieser Studie war, dass Serien unterschwelliger Reize tatsächlich die nachfolgende SS-Wahrnehmung hemmten, jedoch - im Gegensatz zu früheren Ergebnissen für Einzelimpulse – die Reizserien mit einem verringerten Mu-Rhythmus verbunden waren. Daraus schließen wir, dass zentrale Rhythmen höchstwahrscheinlich eine Rolle bei der Wahrnehmungsmodulation durch periphere unterschwellige Reize spielen, dass aber der Zusammenhang zwischen beiden komplexer ist als bisher vermutet (Studie II: Iliopoulos et al. 2020). In der dritten Studie untersuchten wir den Einfluss interozeptiver Signale aus dem Herzen auf die SS-Wahrnehmung. Die Hypothese war, dass die Herzphase und das so genannte Herz-evozierte Potenzial (HEP) die SS-Wahrnehmung modulieren. Wir zeigten, dass die SS-Wahrnehmung während der Diastole besser war als während der Systole und dass die Wahrnehmung in umgekehrtem Verhältnis zur Amplitude des vorausgehenden HEP stand. Für den ersten Effekt legen unsere Daten nahe, dass alle Ereignisse, die zusammen mit der Pulswelle auftreten, vom Gehirn als puls-synchrones peripheres Rauschen angenommen und daher unterdrückt werden. Der zweite Befund wird in Übereinstimmung mit der Literatur am besten dadurch erklärt, dass das HEP ein Marker für das relative Gleichgewicht zwischen Interozeption und Exterozeption darstellt (Studie III: Al et al. 2020). Zusammenfassend zeigen die Ergebnisse dieser Arbeit, wie die SS-Wahrnehmung durch periphere Effekte (Art der Stimulation, Rauschen), zentrale Effekte (Mu-Rhythmus) und interozeptive Signale des Herzens moduliert wird. Das genaue Zusammenspiel zwischen diesen Modulatoren ist ein spannendes Forschungsthema für zukünftige Studien