Christian Keitel

Stimulus-Driven Brain Oscillations in the Alpha Range: Entrainment of Intrinsic Rhythms or Frequency-Following Response?

Human brain activity is rich in rhythms of various characteristic frequencies. The last few decades have seen an increase in their use as an explanatory means, with a vast literature describing manifold correlations between dynamics of brain rhythms and behavioral performance in perceptual and

Independent Effects of Attentional Gain Control and Competitive Interactions on Visual Stimulus Processing

Attention filters behaviorally relevant stimuli from the constant stream of sensory information comprising our environment. Research into underlying neural mechanisms in humans suggests that visual attention biases mutual suppression between stimuli resulting from competition for limited processing resources. As a consequence, processing of an attended stimulus is facilitated. This account makes 2 assumptions: 1) An attended stimulus is released from mutual suppression with competing stimuli and 2) an attended stimulus experiences greater gain in the presence of competing stimuli than when it is presented alone. Here, we tested these assumptions by recording frequency-tagged potentials elicited in early visual cortex that index stimulus-specific processing. We contrasted the processing of a given stimulus when its location was attended or unattended and in the presence or the absence of a nearby competing stimulus. At variance with previous findings, competition similarly suppressed processing of attended and unattended stimuli. Moreover, the magnitude of attentional gain was comparable in the presence or the absence of competing stimuli. We conclude that visuospatial selective attention does not directly modulate mutual suppression between stimuli but instead acts as a signal gain, which biases processing toward attended stimuli independent of competition.

Sustained selective intermodal attention modulates processing of language-like stimuli

Intermodal attention (IA) is assumed to allocate limited neural processing resources to input from one specific sensory modality. We investigated effects of sustained IA on the amplitude of a 40-Hz auditory (ASSR) and a 4.3-Hz visual steady-state response (VSSR). To this end, we concurrently presented amplitude-modulated multi-speech babble and a stream of nonsense letter sets to elicit the respective brain responses. Subjects were cued trialwise to selectively attend to one of the streams for several seconds where they had to perform a lexical decision task on occasionally occurring words and pseudowords. Attention to the auditory stream led to greater ASSR amplitudes than attention to the visual stream. Vice versa, the VSSR amplitude was greater when the visual stream was attended. We demonstrate that IA research by means of frequency tagging can be extended to complex stimuli as used in the current study. Furthermore, we show not only that IA selectively modulates processing of concurrent multisensory input but that this modulation occurs during trial-by-trial cueing of IA. The use of frequency tagging may be suitable to study the role of IA in more naturalistic setups that comprise a larger number of multisensory signals.

Electrophysiological evidence for age effects on sensory memory processing of tonal patterns.

In older adults, difficulties processing complex auditory scenes, such as speech comprehension in noisy environments, might be due to a specific impairment of temporal processing at early, automatic processing stages involving auditory sensory memory (ASM). Even though age effects on auditory temporal processing have been well-documented, there is a paucity of research on how ASM processing of more complex tone-patterns is altered by age. In the current study, age effects on ASM processing of temporal and frequency aspects of two-tone patterns were investigated using a passive listening protocol. The P1 component, the mismatch negativity (MMN) and the P3a component of event-related brain potentials (ERPs) to tone frequency and temporal pattern deviants were recorded in younger and older adults as a measure of auditory event detection, ASM processing, and attention switching, respectively. MMN was elicited with smaller amplitude to both frequency and temporal deviants in older adults. Furthermore, P3a was elicited only in the younger adults. In conclusion, the smaller MMN amplitude indicates that automatic processing of both frequency and temporal aspects of two-tone patterns is impaired in older adults. The failure to initiate an attention switch, suggested by the absence of P3a, indicates that impaired ASM processing of patterns may lead to less distractibility in older adults. Our results suggest age-related changes in ASM processing of patterns that cannot be explained by an inhibitory deficit.

Concurrent visual and tactile steady-state evoked potentials index allocation of inter-modal attention: A frequency-tagging study

We investigated effects of inter-modal attention on concurrent visual and tactile stimulus processing by means of stimulus-driven oscillatory brain responses, so-called steady-state evoked potentials (SSEPs). To this end, we frequency-tagged a visual (7.5Hz) and a tactile stimulus (20Hz) and participants were cued, on a trial-by-trial basis, to attend to either vision or touch to perform a detection task in the cued modality. SSEPs driven by the stimulation comprised stimulus frequency-following (i.e. fundamental frequency) as well as frequency-doubling (i.e. second harmonic) responses. We observed that inter-modal attention to vision increased amplitude and phase synchrony of the fundamental frequency component of the visual SSEP while the second harmonic component showed an increase in phase synchrony, only. In contrast, inter-modal attention to touch increased SSEP amplitude of the second harmonic but not of the fundamental frequency, while leaving phase synchrony unaffected in both responses. Our results show that inter-modal attention generally influences concurrent stimulus processing in vision and touch, thus, extending earlier audio-visual findings to a visuo-tactile stimulus situation. The pattern of results, however, suggests differences in the neural implementation of inter-modal attentional influences on visual vs. tactile stimulus processing.

Visual cortex responses reflect temporal structure of continuous quasi-rhythmic sensory stimulation

Abstract Neural processing of dynamic continuous visual input, and cognitive influences thereon, are frequently studied in paradigms employing strictly rhythmic stimulation. However, the temporal structure of natural stimuli is hardly ever fully rhythmic but possesses certain spectral bandwidths (e.g. lip movements in speech, gestures). Examining periodic brain responses elicited by strictly rhythmic stimulation might thus represent ideal, yet isolated cases. Here, we tested how the visual system reflects quasi‐rhythmic stimulation with frequencies continuously varying within ranges of classical theta (4–7 Hz), alpha (8–13 Hz) and beta bands (14–20 Hz) using EEG. Our findings substantiate a systematic and sustained neural phase‐locking to stimulation in all three frequency ranges. Further, we found that allocation of spatial attention enhances EEG‐stimulus locking to theta‐ and alpha‐band stimulation. Our results bridge recent findings regarding phase locking (“entrainment”) to quasi‐rhythmic visual input and “frequency‐tagging” experiments employing strictly rhythmic stimulation. We propose that sustained EEG‐stimulus locking can be considered as a continuous neural signature of processing dynamic sensory input in early visual cortices. Accordingly, EEG‐stimulus locking serves to trace the temporal evolution of rhythmic as well as quasi‐rhythmic visual input and is subject to attentional bias. HighlightsDynamic visual stimuli constitute large parts of our perceptual experience.Strictly rhythmic dynamics condense in EEG‐recorded mass‐neural activity.We tested how stimuli with fluctuating rhythms reflect in the EEG.We found that the EEG allows tracing two quasi‐rhythmic stimuli in parallel.Dynamics of attended stimuli may be tracked with greater temporal precision.

Flicker-Driven Responses in Visual Cortex Change during Matched-Frequency Transcranial Alternating Current Stimulation

We tested a novel combination of two neuro-stimulation techniques, transcranial alternating current stimulation (tACS) and frequency tagging, that promises powerful paradigms to study the causal role of rhythmic brain activity in perception and cognition. Participants viewed a stimulus flickering at 7 or 11 Hz that elicited periodic brain activity, termed steady-state responses (SSRs), at the same temporal frequency and its higher order harmonics. Further, they received simultaneous tACS at 7 or 11 Hz that either matched or differed from the flicker frequency. Sham tACS served as a control condition. Recent advances in reconstructing cortical sources of oscillatory activity allowed us to measure SSRs during concurrent tACS, which is known to impose strong artifacts in magnetoencephalographic (MEG) recordings. For the first time, we were thus able to demonstrate immediate effects of tACS on SSR-indexed early visual processing. Our data suggest that tACS effects are largely frequency-specific and reveal a characteristic pattern of differential influences on the harmonic constituents of SSRs.

Trial‐by‐trial co‐variation of pre‐stimulus EEG alpha power and visuospatial bias reflects a mixture of stochastic and deterministic effects

Human perception of perithreshold stimuli critically depends on oscillatory EEG activity prior to stimulus onset. However, it remains unclear exactly which aspects of perception are shaped by this pre‐stimulus activity and what role stochastic (trial‐by‐trial) variability plays in driving these relationships. We employed a novel jackknife approach to link single‐trial variability in oscillatory activity to psychometric measures from a task that requires judgement of the relative length of two line segments (the landmark task). The results provide evidence that pre‐stimulus alpha fluctuations influence perceptual bias. Importantly, a mediation analysis showed that this relationship is partially driven by long‐term (deterministic) alpha changes over time, highlighting the need to account for sources of trial‐by‐trial variability when interpreting EEG predictors of perception. These results provide fundamental insight into the nature of the effects of ongoing oscillatory activity on perception. The jackknife approach we implemented may serve to identify and investigate neural signatures of perceptual relevance in more detail.

Sustained splits of attention within versus across visual hemifields produce distinct spatial gain profiles

Visual attention can be focused concurrently on two stimuli at noncontiguous locations while intermediate stimuli remain ignored. Nevertheless, behavioral performance in multifocal attention tasks falters when attended stimuli fall within one visual hemifield as opposed to when they are distributed across left and right hemifields. This “different-hemifield advantage” has been ascribed to largely independent processing capacities of each cerebral hemisphere in early visual cortices. Here, we investigated how this advantage influences the sustained division of spatial attention. We presented six isoeccentric light-emitting diodes (LEDs) in the lower visual field, each flickering at a different frequency. Participants attended to two LEDs that were spatially separated by an intermediate LED and responded to synchronous events at to-be-attended LEDs. Task-relevant pairs of LEDs were either located in the same hemifield (“within-hemifield” conditions) or separated by the vertical meridian (“across-hemifield” conditions). Flicker-driven brain oscillations, steady-state visual evoked potentials (SSVEPs), indexed the allocation of attention to individual LEDs. Both behavioral performance and SSVEPs indicated enhanced processing of attended LED pairs during “across-hemifield” relative to “within-hemifield” conditions. Moreover, SSVEPs demonstrated effective filtering of intermediate stimuli in “across-hemifield” condition only. Thus, despite identical physical distances between LEDs of attended pairs, the spatial profiles of gain effects differed profoundly between “across-hemifield” and “within-hemifield” conditions. These findings corroborate that early cortical visual processing stages rely on hemisphere-specific processing capacities and highlight their limiting role in the concurrent allocation of visual attention to multiple locations.

No changes in parieto-occipital alpha during neural phase locking to visual quasi-periodic theta-, alpha-, and beta-band stimulation

Recent studies have probed the role of the parieto‐occipital alpha rhythm (8–12 Hz) in human visual perception through attempts to drive its neural generators. To that end, paradigms have used high‐intensity strictly‐periodic visual stimulation that created strong predictions about future stimulus occurrences and repeatedly demonstrated perceptual consequences in line with an entrainment of parieto‐occipital alpha. Our study, in turn, examined the case of alpha entrainment by non‐predictive low‐intensity quasi‐periodic visual stimulation within theta‐ (4–7 Hz), alpha‐ (8–13 Hz), and beta (14–20 Hz) frequency bands, i.e., a class of stimuli that resemble the temporal characteristics of naturally occurring visual input more closely. We have previously reported substantial neural phase‐locking in EEG recording during all three stimulation conditions. Here, we studied to what extent this phase‐locking reflected an entrainment of intrinsic alpha rhythms in the same dataset. Specifically, we tested whether quasi‐periodic visual stimulation affected several properties of parieto‐occipital alpha generators. Speaking against an entrainment of intrinsic alpha rhythms by non‐predictive low‐intensity quasi‐periodic visual stimulation, we found none of these properties to show differences between stimulation frequency bands. In particular, alpha band generators did not show increased sensitivity to alpha band stimulation and Bayesian inference corroborated evidence against an influence of stimulation frequency. Our results set boundary conditions for when and how to expect effects of entrainment of alpha generators and suggest that the parieto‐occipital alpha rhythm may be more inert to external influences than previously thought.