Til Ole Bergmann

University Medical Center Mainz · Neuroimaging Center (NIC)

In this issue of Neuron, Helfrich et al. (2017) demonstrate that phase-amplitude coupling (PAC) between slow oscillations and spindles is crucial for memory consolidation, and shifts in its phase relationship may explain age-related deficits in memory performance. These results also suggest a more general function of PAC in synaptic plasticity.

Non-invasive transcranial brain stimulation (NTBS) techniques such as transcranial magnetic stimulation (TMS) and transcranial current stimulation (TCS) are important tools in human systems and cognitive neuroscience because they are able to reveal the relevance of certain brain structures or neuronal activity patterns for a given brain function. I...

Cortical oscillations, such as 8–12 Hz alpha-band activity, are thought to subserve gating of information processing in the human brain. While most of the supporting evidence is correlational, causal evidence comes from attempts to externally drive (“entrain”) these oscillations by transcranial magnetic stimulation (TMS). Indeed, the frequency prof...

During systems-level consolidation, mnemonic representations initially reliant on the hippocampus are thought to migrate to neocortical sites for more permanent storage, with an eminent role of sleep for facilitating this information transfer. Mechanistically, consolidation processes have been hypothesized to rely on systematic interactions between...

Evoked cortical responses do not follow a rigid input-output function but are dynamically shaped by intrinsic neural properties at the time of stimulation. Recent research has emphasized the role of oscillatory activity in determining cortical excitability. Here we employed EEG-guided transcranial magnetic stimulation (TMS) during non-rapid eye mov...

Theta oscillations (4-8 Hz) in frontal cortical regions are present to different degrees across states of consciousness. In sleep, theta is prominent in periods of rapid eye-movement (REM) sleep. Theta has been linked to processes of memory consolidation; however, its mechanistic contribution specifically during REM sleep is not well understood. In...

Transcranial ultrasonic stimulation (TUS) is rapidly emerging as a promising non-invasive neuromodulation technique. TUS is already well-established in animal models, providing foundations to now optimize neuromodulatory efficacy for human applications. Across multiple studies, one promising protocol, pulsed at 1000 Hz, has consistently resulted in...

Thalamocortical sleep spindles, i.e., oscillatory bursts at ~12-15 Hz of waxing and waning amplitude, are a hallmark feature of non-rapid eye movement (NREM) sleep and believed to play a key role in sleep-dependent memory reactivation and consolidation. Generated in the thalamus and projecting to neocortex and hippocampus, they are phasically modul...

Low-intensity Transcranial Ultrasonic Stimulation (TUS) is a non-invasive brain stimulation technique enabling cortical and deep brain targeting with unprecedented spatial accuracy. Given the high rate of adoption by new users with varying levels of expertise and interdisciplinary backgrounds, practical guidelines are needed to ensure state-of-the-...

Concurrent TMS-fMRI provides a step-change in the toolkit of neuroscience research. Using non-invasive perturbation of ongoing human brain activity and simultaneous read-out of its effects across the brain, it permits causal inference into human brain-behaviour relationships with important implications for both fundamental research and clinical app...

TMS is one of the most popular non-invasive brain stimulation techniques. When combined with EEG, it allows researchers to record activity from both the stimulated area and interconnected regions. However, the efficacy of TMS-EEG has been limited by intra- and inter-subject variability, highlighting the necessity of standardized stimulation procedu...

Phase-dependent plasticity has been proposed as a neurobiological mechanism by which oscillatory phase-amplitude cross-frequency coupling mediates memory process in the brain. Mimicking this mechanism, real-time EEG oscillatory phase-triggered transcranial magnetic stimulation (TMS) has successfully induced LTP-like changes in corticospinal excitab...

Transcranial ultrasonic stimulation (TUS) is rapidly emerging as a promising non-invasive neuromodulation technique. TUS is already well-established in animal models, providing foundations to now optimize neuromodulatory efficacy for human applications. Across multiple studies, one promising protocol, pulsed at 1000 Hz, has consistently resulted in...

Transcranial ultrasonic stimulation (TUS) is rapidly emerging as a promising non-invasive neuromodulation technique. TUS is already well-established in animal models, providing foundations to now optimize neuromodulatory efficacy for human applications. Across multiple studies, one promising protocol, pulsed at 1000 Hz, has consistently resulted in...

Background Transcranial evoked potentials (TEPs) measured via electroencephalography (EEG) are widely used to study the cortical responses to transcranial magnetic stimulation (TMS). Immediate transcranial evoked potentials (i-TEPs) have been obscured by pulse and muscular artifacts. Thus, the TEP peaks that are commonly reported have latencies tha...

One of the most critical challenges in using noninvasive brain stimulation (NIBS) techniques for the treatment of psychiatric and neurologic disorders is inter- and intra-individual variability in response to NIBS. Response variations in previous findings suggest that the one-size-fits-all approach does not seem the most appropriate option for enha...

In this editorial we introduce a new non-profit open access journal, Imaging Neuroscience. In April 2023, editors of the journals NeuroImage and NeuroImage:Reports resigned, and a month later launched Imaging Neuroscience. NeuroImage had long been the leading journal in the field of neuroimaging. While the move to fully open access in 2020 represen...

Using concurrent TMS-EEG, Han et al.1 identified temporal and spectral signatures of depression in a prefrontal-orbitofrontal-hippocampal network, which renormalized after rTMS. This highlights the relevance of causal network perturbation for the assessment of disease-related network states and their therapeutic modulation.

Background Transcranial ultrasonic stimulation (TUS) is rapidly emerging as a promising non-invasive neuromodulation technique. TUS is already well-established in animal models, providing foundations to now optimize neuromodulatory efficacy for human applications. Across multiple studies, one promising protocol, pulsed at 1000 Hz, has consistently...

Transcranial magnetic stimulation (TMS) evokes neuronal activity in the targeted cortex and connected brain regions. The evoked brain response can be measured with electroencephalography (EEG). TMS combined with simultaneous EEG (TMS−EEG) is widely used for studying cortical reactivity and connectivity at high spatiotemporal resolution. Methodologi...

Transcranial ultrasonic stimulation (TUS), has been shown to evoke 'visual evoked potential (VEP)-like' potentials on EEG recordings, and also to modulate sensory evoked potentials. However, pulsed TUS is accompanied by an auditory confound, and it is possible that any observed effects were, in-part, evoked by this confound. Therefore, we used ramp...

Background: Local field potentials (LFPs) represent the summation of periodic (oscillations) and aperiodic (fractal) signals. Although previous studies showed changes in beta band oscillations and burst characteristics of the subthalamic nucleus (STN) in Parkinson's disease (PD), how aperiodic activity in the STN is related to PD pathophysiology i...

Recent advancements in real‐time brain stimulation in the sleep field have led to many exciting findings. However, they have also opened up terminological ambiguities about what constitutes “open‐loop”, “closed‐loop”, and “real‐time” designs. Here, we address core theoretical aspects of these terms in the hopes of strengthening future research on t...

Transcranial direct current stimulation (tDCS) has been used for over twenty years to modulate cortical (particularly motor corticospinal) excitability both during (online) and outlasting (offline) the stimulation, with the former effects associated to the latter. However, tDCS effects are highly variable, partially because stimulation intensity is...

Sleep spindles are a hallmark electroencephalographic feature of non‐rapid eye movement sleep, and are believed to be instrumental for sleep‐dependent memory reactivation and consolidation. However, direct proof of their causal relevance is hard to obtain, and our understanding of their immediate neurophysiological consequences is limited. To inves...

One of the most critical challenges in using non-invasive brain stimulation (NIBS) techniques for the treatment of psychiatric and neurologic disorders is inter- and intra-individual variability in response to NIBS. Response variations in previous findings suggest that the one-size-fits-all approach does not seem the most appropriate option for enh...

Sleep spindles are a hallmark electroencephalographic (EEG) feature of non-rapid eye movement (NREM) sleep and believed to be instrumental for sleep-dependent memory reactivation and consolidation. However, direct proof of their causal relevance is hard to obtain, and our understanding of their immediate neurophysiological consequences is limited....

Neuronal oscillations in different frequency bands have been linked to a wide variety of cognitive functions, and may even be a fundamental mechanism of inter-regional communication. For this reason, manipulation of oscillatory activity via brain stimulation is a central goal in neuroscience research. However, the vast majority of studies character...

Low-intensity transcranial electrical stimulation (tES), including alternating or direct current stimulation, applies weak electrical stimulation to modulate the activity of brain circuits. Integration of tES with concurrent functional MRI (fMRI) allows for the mapping of neural activity during neuromodulation, supporting causal studies of both bra...

The slow wave state is a general state of quiescence interrupted by sudden bursts of activity or so-called slow wave events (SWEs). Recently, the relationship between SWEs and blood oxygen level–dependent (BOLD) functional magnetic resonance imaging (fMRI) signals was assessed in rodent models which revealed cortex-wide BOLD activation. However, it...

Transcranial ultrasound stimulation (TUS) holds great potential as a tool to alter neural circuits non-invasively in both animals and humans. In contrast to established non-invasive brain stimulation methods, ultrasonic waves can be focused on both cortical and deep brain targets with the unprecedented spatial resolution as small as a few cubic mil...

Non-invasive brain stimulation (NIBS) experiments involve many recurring procedures that are not sufficiently standardized in the community. Given the diversity in experimental design and experience of the investigators, automated but yet flexible data collection and analysis tools are needed to increase objectivity, reliability, and reproducibilit...

This handbook is currently in development, with individual articles publishing online in advance of print publication. At this time, we cannot add information about unpublished articles in this handbook, however the table of contents will continue to grow as additional articles pass through the review process and are added to the site. Please note...

Understanding the neural correlates of risk-sensitive skin conductance responses can provide insights into their connection to emotional and cognitive processes. To provide insights into this connection, we studied the cortical correlates of risk-sensitive skin conductance peaks using electroencephalography. Fluctuations in skin conductance respons...

The experimental manipulation of neural activity by neurostimulation techniques overcomes the inherent limitations of correlative recordings, enabling the researcher to investigate causal brain-behavior relationships. But only when stimulation and recordings are combined, the direct impact of the stimulation on neural activity can be evaluated. In...

Background: Sleep consolidates declarative memory by repeated replay linked to the cardinal oscillations of non-rapid eye movement (NonREM) sleep. However, there is so far little evidence of classical glutamatergic plasticity induced by this replay. Rather, we have previously reported that blocking N-methyl-D-aspartate (NMDA) or α-amino-3-hydroxy-...

The perturbational complexity index (PCI) measures the spatiotemporal dynamics of transcranial magnetic stimulation (TMS)-evoked electroencephalography (EEG) potentials (TEPs). High PCI values reflect the joint presence of integration and differentiation in thalamocortical networks of conscious brains. Low PCI values have been reported during natur...

Recent studies suggest that transcranial electrical stimulation (tES) can be performed during functional magnetic resonance imaging (fMRI). The novel approach of using concurrent tES-fMRI to modulate and measure targeted brain activity/connectivity may provide unique insights into the causal interactions between the brain neural responses and psych...

This handbook is currently in development, with individual articles publishing online in advance of print publication. At this time, we cannot add information about unpublished articles in this handbook, however the table of contents will continue to grow as additional articles pass through the review process and are added to the site. Please note...

Background Low intensity transcranial electrical stimulation (tES), including alternating or direct current stimulation (tACS or tDCS), applies weak electrical stimulation to modulate brain circuits. Integration of tES with concurrent functional magnetic resonance imaging (fMRI) allows neuromodulation of brain regions while mapping network function...

Background Sleep consolidates declarative memory by repeated replay linked to the cardinal oscillations of NonREM sleep. However, there is so far little evidence of classical glutamatergic plasticity induced by this replay. Rather, we have previously reported that blocking NMDA or AMPA receptors does not affect sleep-dependent consolidation of decl...

Noninvasive brain stimulation (NIBS) techniques, such as transcranial magnetic stimulation or transcranial direct and alternating current stimulation, are advocated as measures to enable causal inference in cognitive neuroscience experiments. Transcending the limitations of purely correlative neuroimaging measures and experimental sensory stimulati...

Alpha oscillations (8-14 Hz) are assumed to gate information flow in the brain by means of pulsed inhibition, i.e., the phasic suppression of cortical excitability and information processing once per alpha cycle, resulting in stronger net suppression for larger alpha amplitudes due to the assumed amplitude asymmetry of the oscillation. While there...

Frequency tagging has been widely used to study the role of visual selective attention. Presenting a visual stimulus flickering at a specific frequency generates so-called steady-state visually evoked responses. However, frequency tagging is mostly done at lower frequencies (<30 Hz). This produces a visible flicker, potentially interfering with bot...

Ongoing brain activity has been implicated in the modulation of cortical excitability. The combination of electroencephalography (EEG) and transcranial magnetic stimulation (TMS) in a real-time triggered setup is a novel method for testing hypotheses about the relationship between spontaneous neuronal oscillations, cortical excitability, and synapt...

Brain responses to transcranial magnetic stimulation (TMS) recorded by electroencephalography (EEG) are emergent noninvasive markers of neuronal excitability and effective connectivity in humans. However, the underlying physiology of these TMS‐evoked EEG potentials (TEPs) is still heavily underexplored, impeding a broad application of TEPs to study...

Transcranial Magnetic Stimulation (TMS) excites populations of neurons in the stimulated cortex, and the resulting activation may spread to connected brain regions. The distributed cortical response can be recorded with electroencephalography (EEG). Since TMS also stimulates peripheral sensory and motor axons and generates a loud "click" sound, the...

Ongoing brain activity has been implicated in the modulation of cortical excitability. The combination of electroencephalography (EEG) and transcranial magnetic stimulation (TMS) in a real-time triggered setup is a novel method for testing hypotheses about the relationship between spontaneous neuronal oscillations, cortical excitability, and synapt...

Low frequency oscillations such as alpha (8-12 Hz) are hypothesized to rhythmically gate sensory processing, reflected by 40-100 Hz gamma band activity, via the mechanism of pulsed inhibition. We applied transcranial alternating current stimulation (TACS) at individual alpha frequency (IAF) and flanking frequencies (IAF-4 Hz, IAF+4 Hz) to the occip...

Transcranial Magnetic Stimulation (TMS) excites populations of neurons in the stimulated cortex, and the resulting activation may spread to connected brain regions. The distributed cortical response can be recorded with electroencephalography (EEG). Since TMS also stimulates peripheral sensory and motor axons and generates a loud click sound, the T...

Background: Alpha (8-14 Hz) oscillatory power is linked to cortical excitability and corresponding modulations of sensory evoked potentials and perceptual detection performance. In somatosensory cortex (S1), negative linear and inverted U-shape relationships exist, whereas its effect on the primary motor cortex (M1) is hardly known. Objective: W...

The capacity to externally control transcranial magnetic stimulation (TMS) devices is becoming increasingly important in brain stimulation research. Here we introduce MAGIC (MAGnetic stimulator Interface Controller), an open-source MATLAB toolbox for controlling Magstim and MagVenture stimulators. MAGIC includes a series of MATLAB functions which a...

Introduction Focal transcranial magnetic stimulation (TMS) induces somatosensory input due to excitation of peripheral trigeminal nerve branches as well as auditory input caused by the loud click produced during stimulus discharge. In a recent study, we have demonstrated that these peripheral sources of cortical activation make a strong contributio...

Alpha oscillations (8-12 Hz) are hypothesized to rhythmically gate sensory processing, reflected by activity in the 40-100 Hz gamma band, via the mechanism of pulsed inhibition. We applied transcranial alternating current stimulation (TACS) at individual alpha frequency (IAF) and flanking frequencies (IAF-4 Hz, IAF+4 Hz) to the occipital cortex of...

According to the influential “pulsed inhibition hypothesis” (Jensen and Mazaheri, FHN 2010), the 8–14 Hz alpha oscillation, termed μ-rhythm in the sensorimotor cortex, is driven by bouts of inhibition (or deflections of the cortical excitation-inhibition balance towards inhibition), rhythmically suppressing neural processing during the inhibitory p...

Objectives Transcranial Magnetic Stimulation (TMS) is capable to non-invasively stimulate the human cortex. Electroencephalography (EEG) can record the cortical response evoked by TMS (TEPs), which are a summation of the brain responses to the TMS-induced electric field in the cortex, and to the multisensory peripheral stimulation derived from the...

Brain responses to transcranial magnetic stimulation (TMS) as measured with electroencephalography (EEG) have so far been assessed either by TMS-evoked EEG potentials (TEPs), mostly reflecting phase-locked neuronal activity, or time-frequency-representations (TFRs), reflecting oscillatory power arising from a mixture of both evoked (i.e., phase-loc...

Introduction Alpha oscillations (8–12 Hz) are proposed to mediate ‘attentional gating’ by suppressing neuronal processing in task-