Biology subjects

Biabani, M.

Publications and source records attributed to Biabani, M..

18 recordsLinked to original sources

bioRxiv2026

Repetitive suprathreshold paired-pulse transcranial magnetic stimulation alters corticospinal excitability

Background: Suprathreshold paired-pulse transcranial magnetic stimulation (TMS) at interstimulus intervals (ISIs) of 20-30 ms evokes cumulative online increases in corticospinal excitability. Whether these changes persist beyond stimulation remains unclear. This study investigated whether repetitive paired-pulse TMS targeting periods of net excitation or inhibition induces offline changes in corticospinal excitability. Methods: Nineteen healthy participants completed three sessions, separated by at least one week, involving 40 pairs of repetitive paired-pulse TMS delivered to the left primary motor cortex at an ISI of 20 ms (rPPS20) or 100 ms (rPPS100), or a single-pulse control condition (rSPS). Offline changes in corticospinal excitability were assessed using motor evoked potential (MEP) amplitudes recorded before and after stimulation. Results: During stimulation, MEP amplitudes decreased across trials in the rPPS100 condition, although online changes did not differ significantly between conditions. Following stimulation, MEP amplitudes were significantly reduced for up to 40 minutes after rPPS100 (p = 0.017) but were not changed following rPPS20 or rSPS (all p > 0.05). Conclusion: Contrary to our hypothesis, repetitive paired-pulse TMS targeting periods of net excitation did not induce offline changes in corticospinal excitability. In contrast, stimulation targeting long-interval intracortical inhibition (LICI) produced sustained reductions in corticospinal excitability, suggesting that repetitive LICI-inducing stimulation can induce lasting plasticity within the corticospinal system.

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bioRxiv2026

Context Shapes Alpha-Band Electrophysiological Signatures Under Psilocybin

Background: Psilocybin is a serotonergic psychedelic that provides a pharmacological tool for examining neural activity associated with altered states of consciousness. Existing studies report reduced power (4-30 Hz) in posterior regions and altered functional connectivity (FC) at rest, but have typically involved small samples and coarse anatomical parcellations. Additionally, perceptual engagement has been shown to attenuate psilocybin-related whole-brain FC changes. However, how naturalistic context modifies psilocybin's frequency-specific electrophysiological effects remains incompletely characterised. Methods: We analysed the largest psilocybin electroencephalography (EEG) dataset to date (n = 56), with matched baseline recordings acquired before psilocybin administration. EEG was recorded during movie viewing, eyes-closed rest, guided meditation, and music listening. Cortical source activity was estimated using minimum-norm estimation and analysed across six canonical frequency bands (1-80 Hz). Spectral power, alpha peak frequency (APF), and magnitude-squared coherence within and between seven functional networks were quantified. Psilocybin effects were assessed using permutation testing with FDR correction, and linear mixed-effects models separated the contributions of drug, context, and their interaction to alpha-band connectivity. Results: At rest, psilocybin decreased alpha (8-13 Hz) power, increased gamma-range power, and elevated APF. Alpha-band FC decreased within and between higher-order networks implicated in top-down control. Relative to eyes-closed rest, psilocybin-context interactions showed that structured sensory environments attenuated or reversed psilocybin-related alpha-band connectivity changes. Conclusions: These findings identify alpha-band activity as a key electrophysiological correlate of psilocybin-related changes in cortical communication. Structured sensory input reshapes psilocybin-related cortical dynamics, particularly alpha-band connectivity across large-scale cortical networks, with the strength and direction of this modulation differing by context.

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bioRxiv2026

An Integrated EEG, TMS-EEG and Behavioural Dataset for Investigating the Neural Correlates of Working Memory

Working memory is a fundamental process that underlies cognition. Accordingly, the neural mechanisms that support working memory performance are of great interest in cognitive neuroscience. We present a publicly accessible dataset comprising 123 healthy adults (mean age = 28.58 years, SD = 7.56; age range = 18-46 years; 75 females) to facilitate investigation of the neural mechanisms underlying working memory performance in humans. Across two days of assessment, participants underwent a battery of cognitive tasks assessing working memory and other cognitive domains (day 1), followed by 62-channel EEG recordings (day 2). EEG data were acquired at rest, during a visual working memory task, and following single-pulse transcranial magnetic stimulation (TMS-EEG) targeting key brain regions involved in working memory. This article provides a detailed overview of the study design, methodology, and data characteristics.

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bioRxiv2025

Cumulative timing-dependent changes in corticospinal excitability during suprathreshold paired-pulse transcranial magnetic stimulation

Transcranial magnetic stimulation (TMS) is widely used to assess inhibitory and facilitatory circuits within the primary motor cortex. However, accumulating evidence suggests that even brief TMS paradigms may induce unintended changes in corticospinal excitability. Here, we examined whether suprathreshold paired-pulse TMS delivered at inter-stimulus intervals (ISIs) associated with intracortical facilitation or long-interval cortical inhibition (LICI) elicits cumulative changes in motor-evoked potential (MEP) amplitude. In experiment 1, we reanalysed data from 17 participants who received 20 suprathreshold paired-pulses at eight ISIs (10-200 ms) and 40 unconditioned single pulses. Stimulation was pseudo-randomised and distributed evenly across four blocks. A linear mixed-effects model assessed trial-wise changes in MEP amplitude across ISIs. We replicated the design in an independent sample (n=10, experiment 2). A significant trial-by-ISI interaction was observed in both cohorts. Specifically, MEP amplitudes increased across trials for ISIs of 20 and 30 ms (p<0.05), but remained stable at LICI-related ISIs (100-150 ms). A similar increase was also seen with single-pulse TMS. These findings demonstrate that suprathreshold paired-pulse TMS at short ISIs can cumulatively enhance corticospinal excitability during stimulation. Furthermore, the results suggest potential for using these protocols not just for probing cortical circuits, but also as interventions to modulate motor system excitability.

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bioRxiv2025

Psychedelics Align Brain Activity with Context

Psychedelics can profoundly alter consciousness by reorganising brain connectivity; however, their effects are context-sensitive. To understand how this reorganisation depends on context, we collected and comprehensively analysed the largest psychedelic neuroimaging dataset to date. Sixty-two adults were scanned with functional MRI and EEG during rest and naturalistic stimuli (meditation, music, and movie), before and after ingesting 19 mg of psilocybin (functional MRI {approx}80 min post-dose; EEG {approx}150 min post-dose). Half the participants ranked the experience among the most meaningful of their lives. Under psilocybin, functional MRI and EEG signals recorded during eyes-closed conditions became similar to those recorded during an eyes-open condition. Global functional connectivity increased in associative regions and decreased in sensory areas. Using machine learning to represent neural activity as low-dimensional trajectories, we found that psilocybin reorganised these into structured, context-sensitive patterns of brain activity that reflected both experimental condition and the quality of subjective experience, revealing an organisation that was missed by time-averaged connectivity measures. Under psilocybin, brain networks that ordinarily segregate internal and external processing coherently integrated and aligned neural dynamics with context. This context-alignment manifested as distinct and cohesive neural trajectories in participants reporting positively felt self- and boundary-dissolving effects, corresponding to the felt experience of being part of the environment, which we refer to as embeddedness--the subjective experience of being continuous with, rather than separate from, the surrounding environment. The strength of this context-alignment was associated with next-day mindset change, bridging the neural, experiential, and therapeutic dimensions of the psychedelic state. These findings show that the organisation of brain activity covaries with the experiential coherence of the psychedelic state, and provide a systems-level framework for how context-sensitive brain dynamics link neurobiology to subjective experience and behavioural change.

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bioRxiv2024

Default mode and motor networks facilitate early learning of implicit motor sequences: a multimodal MR spectroscopy and fMRI study

Learning new motor skills is a fundamental process that involves the sequencing of actions. Skill develops with practice and time, and manifests as performance that is fast and accurate. Although we know that learning can occur through an implicit process in the absence of conscious awareness, and across multiple temporal scales, the precise neural mechanisms mediating implicit motor sequence learning remain poorly understood. Similarly, the capacity for interventions with known influence on learning and memory, such as cardiovascular exercise, to facilitate implicit learning is yet to be clearly established. Here, we investigated the neuroplasticity of implicit motor sequence learning and the effect of acute exercise priming. Healthy adults (39.5% female) aged 22.55 {+/-} 2.69 years were allocated to either a high-intensity exercise (HIIT) group (n = 16) or to a very low-intensity control group (n = 17). Following exercise, participants performed a serial reaction time task. MR spectroscopy estimates of sensorimotor gamma-aminobutyric acid (GABA) were acquired before and after exercise, and during task performance, and resting-state fMRI was acquired at the end of the protocol. We show that early stages of learning are linked to default mode network connectivity, while the overall degree of learning following sustained practice is associated with motor network connectivity. Sensorimotor GABA concentration was linked to the early stages of learning, and GABA concentration was modulated following HIIT, although the two were not related. Overall, via integration of multiple neuroimaging modalities we demonstrate that interactions between hippocampal and motor networks underlie implicit motor sequence learning. Key points summary- Motor learning occurs across different temporal scales and can arise implicitly in the absence of conscious awareness. - Explicit motor learning is linked to the brains primary inhibitory neurotransmitter, GABA, and interactions across motor and hippocampal networks. - Whether these same neural mechanisms are implicated in implicit learning is unclear. Similarly, the capacity to influence learning via priming with cardiovascular exercise is yet to be clearly established. - We show that early implicit learning is underpinned by default mode network connectivity and sensorimotor GABA concentration, while total learning following sustained practice is linked to motor network connectivity. We also found that HIIT exercise elevated sensorimotor GABA concentration, but not the magnitude of implicit learning. - Overall, our results highlight shared involvement of default mode and motor networks in implicit motor sequence learning.

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bioRxiv2024

Supplementary motor area disinhibition during motor sequence learning: A TMS-EEG study

BackgroundIn primary motor cortex, changes in excitatory and inhibitory neurotransmission (E:I balance) accompany motor sequence learning. In particular, there is an early reduction in inhibition (i.e., disinhibition). The supplementary motor area (SMA) is a key brain region involved in the learning of sequences, however the neurophysiological mechanisms within SMA which support motor sequence learning remain poorly understood. Disinhibition may also occur in SMA, but this possibility remains unexamined. ObjectiveWe investigated disinhibition within SMA during motor sequence learning using combined transcranial magnetic stimulation (TMS) and electroencephalography (EEG). MethodsTwenty-nine healthy adults practiced a sequential motor task. TMS-evoked potentials (TEPs) resulting from SMA stimulation were measured with EEG before, during, and after practice. The N45 TEP peak was our primary measure of disinhibition. Furthermore, the slope of aperiodic EEG activity was included as an additional E:I balance measure. ResultsSignificant improvements in task performance (i.e., learning) occurred with practice. We observed smaller N45 amplitudes during early learning relative to baseline (both p < .01), indicative of disinhibition. Intriguingly, aperiodic exponents increased as learning progressed and were associated with greater sequence learning (p < .05). ConclusionOur results show disinhibition within SMA during the planning phase of motor sequence learning and thus provide novel understanding on the neurophysiological mechanisms within higher-order motor cortex that accompany new sequence learning.

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bioRxiv2023

Neurophysiology of perceptual decision-making and its alterations in attention deficit hyperactivity disorder (ADHD)

Despite the prevalence of ADHD, efforts to develop a detailed understanding of the neuropsychology of this neurodevelopmental condition are complicated by the diversity of interindividual presentations and the inability of current clinical tests to distinguish between its sensory, attentional, arousal or motoric contributions. Identifying objective methods that can decompose the clinical heterogeneity of ADHD is a long-held goal that is hoped to advance our understanding of etiological processes and potentially aid the development of personalized treatment approaches. Here, we examine key neuropsychological components of ADHD within an electrophysiological (EEG) perceptual decision-making paradigm that is capable of isolating distinct neural signals of several key information processing stages necessary for sensory-guided actions from attentional selection to motor responses. We show that compared to typically developing children, children with ADHD displayed slower and less accurate performance, which was driven by the atypical dynamics of discrete electrophysiological signatures of attentional selection, the accumulation of sensory evidence, and strategic adjustments reflecting urgency of response. These findings offer an integrated account of decision-making in ADHD and establish discrete neural signals that can be used to understand the wide range of neuropsychological performance variations in individuals with ADHD. Significance StatementThe efficacy of diagnostic and therapeutic pathways in ADHD is limited by our incomplete understanding of its neurological basis. One promising avenue of research is the search for basic neural mechanisms that may contribute to the variety of cognitive challenges associated with ADHD. We developed a mechanistic account of differences in a fundamental cognitive process by integrating across neurocognitive, neurophysiological (i.e., EEG), and computational levels of analysis. We detected distinct neural changes in ADHD that explained altered performance (e.g., slowed and less accurate responses). These included changes in neural patterns of attentional selection, sensory information processing, and response preparation. These findings enhance our understanding of the neurophysiological profile of ADHD and may offer potential targets for more effective, personalized interventions.

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bioRxiv2023

Characterising the contribution of auditory and somatosensory inputs to TMS-evoked potentials following stimulation of prefrontal, premotor and parietal cortex

Transcranial magnetic stimulation (TMS) results in a series of evoked potentials (TEPs) in electroencephalography (EEG) recordings. However, it remains unclear whether these responses reflect neural activity resulting from transcranial stimulation of the cortex, or from the sensory experiences of TMS. Across three experiments (total n = 135), we recorded EEG activity following TMS to the dorsolateral prefrontal cortex, premotor cortex, and parietal cortex as well as a sensory control condition (stimulation of the shoulder or electrical stimulation of the scalp with a click sound). We found that TEPs showed a stereotypical frontocentral N100/P200 complex following TMS of all cortical sites and control conditions, regardless of TMS intensity or the type of sensory control. In contrast, earlier TEPs (<60 ms) showed site-specific characteristics which were largest at the site of stimulation. Self-reported sensory experiences differed across sites, with prefrontal stimulation resulting in stronger auditory (click sound perception) and somatosensory input (scalp muscle twitch, discomfort) than premotor or parietal stimulation, a pattern that was reflected in the amplitude of later (N100/P200), but not earlier (<60 ms) TEP peak amplitudes. Later TEPs were also larger in individuals who experienced stronger click sound perception and, to a lesser extent, TMS-evoked scalp muscle twitches. Increasing click sound perception by removing auditory masking increased N100/P200 amplitudes without altering earlier peaks, an effect which was more prominent at sites with more successful masking. Together, these findings suggest that the frontocentral N100/P200 complex represents a generalised sensory response resulting from TMS-related auditory and somatosensory input. In contrast, early TEP peaks likely reflect activity resulting from transcranial stimulation of the cortex. The results have important implications for designing and interpreting TEP studies, especially when comparing TEPs between stimulation sites and participant groups showing differences in sensory experiences following TMS.

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bioRxiv2022

Introducing RELAX (the Reduction of Electroencephalographic Artifacts): A fully automated pre-processing pipeline for cleaning EEG data - Part 1: Algorithm and Application to Oscillations

Electroencephalographic (EEG) data is typically contaminated with non-neural artifacts which can confound the results of experiments. Artifact cleaning approaches are available, but often require time-consuming manual input and significant expertise. Advancements in artifact cleaning often only address a single artifact, are only compared against a small selection of pre-existing methods, and seldom assess whether a proposed advancement improves experimental outcomes. To address these issues, we developed RELAX (the Reduction of Electroencephalographic Artifacts), an automated EEG cleaning pipeline implemented within EEGLAB that reduces all artifact types. RELAX cleans continuous data using Multiple Wiener filtering [MWF] and/or wavelet enhanced independent component analysis [wICA] applied to artifacts identified by ICLabel [wICA_ICLabel]). Several versions of RELAX were tested using three datasets containing a mix of cognitive and resting recordings (N = 213, 60 and 23 respectively). RELAX was compared against six commonly used EEG cleaning approaches across a wide range of artifact cleaning quality metrics, including signal-to-error and artifact-to-residue ratios, measures of remaining blink and muscle activity, and the amount of variance explained by experimental manipulations after cleaning. RELAX with MWF and wICA_ICLabel showed amongst the best performance for cleaning blink and muscle artifacts while still preserving neural signal. RELAX with wICA_ICLabel (and no MWF) may perform better at detecting the effect of experimental manipulations on alpha oscillations in working memory tasks. The pipeline is easy to implement in MATLAB and freely available on GitHub. Given its high cleaning performance, objectivity, and ease of use, we recommend RELAX for data cleaning across EEG studies.

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bioRxiv2022

Introducing RELAX (the Reduction of Electroencephalographic Artifacts): A fully automated pre-processing pipeline for cleaning EEG data - Part 2: Application to Event-Related Potentials

Electroencephalography (EEG) is commonly used to examine neural activity time-locked to the presentation of a stimulus, referred to as an Event-Related Potential (ERP). However, EEG is also influenced by non-neural artifacts, which can confound ERP comparisons. Artifact cleaning can reduce artifacts, but often requires time-consuming manual decisions. Most automated cleaning methods require frequencies <1Hz to be filtered out of the data, so are not recommended for ERPs (which often contain <1Hz frequencies). In our companion article, we introduced RELAX (the Reduction of Electroencephalographic Artifacts), an automated and modular cleaning pipeline that reduces artifacts with Multiple Wiener Filtering (MWF) and/or wavelet enhanced independent component analysis (wICA) applied to artifact components detected with ICLabel (wICA_ICLabel) (Bailey et al., 2022). To evaluate the suitability of RELAX for data cleaning prior to ERP analysis, multiple versions of RELAX were compared to four commonly used EEG cleaning pipelines. Cleaning performance was compared across a range of artifact cleaning metrics and in the amount of variance in ERPs explained by different conditions in a Go-Nogo task. RELAX with MWF and wICA_ICLabel cleaned the data the most effectively and produced amongst the most dependable ERP estimates. RELAX with wICA_ICLabel only or MWF_only may detect experimental effects better for some ERP measures. Importantly, RELAX can high-pass filter data at 0.25Hz, so is applicable to analyses involving ERPs. The pipeline is easy to implement via EEGLAB in MATLAB and is freely available on GitHub. Given its performance, objectivity, and ease of use, we recommend RELAX for EEG data cleaning. The MATLAB code, the supplementary materials, and a simple instruction manual explaining how to implement the RELAX pipeline can be downloaded from https://github.com/NeilwBailey/RELAX/releases. A condition of use of the pipeline is that the version of the pipeline used is referred to as RELAX_[pipeline], for example "RELAX_MWF_wICA" or "RELAX_wICA_ICLabel", and that the current paper be cited, as well as the dependencies used. These dependencies are likely to include: EEGLAB (Delorme & Makeig, 2004), fieldtrip (Oostenveld et al., 2011), the MWF toolbox (Somers et al., 2019), fastICA (Hyvarinen, 1999), wICA (Castellanos & Makarov, 2006), ICLabel (Pion-Tonachini et al., 2019), and PREP (Bigdely-Shamlo et al., 2015) See our companion article for the application of RELAX to the study of oscillatory power (Bailey et al., 2022).

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bioRxiv2022

Task-related changes in aperiodic activity are related to visual working memory capacity independent of event-related potentials and alpha oscillations

Research using electroencephalography (EEG) has shown that individual differences in visual working memory capacity are related to slow-wave event-related potentials (ERPs) and suppression of alpha-band oscillatory power during the delay period of memory tasks. However, recent evidence suggests that changes in non-oscillatory (aperiodic) features of the EEG signal are related to working memory performance. We assessed several features of task-related changes in aperiodic activity including its spatial distribution, the effect of memory load, and the relationships between aperiodic activity, memory capacity, slow-wave ERPs, and alpha suppression. Eighty-four healthy individuals performed a continuous recall working memory (WM) task consisting of 2, 4 or 6 coloured squares while EEG was recorded. Aperiodic activity during a baseline and WM delay period was quantified by fitting a model to the background of the EEG power spectra, which returned parameters describing the slope (exponent) and broadband offset of the spectra. The aperiodic exponent decreased (i.e., slope flattened) in lateral parieto-occipital electrodes but increased (i.e., slope steepened) in fronto-central electrodes during the WM delay period, whereas the offset decreased over parieto-occipital electrodes. These task-related changes in aperiodic activity did not differ between memory loads. Larger increases in the aperiodic exponent were associated with higher working memory capacity measured from both the WM task and a separate battery of complex span tasks, a relationship that was independent of slow-wave ERPs and alpha suppression. Our findings suggest that WM task-related changes in aperiodic activity are region specific and reflect an independent neural mechanism that is important for general working memory ability.

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bioRxiv2021

Designing and comparing cleaning pipelines for TMS-EEG data: a theoretical overview and practical example

Combining transcranial magnetic stimulation (TMS) with electroencephalography (EEG) is growing in popularity as a method for probing the reactivity and connectivity of neural circuits in basic and clinical research. However, using EEG to measure the neural responses to TMS is challenging due to the unique artifacts introduced by combining the two techniques. In this paper, we overview the artifacts present in TMS-EEG data and the offline cleaning methods used to suppress these unwanted signals. We then describe how open science practices, including the development of open-source toolboxes designed for TMS-EEG analysis (e.g., TESA - the TMS-EEG signal analyser), have improved the availability and reproducibility of TMS-EEG cleaning methods. We provide theoretical and practical considerations for designing TMS-EEG cleaning pipelines and then give an example of how to compare different pipelines using TESA. We show that changing even a single step in a pipeline designed to suppress decay artifacts results in TMS-evoked potentials (TEPs) with small differences in amplitude and spatial topography. The variability in TEPs resulting from the choice of cleaning pipeline has important implications for comparing TMS-EEG findings between research groups which use different online and offline approaches. Finally, we discuss the challenges of validating cleaning pipelines and recommend that researchers compare outcomes from TMS-EEG experiments using multiple pipelines to ensure findings are not related to the choice of cleaning methods. We conclude that the continued improvement, availability, and validation of cleaning pipelines is essential to ensure TMS-EEG reaches its full potential as a method for studying human neurophysiology. HighlightsO_LIConcurrent TMS-EEG is challenging due to artifacts in the recorded signals. C_LIO_LIWe overview offline methods for cleaning TEPs and provide tips on pipeline design. C_LIO_LIWe use TESA to compare pipelines and show changing a single step alters TEPs. C_LIO_LIWe discuss the challenges in validating pipelines for TMS-EEG analysis. C_LIO_LIWe suggest using multiple pipelines to minimise the impact of method choice on TEPs. C_LI

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bioRxiv2021

No evidence for changes in GABA concentration, functional connectivity, or working memory following continuous theta burst stimulation over dorsolateral prefrontal cortex

Continuous theta burst stimulation (cTBS) is thought to reduce cortical excitability and modulate functional connectivity, possibly by altering cortical inhibition at the site of stimulation. However, most evidence comes from the motor cortex and it remains unclear whether similar effects occur following stimulation over other brain regions. We assessed whether cTBS over left dorsolateral prefrontal cortex altered gamma aminobutyric acid (GABA) concentration, functional connectivity and brain dynamics at rest, and brain activation and memory performance during a working memory task. Seventeen healthy individuals participated in a randomised, sham-controlled, cross-over experiment. Before and after either real or sham cTBS, magnetic resonance spectroscopy was obtained at rest to measure GABA concentrations. Functional magnetic resonance imaging (fMRI) was also recorded at rest and during an n-back working memory task to measure functional connectivity, regional brain activity (low-frequency fluctuations), and task-related patterns of brain activity. We could not find evidence for changes in GABA concentration (P=0.66, Bayes factor [BF10]=0.07), resting-state functional connectivity (P(FWE)>0.05), resting-state low-frequency fluctuations (P=0.88, BF10=0.04), blood-oxygen level dependent activity during the n-back task (P(FWE) >0.05), or working memory performance (P=0.13, BF10=0.05) following real or sham cTBS. Our findings add to a growing body of literature suggesting the effects of cTBS are highly variable between individuals and question the notion that cTBS is a universal inhibitory paradigm.

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bioRxiv2020

Assessing cerebellar-cortical connectivity using concurrent TMS-EEG: A Feasibility Study

BackgroundCombined single-pulse transcranial magnetic stimulation (TMS) and electroencephalography (EEG) has been used to probe the features of local networks in the cerebral cortex. Here we investigate whether we can use this approach to explore long-range connections between the cerebellum and cerebral cortex. ObjectiveTo assess the feasibility of using cerebellar TMS-EEG for the exploration of cerebellar-cerebral network dynamics. MethodsTen healthy adults received single-pulse suprathreshold TMS to the cerebellum and an occipital/parietal control site with double-cone and figure-of-eight coils while cerebral activity was recorded. A multisensory electrical control condition was used to simulate the sensation of the double-cone coil at the cerebellar site. Two cleaning pipelines were compared, and the spatiotemporal relationships of the EEG output between conditions were examined at sensor and source levels. ResultsCerebellar stimulation with the double-cone coil resulted in large artefacts in the EEG trace. The addition of SOUND filtering to the cleaning pipeline improved the signal such that further analyses could be undertaken. The cortical potentials evoked by the active TMS conditions showed strong relationships with the responses to the multisensory control condition after ~50 ms. A distinct parietal component at ~42 ms was found following cerebellar double-cone stimulation. ConclusionsCerebellar double-cone stimulation produces large artefacts in the EEG. Cerebellar-specific responses could not be reliably differentiated from sensory evoked potentials after ~50 ms. While evoked potentials differed across conditions at early latencies, it is unclear as to whether these represented TMS-related network activation of the cerebellarthalamocortical tract, or whether components were dominated by sensory contamination and/or coil-driven artefacts. Further work will be required to clarify the specific contribution of cerebellar-cortical connectivity to the observed early latency signals.

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bioRxiv2019

Do TMS evoked responses measured from scalp and hand represent the same cortical mechanisms?

Transcranial magnetic stimulation (TMS) is a powerful tool to investigate cortical circuits. Changes in cortical excitability following TMS are typically assessed by measuring changes in either conditioned motor-evoked potentials (MEPs) following paired-pulse TMS over motor cortex or evoked potentials measured with electroencephalography following single-pulse TMS (TEPs). However, it is unclear whether these two measures of cortical excitability index the same cortical response. Twenty-four healthy participants received local and interhemispheric paired-pulse TMS over motor cortex with eight inter-pulse intervals, suband suprathreshold conditioning intensities, and two different pulse waveforms, while MEPs were recorded from a hand muscle. TEPs were also recorded in response to single-pulse TMS using the conditioning pulse alone. The relationships between TEPs and conditioned-MEPs were evaluated using metrics sensitive to both their magnitude at each timepoint and their overall shape across time. The impacts of undesired sensory potentials resulting from TMS pulse and muscle contractions were also assessed on both measures. Both conditioned-MEPs and TEPs were sensitive to re-afferent somatosensory activity following motor-evoked responses, but over different post-stimulus timepoints. Moreover, the amplitude of low-frequency oscillations in TEPs was strongly correlated with the sensory potentials, whereas early and local high-frequency responses showed minimal relationships. Accordingly, conditioned-MEPs did not correlate with TEPs in the time domain but showed high shape similarity with the amplitude of high-frequency oscillations in TEPs. Therefore, despite the effects of sensory confounds, the TEP and MEP measures share a response component, suggesting that they index a similar cortical response and perhaps the same neuronal populations.

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bioRxiv2018

Characterizing and minimizing the contribution of sensory inputs to TMS-evoked potentials

BackgroundTranscranial magnetic stimulation (TMS) evokes voltage deflections in electroencephalographic (EEG) recordings, known as TMS-evoked potentials (TEPs), which are increasingly used to study brain dynamics. However, the extent to which TEPs reflect activity directly evoked by magnetic rather than sensory stimulation is unclear.\n\nObjectiveTo characterize and minimize the contribution of sensory inputs to TEPs.\n\nMethodsTwenty-four healthy participants received TMS over the motor cortex using two different intensities (below and above cortical motor threshold) and waveforms (monophasic, biphasic). TMS was also applied over the shoulder as a multisensory control condition. Common sensory attenuation measures, including coil padding and noise masking, were adopted. We examined spatiotemporal relationships between the EEG responses to the scalp and shoulder stimulations at sensor and source levels. Furthermore, we compared three different filters (independent component analysis, signal-space projection with source informed reconstruction (SSP-SIR) and linear regression) designed to attenuate the impact of sensory inputs on TEPs.\n\nResultsThe responses to the scalp and shoulder stimulations were correlated in both temporal and spatial domains, especially after [~]60 ms, regardless of the intensity and stimuli waveform. Among the three filters, SSP-SIR showed the best trade-off between removing sensory-related signals while preserving data not related to the control condition.\n\nConclusionsThe findings demonstrate that TEPs elicited by motor cortex TMS reflect a combination of transcranially and peripherally evoked brain responses despite adopting sensory attenuation methods during experiments, thereby highlighting the importance of adopting sensory control conditions in TMS-EEG studies. Offline filters may help to isolate the transcranial component of the TEP from its peripheral component, but only if these components express different spatiotemporal patterns. More realistic control conditions may help to improve the characterization and attenuation of sensory inputs to TEPs, especially in early responses.

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bioRxiv2018

TMS-evoked EEG potentials from prefrontal and parietal cortex: reliability, site specificity, and effects of dextromethorphan

Measuring the brains response to transcranial magnetic stimulation (TMS) with electroencephalography (EEG) offers a unique insight into the local cortical circuits and networks activated following stimulation, particularly in non-motor regions where less is known about TMS physiology. However, the mechanisms underlying TMS-evoked EEG potentials (TEPs) remain largely unknown. We assessed TEP reliability, site-specificity, and sensitivity to changes in excitatory neurotransmission mediated by n-methyl-d-aspartate (NMDA) receptors following stimulation of non-motor regions. In fourteen male volunteers, resting EEG and TEPs from prefrontal (PFC) and parietal (PAR) cortex were measured before and after administration of either dextromethorphan (an NMDA receptor antagonist) or placebo across two sessions separated by at least a week in a double-blinded pseudo-randomised crossover design. At baseline, TEPs showed lower within-than between-subject variability for both stimulation sites across sessions, demonstrating the reliability of non-motor TEPs within individuals. There were differences in amplitude between PFC and PAR TEPs across a wide time range (15-250 ms), however the signals were correlated after ~80 ms, suggesting that early peaks reflect site-specific activity, whereas late peaks reflect activity patterns less dependent on the stimulated sites. TEPs were not altered following dextromethorphan compared to placebo, however low frequency resting oscillations were reduced in power. Our findings suggest that TEPs from PFC and PAR: 1) are reliable within and variable between individuals; 2) reflect stimulation site specific activity across early time periods (<80 ms); and 3) are not sensitive to changes in NMDA receptor-mediated neurotransmission.

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