Publications

You can also find my articles on my Google Scholar profile.

Resting-state is not enough: alpha and mu rhythms change shape across development, but lack diagnostic sensitivity

In the field of brain rhythms, one of the rock-solid empirical findings is that oscillatory frequency increases across development, for alpha and mu rhythms. In this new 〰️preprint〰️, we also look at waveform shape across development & neurodevelopmental disorders. Read more

A Bender, B Voytek*, N Schaworonkow*, "Resting-state is not enough: alpha and mu rhythms change shape across development, but lack diagnostic sensitivity." bioRxiv, 2023. https://www.biorxiv.org/content/10.1101/2023.10.13.562301v1

An #EEGManyLabs study to test the role of the alpha phase on visual perception (a replication and new evidence)

Recent studies have debated whether alpha oscillations are crucial for how the brain processes sensory information through periods of varying excitability. To investigate, this #EEGManyLabs project is replicating a key study by Mathewson et al. (2009), which found that the visibility of visual stimuli and the amplitude of visual evoked potentials like N1, P1, P2, and P3, depend on their timing relative to alpha phases. If the replication is successful, a new experiment will explore if these effects rely on the predictability of the stimulus onset time.

M Ruzzoli, M Torralba, N Molinaro, C Benwell, D Berkowitz, D Brignani, L Falciati, L Greenwood, A Harris, C Huber-Huber, B Jack, C Keitel, M Kopčanová, C Madan, K Mathewson, S Mishra, S Mishra, P Morucci, N Myers, N Myers, F Nannetti, S Nara, J Pérez-Navarro, T Ro, N Schaworonkow, J Snyder, S Soto-Faraco, N Srinivasan, D Trübutschek, U Ajmeria, A Zazio, F Mushtaq, Y Pavlov, D Veniero "An #EEGManyLabs study to test the role of the alpha phase on visual perception (a replication and new evidence)." OSF, 2023. https://osf.io/3dhpx

Overcoming harmonic hurdles: Genuine beta-band rhythms vs. contributions of alpha-band waveform shape

Beta-band activity in the human cortex as recorded with noninvasive electrophysiology is of diverse origin. In addition to genuine beta-rhythms, there are numerous nonsinusoidal alpha-band rhythms present in the human brain, which will result in harmonic beta-band peaks. This type of activity has different temporal and response dynamics than genuine beta-rhythms. Here, it is argued that in the analysis of higher-frequency rhythms, the relationship to lower-frequency rhythms needs to be clarified. Only in that way we can arrive at strong, methodologically valid interpretations of potential functional roles and generative mechanisms of neural oscillations.

N Schaworonkow, "Overcoming harmonic hurdles: Genuine beta-band rhythms vs. contributions of alpha-band waveform shape." Imaging Neuroscience, 2023. https://direct.mit.edu/imag/article/doi/10.1162/imag_a_00018/

Oscillatory waveform shape and temporal spike correlations differ across bat frontal and auditory cortex

A fun collaboration with my colleague Francisco: This study investigates how the waveform shape of neural oscillations varies between different but functional related cortical areas in the brain, using simultaneous local field potential (LFP) recordings in the auditory and frontal cortices of awake bats. It finds significant differences in waveform shape, even for rhythmic activities of similar frequency, between these regions. Additionally, the study observes consistent variability in waveform shape across individual cycles and a higher correlation between spikes and LFPs in regions with more asymmetric waveforms, particularly in the frontal cortex. These findings suggest that oscillatory activity dynamics are distinct in different cortical areas, reflecting the anatomical and functional diversity of neural circuits.

F García-Rosales, N Schaworonkow*, J Hechavarria*, "Oscillatory waveform shape and temporal spike correlations differ across bat frontal and auditory cortex." Journal of Neuroscience, 2024. https://www.jneurosci.org/content/44/10/e1236232023

Corticospinal excitability is highest at the early rising phase of sensorimotor µ-rhythm

This study explores the relationship between the sensorimotor µ-rhythm and cortical excitability, using transcranial magnetic stimulation (TMS) and EEG in 52 adults. It examines whether the negative peak of these oscillations corresponds to the highest corticospinal excitability and if this is consistent across individuals. The research found that the highest excitability occurs during the early rising phase of the µ-rhythm, slightly delayed from the negative peak. This phase of highest excitability was consistent among participants and unaffected by different EEG-cleaning methods. The study results, showing consistent phases of highest excitability across individuals using a standard EEG setup, hold promise for clinical applications in personalized brain interventions, particularly in the motor system. Further research is needed to see if similar findings apply to other brain areas and oscillations.

C Zrenner, G Kozák, N Schaworonkow, J Metsomaa, D Baur, D Vetter, D Blumberger, U Ziemann, P Belardinelli, "Corticospinal excitability is highest at the early rising phase of sensorimotor µ-rhythm." NeuroImage, 2023. https://www.sciencedirect.com/science/article/pii/S1053811922009260

Oscillations and aperiodic activity: Evidence for dynamic changes in both during memory encoding

EEG studies show that human brain activity involves rhythmic oscillations linked to various cognitive processes and diseases. Recent research, however, highlights the importance of non-oscillatory, aperiodic neural activity, which is connected to the balance of neuronal excitatory and inhibitory signaling. Traditional analysis methods often mix up oscillations and aperiodic activity, obscuring their distinct roles in perception, cognition, and disease. This study reanalyzes intracranial human EEG data from Fellner et al., 2019, using techniques to separately analyze oscillations and aperiodic activity. The findings reveal that human memory encoding involves rapid co-modulation of both oscillatory and aperiodic activity, suggesting that these two processes, while independent, are interconnected and play significant roles in human cognition.

M Preston, N Schaworonkow, B Voytek, "Oscillations and aperiodic activity: Evidence for dynamic changes in both during memory encoding." bioRxiv, 2022. http://biorxiv.org/lookup/doi/10.1101/2022.10.04.509632

Is sensor space analysis good enough? Spatial patterns as a tool for assessing spatial mixing of EEG/MEG rhythms

Using simulations and some EEG & MEG data, we visualized how sensor space activity results from the contribution of many different types of alpha-rhythms. Visual alpha rhythms are strong and also contribute a lot of activity of frontal sensors, the extent of it may be surprising.

N Schaworonkow, V Nikulin, "Is sensor space analysis good enough? Spatial patterns as a tool for assessing spatial mixing of EEG/MEG rhythms." NeuroImage, 2022. https://www.sciencedirect.com/science/article/pii/S105381192200221X

Enhancing oscillations in intracranial electrophysiological recordings with data-driven spatial filters

I got to look at a rare type of recordings from the brain, intracranial EEG. Because there are many different types of oscillations & other types of activity present at a given moment in time, they may overlap in space and time. Here we explore if there is a better way of extracting oscillations in this type of data, trying to reconstruct activity generators from the signal recorded on electrodes.

N Schaworonkow, B Voytek, "Enhancing oscillations in intracranial electrophysiological recordings with data-driven spatial filters." PLOS Computational Biology, 2021. https://dx.plos.org/10.1371/journal.pcbi.1009298

Methodological considerations for studying neural oscillations

Neural oscillations, crucial for understanding cognitive tasks, require careful analysis due to the complexities of field potential data. This article outlines seven key considerations for their study:
(1) confirming oscillation presence;
(2) accurately defining oscillation bands;
(3) accounting for overlapping aperiodic activity;
(4) addressing temporal variability;
(5) considering non-sinusoidal waveform shape;
(6) distinguishing between spatially overlapping rhythms; and
(7) ensuring a sufficient signal-to-noise ratio.
These points are essential for avoiding misinterpretation and improving the reliability of neural oscillation research.

T Donoghue, N Schaworonkow, B Voytek, "Methodological considerations for studying neural oscillations." European Journal of Neuroscience, 2021. https://onlinelibrary.wiley.com/doi/10.1111/ejn.15361

Longitudinal changes in aperiodic and periodic activity in electrophysiological recordings in the first seven months of life

Adults have pronounced alpha-oscillations in the EEG, but young infants lack them. Oscillations emerge only gradually during the first year of life. so, in the EEG power spectrum of a 2-month year old baby, there are no pronounced peaks, they show a 1/f-power spectrum, where the spectral power decreases with increasing frequency. In this article, we looked at changes to this type of aperiodic activity across age and found a very robust decrease in the aperiodic exponent. Quanta Magazine features our work in a popular science article describing this type of brain activity: “Brain’s ‘Background Noise’ May Hold Clues to Persistent Mysteries”.

N Schaworonkow, B Voytek, "Longitudinal changes in aperiodic and periodic activity in electrophysiological recordings in the first seven months of life." Developmental Cognitive Neuroscience, 2021. http://www.sciencedirect.com/science/article/pii/S1878929320301420

Spatial neuronal synchronization and the waveform of oscillations: Implications for EEG and MEG

EEG/MEG oscillations have been treated as sinusoids up until recently. But they feature some beautiful waveforms, strongly deviating from a sine wave. In this article with my colleague Vadim Nikulin from the Max-Planck-Institute in Leipzig, we proposed measures of waveform shape in the time domain and explored these measures in a large EEG dataset.

N Schaworonkow, V Nikulin, "Spatial neuronal synchronization and the waveform of oscillations: Implications for EEG and MEG." PLOS Computational Biology, 2019. https://journals.plos.org/ploscompbiol/article?id=10.1371/journal.pcbi.1007055

EEG-triggered TMS reveals stronger brain state-dependent modulation of motor evoked potentials at weaker stimulation intensities

Ongoing brain activity influences behavior, but most non-invasive research in that direction relies on correlational analysis. With transcranial magnetic stimulation, a causal investigation is possible. A large portion of my PhD was concerned with brain-state dependent stimulation: we apply stimulation depending on the phase of the sensorimotor rhythm and observe large modulation of evoked muscle responses depending on the phase. Interestingly, in experiments this is only the case for some subjects, so the factors necessary for the modulation still need to be uncovered. Read more

N Schaworonkow, J Triesch, U Ziemann, C Zrenner, "EEG-triggered TMS reveals stronger brain state-dependent modulation of motor evoked potentials at weaker stimulation intensities." Brain Stimulation, 2019. https://linkinghub.elsevier.com/retrieve/pii/S1935861X18303115

μ-Rhythm Extracted With Personalized EEG Filters Correlates With Corticospinal Excitability in Real-Time Phase-Triggered EEG-TMS

Combining EEG and TMS, this study explores how ongoing brain activity affects cortical excitability. It compares two methods for extracting brain signals: spatial-spectral decomposition (SSD) and traditional local C3-centered Laplacian filters, focusing on the sensorimotor μ-rhythm. Both extraction methods led to a comparable degree of MEP amplitude modulation by phase of the sensorimotor μ-rhythm at the time of stimulation. This is important for accurately detecting EEG features in real-time, especially in brain regions without established benchmark methods to extract oscillatory phase.

N Schaworonkow, P Caldana, P Belardinelli, U Ziemann, T Bergmann, C Zrenner, "μ-Rhythm Extracted With Personalized EEG Filters Correlates With Corticospinal Excitability in Real-Time Phase-Triggered EEG-TMS." Frontiers in Neuroscience, 2018. https://www.frontiersin.org/article/10.3389/fnins.2018.00954/full

Simulation of electromyographic recordings following transcranial magnetic stimulation

Transcranial magnetic stimulation (TMS) is a technique that enables noninvasive manipulation of neural activity and holds promise in both clinical and basic research settings. The effect of TMS on the motor cortex is often measured by electromyography (EMG) recordings from a small hand muscle. However, the details of how TMS generates responses measured with EMG are not completely understood. We aim to develop a biophysically detailed computational model to study the potential mechanisms underlying the generation of EMG signals following TMS.

B Moezzi, N Schaworonkow, L Plogmacher, M Goldsworthy, B Hordacre, M McDonnell, N Iannella, M Ridding, J Triesch, "Simulation of electromyographic recordings following transcranial magnetic stimulation." Journal of Neurophysiology, 2018. https://www.physiology.org/doi/10.1152/jn.00626.2017

Ongoing brain rhythms shape I-wave properties in a computational model

This study explores how ongoing brain activity, like sensorimotor rhythms, affects responses to transcranial magnetic stimulation (TMS). Using a computational model, simulating TMS-induced I-waves, the impact of the power and phase of these rhythms is examined. The model shows that TMS responses vary significantly with the phase and power of ongoing activity, showing the strongest response at maximum depolarization of layer 5 neurons. The degree of phase-modulation is also intensity-dependent, with lower intensities showing stronger modulation. The model predicts that responses to TMS are highly variable for low stimulation intensities if ongoing brain rhythms are not taken into account. Closed-loop TMS-EEG holds promise for obtaining more reliable TMS effects.

N Schaworonkow, J Triesch, "Ongoing brain rhythms shape I-wave properties in a computational model." Brain Stimulation, 2018. https://linkinghub.elsevier.com/retrieve/pii/S1935861X18300937

A multi-scale computational model of the effects of TMS on motor cortex

This study presents a computational model to understand how transcranial magnetic stimulation (TMS) activates motor cortex pyramidal cells. Using MRI-based head models, it predicts electric field effects on neurons, showing layer 3 pyramidal neurons are more easily stimulated than layer 5. The model also indicates how coil orientation influences cortical activation. This work aids in refining TMS applications in clinical and research contexts.

H Seo, N Schaworonkow, S Jun, J Triesch, "A multi-scale computational model of the effects of TMS on motor cortex." F1000Research, 2017. https://f1000research.com/articles/5-1945/v3

Power-law dynamics in neuronal and behavioral data introduce spurious correlations: Power-Law Dynamics in Neuronal and Behavioral Data

Neuronal dynamics as well as behavioral output exhibit power-law dynamics. If not properly accounted for, this leads to the inflation of correlations calculated between those measures. Here, we suggest using a surrogate data procedure to counter that.

N Schaworonkow, D Blythe, J Kegeles, G Curio, V Nikulin, "Power-law dynamics in neuronal and behavioral data introduce spurious correlations: Power-Law Dynamics in Neuronal and Behavioral Data." Human Brain Mapping, 2015. http://doi.wiley.com/10.1002/hbm.22816