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xxiv List of Figures
Fig. 15.6 Facial muscle activity associated with smiling and laughing
produces characteristic artifacts in EEG recordings. Typically showing as slow signal drifts. The gure illustrates these distortions across a subset of 64 EEG channels, highlighting their widespread impact on scalp recordings. The artifacts are most prominent in frontal and temporal regions, where the muscle
activity interferes with the EEG bandwidth ......................... 185
Fig. 15.7 EEG signal distortion caused by higher impedance values,
manifesting in increased sensitivity to high-frequency interference. Electrode and cable movements due to whole body movement introduce random waves of higher amplitudes, which are further masked by 50 Hz line noise. A few EEG channels with
these described artifacts are marked ............... .................. 189
Fig. 15.8 Regular bumps in EEG recordings during running reect head
motion-induced artifacts, with their amplitude and frequency closely tied to the intensity and acceleration of movemen t. Channels are arranged from frontal to posterior regions, progressing from the left to the right hemisphere. The artifact is especially visible in the second half of the window on the frontal
electrodes ....... ....................................................... 190
Fig. 15.9 Cable movement artifacts, compounded by 50 Hz line noise, often
result from higher electrode impedance. Mechanical disturbances, like vibrations, tugging, or loose connections, can introduce
voltage shifts ... .. . .. ... ... .. . .. ... ... .. . .. ... ... .. ... ... ... .. ... ... ... 191
Fig. 17.1 Magnitude response plot of a 4th-order IIR Butterworth low-pass
lter, based on the half-power cutoff denition. About 70.7% of
the original amplitude is retained at the cutoff frequency ......... 216
Fig. 17.2 Topographic maps of blinks and eye movements. (a) Blinks are
prominent bell-shaped deections over the data. They show a predominantly frontal topography. (b) Lateral eye movements show a box-shaped morphology, appear strongest in frontopolar and frontotemporal channels, and typically show opposite polarity on each side of the scalp. Note: the interval between the vertical
green lines is 1 s .. . .. ... .. . .. ... ... .. ... .. . .. ... ... .. ... ... .. ... ... .. . 217
Fig. 18.1 Pendulu
m motion and corresponding sinusoidal oscillation. (a) Pendulum oscillating in a vertical plane, with its bobs displacement on the x-axis restricted to the range -A to A. (b )A sinusoidal function describes the pendulums movement along the
x-axis within this range .......................................... ..... 229
List of Figures xxv
Fig. 18.2 Composite of sinusoids at 5, 10, and 15 Hz and corresponding
frequency representation (a). Sinusoids at three different frequencies: 5, 10, and 15 Hz. (b) Composite signal obtained by summing the three sinusoids in panel a. (c) Power spectrum of the composite signal shown in panel b, showing peaks at the
corresponding frequencies ... ......................................... 231
Fig. 18.3 Dot product between two vectors at different angles. Each panel
illustrates the dot product between vectors A (red) and B (blue). The green vector represents the projection of A onto B, quantifying the component of A that lies in the direction of B. As the angle θ increases, the projection shortens and can become negative, as seen in panel c. (a ) θ = 30 ° (π/6) (b) θ = 90 ° (π/2)
(c) θ = 150 ° (5π /6) .................................................. 232
Fig. 18.4 Representation of the Fourier analysis of sinusoids. (a) A 3 Hz
sine wave (left) and its spectrum (right), showing a single peak at 3 Hz. (b) An 8 Hz sine wave (left) and its spectrum (right), showing a single peak at 8 Hz. (c) The sum of the 3 Hz and 8 Hz sine waves (left). The corresponding spectrum (right) displays two
peaks, reecting the presence of both frequency components . . . . 236
Fig. 19.1 Schematic of the ERP logic. Each trial contains ongoing task-
unrelated background activity and evoked activity that is due to the stimulation. Averaging over many repetitions reduces the ongoing activity and amplies the evoked activity in the nal
ERP .................................................... ................ 240
Fig. 19.2 A simplied overview of the waveform pattern of ERP
components. The gure is based on an auditory oddball paradigm and the ERP at CPz is displayed. Sensory components and a P3 are visible. Note that negative is plotted downward. The approximate time ranges in which the different classes of ERP
components fall are indicated at the bottom ........................ 245
Fig. 19.3 Overview of typical measures extracted from ERPs based on a P3
component. From the peak of a component, both the peak amplitude (a) and the peak latency (b) can be determined. Aggregated measures like the mean amplitude (c) or the area under the curve (d) can be computed within a dened time window around the component. S Stimulus, A Amplitude,
t Time .................................................................. 252
Fig. 20.1 Schema
The right-to-left arrow represents the EEG voltage generation model; source dipole currents on the right spread throughout the tissue, producing a voltage pattern on the left. The left-to-right arrow indicates the inverse problem: the estimation of the most likely current source pattern that could h ave generated the
observed EEG voltage scalp pattern ................................. 259
tic representation of EEG forward and inverse problems.
xxvi List of Figures
Fig. 20.2 Typical EEG source analysis owchart. The source model
determines the subsequent pipeline. If the model is discrete, we need to go the route of dipole modeling and clustering and decide how to perform subsequent single-participant and group analyses on dipoles or clusters of dipoles. If the source model is distributed, we build a participant-specic or template head model and calculate the lead eld using a forward solver (see Table 20.1). Next, we use an inverse solver to estimate the source maps for selected time points of interest in the EEG (see Table 20.2). Then, we can perform connectivity analysis on the source time series. Statistical analyses can be performed on sources or connect ivity features at the participant level or in groups. An important caveat for group analyses is that unless we use the same template head model for all participants, we need to co-register participant source spaces to ensure that the features on which we perform
statistics correspond to the same anatomical structures ............ 260
Fig. 20.3 Statistical inference in the source space. From left to right, the
gure displays a collection of EEG topographies corresponding to the same latency but a different trial. Then we use the inverse solver to estimate single-trial source maps. Afterwards, the estimated source maps are collected into a matrix of sources by trials, which can be used for statistical inference. Note that in this case we group the single-trial EEG topographies into the columns of matrix V; consequently, the error term uses the Frobenius norm (kV - KJk
) instead of the Euclidean norm, and we solve for the
F
matrix J, where each column represents the corresponding single-
trial source estimate ................................................... 264
Fig. 21.1 Schematic of a basic online visualization of the acquired EEG
signals. In parallel, a classier translates the brain activity into a bar graph that the participant is trying to control. Successful (and unsuccessful) control attempts are immediately shown as
feedback, which can in turn alter brain activity .................... 271
Fig. 21.2 Select
ion of possible choices when designing an experiment and deciding what kind of hardware and software to use. The choices can be affected by many different factors (e.g., project-related requirements, personal experience, or simply by having access to specic equipment). In this example, the selected setup for the experiment is driven by practical considerations such as reputation and availability in the lab, but also the fulllment of certain requirements, such as the electrode and ampl ier type, data quality, sampling rate, and number of channels. Raw data can be accessed via an LSL connector provided by the manuf acturer, and a mix of E-Prime
®
and MATLAB® is used to train, visualize, and analyze data. Refer to Chap. 12: Hardware for Recording EEG and Peripheral Physiology and Chap. 13: Software for Recording
EEG and Peripheral Physiology for more information . .. . .. . .. . .. 276
List of Figures xxvii
Fig. 22.1 (a) Averaged ERPs for the oddball trials for one participant in an
oddball experiment in which each participant pressed one button for frequently occurring digits and another button for rarel y occurring letters (or vice versa). Time zero is stimulus onset. (b) Single-trial EEG epochs from the oddball trials that were averaged together to create the waveform in panel A. If the amplitude of the P3 wave is scored as the mean voltage between 300 and 500 ms in the averaged ERP waveform, the analytic standardized measurement error (aSME) can be computed by measuring the mean voltage between 300 and 500 ms in the single-trial EEG epochs and applying the equation shown between panels A and B. (c) Illustration of the concepts of precision, accuracy, and bias for measuring the weight of an object. (d) Conceptual approach for understanding the SME, in which the same participant is tested in 10,000 different replications of the oddball experiment (assuming no fatigue or learning). For each replication, the oddball trials are averaged together, and the mean voltage between 300 and 500 ms is measured from the averaged ERP waveform. The histogram shows the P3 amplitude scores obtained from each of these 10,000 replications. The SME is the standard deviation (SD) of these
10,000 scores .......................................................... 280
Fig. 22.2 Example of how the effect of noise on a score depends on the
scoring method. (a) ERP waveform with no noise. (b) Sa me waveform as A, but wi th high-frequency contamination added. The high-frequency noise distorts the peak amplitude between 300 and 500 ms but has relatively little effect on the mean voltage
during this measurement window .. . .. .. ... .. ... .. .. ... .. ... .. .. ... . 288
Fig. 22.3 Boo
tstrapped standardized measurement error (SME) values from the seven ERP components in the ERP CORE dataset (Kappenman et al., 2021), as derived by Zhang and Luck (2023). The number of trials per condition is given for each component. The SME values were obtained via bootstrapping from the difference wave used to isolate a given component (e.g., oddball minus standard for P3b). Separate SME values were obtained from each participant at the optimal electrode site for each component, and these SME values were then averaged across participants, with error bars indicating the standard error of the mean SME value across participants. Separate SME values are shown for two different amplitude scoring methods (a: time­window mean amplitude and peak amplitude) and for two different latency scoring methods (b: 50% area latency and peak
latency) . ... .... ... ... .... ... ... .... ... ... .... ... ... .... ... ... ... . ... ... . 290
xxviii List of Figures
Fig. 22.4 Effects of low-pass and high-pass lters on real and articial ERP
waveforms. (a) Averaged ERP wave form from an actual research participant, with and without the application of a noncausal Butterworth low-pass lter with a 5 Hz half-amplitude cutoff and a slope of 48 dB/octave. (b) Articial ERP waveform, with and without the same low-pass lter as in (a). (c) Averaged ERP waveform, with and without the application of a noncausal Butterworth high-pass lter with a 2 Hz half-amplitude cutoff and a slope of 12 dB/octave. (d) Articial ERP waveform, with and without the same low-pass lter as in panel C. Note that both the low-pass and high-pass lters reduce the size of the signal as well as reducing the noise. Note also that the high-pass lter produce s artifactual peaks (highlighted with blue circles) that are more
easily observed in the articial waveforms ... .. .. ... .. ... .. .. ... .. . 293
Fig. 23.1 Example of the subtraction method applied to alpha (8–12 Hz)
power during an eyes-closed and an eyes-open condition. On top, the spectrum plot shows a clear power reduction effect after eye opening. When mapping alpha over the scalp, anterior alpha levels common to both conditions occlude the alpha reduction localization. The contrast is created by subtracting the spectra of the eyes-closed from the eyes-open condit ion. This eliminates the common anterior activity and isolates the alpha power reduction
effect to the occipital region ......................................... 308
Fig. 24.1 EEG features as biomarkers. Different EEG features such as
event-related potentials, frequency patterns of neural oscillations, and EEG connectivity can be used as potential biomarkers. These features can be used as biomarkers. Diagnostic biomarkers are used to identify diseases within a population. Prognostic biomarkers are classied as biomarkers that indicate the probability of a disease occurring or progressing. A predictive biomarker helps identify the probability of a group of individuals
that respond to a treatmen t .. ... ... .. ... ... ... .. ... ... ... .. . .. ... ... .. 320
Fig. 24.2 Exa
mple demonstrating real-time analysis of the bispectral index. (a) The raw EEG data is analyzed in near real time to monitor the level of anesthesia and adjust the level of drug administration. (b) The bispectral index is a value ranging from 0 (no brain activity) to 100 (fully awake). The BIS value increases as the brain wave amplitude decreases and the frequency increases. The awake state has a BIS index of 90 and is characterized by small amplitude, fast frequency waves. Deep anesthesia occurs around a score of 30 and is characterized by large amplitude and slow frequency oscillations. Part B of this gure has been reproduced by Youse-
Banaem et al. (2020) with permission .... ........................... 323
List of Figures xxix
Fig. 26.1 The 2025 updated ILAE classication of seizure types categorise
them into further subtypes based on consciousness i.e., preserved consciousness seizure or impaired consciousness seizure. If a seizures starts as focal and then propagates to become a tonic clonic seizure it is classied as focal to bilateral tonic clonic seizure. Under unknown, the subtypes are preserved consciousness seizure, impaired consciousness seizure, and bilateral tonic-clonic seizure. Under generalized, the subtypes are absence seizures, generalized tonic-clonic seizures, and other
generalized seizures ................................................... 352
Fig. 26.2 Patterns of seizure onset frequently observed on scalp EEG. (a)
Rhythmical activity evolving theta, delta, alpha frequencies; (b) Rhythmical spiking; (c) Spike waves; (d) Electro-decremental onset characterized by low-voltage fast activity: (e) Clinical
seizure without a clear EEG correlate ... ... ... ... .. ... ... .. . .. ... ... 355
Fig. 26.3 The eight seizure onset patterns from SEEG, with red asterisks
marking the seizure onset point. (a) Low-voltage fast activity (LVFA). (b) Preictal spiking that transitions into LVFA. (c) A brief burst of polyspikeshigh-frequency (>12 Hz), high­amplitude, short-duration activity lasting less than 5 seconds followed by LVFA. (d) A slow wave or baseline shift (comparable to a DC shift), which precedes LVFA. (e) Rhythmic spikes and spike-wave discharges at low frequencies above 6 Hz and consistently below 14 Hz. (f) Sharp theta or alpha activity, represented by low-frequency sinusoidal waveforms. (g) Sharp beta-frequency activity with beta-band sinusoidal oscillations. (h) A delta-brush pattern characterized by bursts of low-amplitude, rapid gamma-frequency activity superimposed on low-frequency
delta sinusoidal waves ............... ................................. 356
Fig. 26.4 Exa
mples of icEEG implantations. (a) A brain model with a mixed subdural grid, where yellow dots represent individual contacts, and orange and blue dots indicate the position of depth electrodes. (b) A T1 MRI image showing SEEG implantation targeting the right orbitofrontal and mesial frontal regions and
cingulum, marked by yellow dots . .................................. 359
xxx List of Figures
Fig. 27.1 Illustrative data showing how an episode of subtle bradycardia is
preceded by a hypopnoea an d EEG attenuation. Top panel: Heartbeat (RR) intervals (asterisks) extracted from a 19 min recording segment from an infant of 35 weeks corrected gestational age (gestational age + postnatal age) using the EEG-Beats toolbox. Note the RR intervals variability, including intermittent bradycardias with one example shaded pink. Bottom panel: EEG, electrocardiography (ECG) and respiratory movement (Resp) recordings associated with the shaded example bradycardia. The infant is awake, 3 min before a transition into REM sleep. No lters applied except 50 Hz notch lter; DC offset
removed. Midline central referential EEG montage ................ 368
Fig. 27.2 Examples of annotated events occurring during home EEG
monitoring. Annotations collated from a deidentied research database of multiple home-video EEG sessions carried out in paediatric populations at King s College Hospital. For illustrative purposes, annotations are superimposed onto a 20 min segment of EEG, electrocardiography (ECG), and electromyography (EMG) activity from a patient of 10 years of age undergoing 24-h home­video EEG monitoring. The child is awake. Midline central
referential EEG montage . ... .. ... ... .. ... ... .. . .. ... .. . .. ... ... .. . .. . 371
Fig. 28.1 Exa
sleep. (a) An example hypnogram. The hypnogram depicts the cyclic alternation between NREM and REM sleep roughly every 90 min. N3 predominates early in the night, whereas the proportions of N2 and REM sleep increase in the latter half of the night. (b) Representative recordings of EEG, horizontal and vertical EOG, and EMG across different sleep stages, including wakefulness, N1, N2, N3, and REM. During wakefulness, alpha waves (8–13 Hz) appear continuously. N1 is a drowsy state in which alpha activity diminishes and theta waves (4–8 Hz) emerge in EEG signals, while EOG shows slow eye movements (SEMs). N2 is characterized by sleep spindles (11–16 Hz) and K-complexes. N3 shows large, synchronous slow waves (0.5–4 Hz). During REM sleep, EEG signals exhibit mixed high­frequency, low-amplitude activity resembling that of N1 or wakefulness. EOG displays periodic bursts of rapid eye movements, whereas EMG recordings show markedly reduced muscle tone. Negative polarity is up in the scale. hEOG horizontal
EOG, vEOG vertical EOG ........................................... 387
mples of a hypnogram and physiological recordings during
List of Figures xxxi
Fig. 28.2 Asymmetric sleep related to the rst-night effect in humans. (a)
Interhemispheric differences in slow-wave activity in the default mode network. Red, left hemisphere; blue, right hemisphere. (b) Relationship between the sleep onset latency and the asymmetry index on Day 1. (c) Relationship between the sleep onset latency
and the asymmetry index on Day 2 .. . .............................. 391
Fig. 28.3 E/I balance during sleep. (a) The E/I balance changes during
NREM (orange) and REM (blue) sleep. During NREM sleep, the E/I balance is signicantly higher than during the wake baseline, whereas during REM sleep, the E/I balance is signicantly lower than the wake baseline. (b) The correlation between E/I balance changes during NREM sleep and ofine performance gains. Red, NREM + REM group. Gray, NREM-only group. (c) The correlation between E/I balance changes during REM sleep and
stabilization of learning . .............................................. 395
Fig. 28.4 CSF dynamics during sleep. (a) CSF signal changes to slow
waves and sleep spindles during light NREM sleep. During light NREM sleep, sleep spindles were correlated with signicant CSF signal changes at 6 s after onset, whereas slow waves preceded a large CSF signal peak at 8 s after onset. (b) CSF signals changes to slow waves and sleep spindles during deep NREM sleep or slow­wave sleep. During slow-wave sleep, slow waves triggered CSF signal changes peaking at 5.5 s, in which latency was signicantly shorter and smaller in amplitude. Sleep spind les preceded CSF signal changes at 4 s after their onset, which lasted for
approximately 10 s . ................................................... 397
Fig. 28.5 Brain regions recruited during various stages of sleep and for
arousals. The red-highlighted parts indicate brain areas activated to slow waves during light NREM (top row) and slow-wave sleep (the second row), rapid eye movements during REM sleep (the third row), and arousals (the bottom row ). Light NREM sleep activates sensory and motor regions including the parietal and visual cortices and cerebellum. Slow-wave sleep engages regions involved in plasticity and homeostatic processing including the prefrontal and hippocampal regions, striatum, and amygdala. REM sleep recruits visual and plasticity circuits including the visual areas, the thalamus, the hippocampus, the striatum, and the amygdala. Arousals accompanied widespread brain activation not
localized to specic brain regions ................................... 398
Fig. 29.1 A traditionalEEG system, the Brain Products actiChamp
(Brain Products GmbH, Gilching, Germany) . ... ... ................ 404
Fig. 29.2 A mobileEEG system, the Brain Products X.on (Brain
Products GmbH, Gilching, Germany) ............................... 405
Fig. 29.3 EEG
being recorded while riding a bike ... . .. ... ... .. ... ... .. ... ... 406
xxxii List of Figures
Fig. 29.4 The Muse S Athena. A combined EEG and fNIRS system ....... 407
Fig. 29.5 The P300 measured at Pz (top panel) TP9 and TP10 (middle
panel) with a Brain Products ActiChamp and an InterAxon Muse (bottom panel). (From Choosing Muse, Krigolson et al., 2017) . . 412
Fig. 29.6 P300 amplitude is reduced following a simulated 12 h night on
call . .... ... ... ... . ... ... ... . ... ... ... .... ... ... ... . ... ... ... . ... ... ... ... 414
Fig. 29.7 Exercising outside enhances P300 amplitude after a 15 min
walk .......................... .......................................... 416
Fig. 29.8 Decreased EEG beta power predicts baseball batting
performance . ... .. ... ... ... .. ... ... ... .. ... ... ... .. ... ... ... .. . .. ... ... 417
Fig. 30.1 Mobile Brain/Body Imaging (MoBI) system setup, experimental
workow in representative art-science performances. (a) Standard MoBI conguration with 32-ch EEG (BrainAmp DC, 28 scalp + 4 EOG, 1 kHz) and IMUs (APDM Opal, 9-axis, 128 Hz), housed in a wearable pack adapted to each performance. (b) The Slowest Wave performance with. (c) Brain On Nature project. (d) Balinese Gamelan, Brain, Mind and Body performance in collaboration with Udayana University and Institut Seni Indonesia (ISI) Denpasar. (e) Typical experimental timeline illustrating impedance check, resting baseline (eyes open/eyes closed), and
the experimental task phase ... ... ... ... ... .. ... ... ... ... ... .. ... ... .. 428
Fig. 30.2 Mobile Brain/Body Imaging (MoBI) system architecture. A
typical setup integrating EEG (BrainAmp DC, Brain Products GmbH, Gilching, Germany), IMUs (APDM Opal), audiovisual recording, and a synchronization layer (SyncBox) to enable multimodal, time-aligned data collection in naturalistic environments. The system is adapted per study to align with task
demands, movement constraints, and aesthetic requirements . . . . . 430
Fig. 30.3 A visual
timeline illustrating the Brain + Arts collaborative
projects from 2014 to 2025 .......................................... 433
Fig. 30.4 Multim
odal data processing pipeline for MoBI studies in naturalistic settings. The framework supports synchronized preprocessing and artifact removal across video, EEG, EOG, and IMU signals. It consists of four stages: Stage 0 (Data Understanding and Annotation), Stage 1 (Eye Artifact Removal), Stage 2 (Motion Artifact Removal), and Stage 3 (Source-Space Artifact Removal). This full pipeline is used for ofine data cleaning and analysis. A simplied real-time implementation using H-innity ANC is currently employed during live performances for visualization, but is not yet congured for
closed-loop neurofeedback or BCI control ......................... 440
List of Figures xxxiii
Fig. 31.1 Grand average power at electrode POz before, during, and after 20
min of 10 Hz tACS (B) or sham (A) during a visual oddball task. The orange lines depict the average power before stimulation, the gray lines depict recovered power during stimulation, and the green lines depict the power after stimulation. While there are no signicant differences during sham stimulation around 10 Hz between conditions during sham stimulation, during tACS an increase is visible during (ISI) and after tACS stimulation (adapted from Helfrich et al. (2014b) and published with
permission by Elsevier) . .............................................. 455
Fig. 31.2 Average power at electrode Pz before and after 20 min of IAF
tACS (top) and sham (bottom) during an eyes-open auditory detection task. The dashed lines indicate the pre-stimulation power, solid lines indicate post-stimulation power, and the shaded areas represent the standard error of the mean. Even after sham, alpha power increases over time from pre- to post-intervention (blue spectra). After tACS, this increase is signicantly stronger (red spectra). This nicely illustrates why a sham group is necessary for tACS (adapted from Neuling et al. (2013) under the Creative
Commons Attribution License (CC BY 3.0)) .. .. .. . . . . . .. .. .. .. .. . 459
Fig. 31.3 Basic
principle of dividing spectral data into periodic and aperiodic components. (a) Original total spectrum composed of aperiodic activity (1/f-oor) and two oscillatory signals (visible as peaks in the spectrum). The magnitude of the peaks represents the magnitude of the periodic activity plus the underlying noise oor. (b) The same spectrum as viewed on a double-logarithmic scale (blue line). On this scale the aperiodic noise oor resembles a straight line. Through iterative tting procedures, the total spectrum is divided into an aperiodic t (red) and an oscillatory t (yellow). (c) Subtracting the aperiodic t from the total spectrum (transparent blue curve) yields the corrected purely periodic spectrum (solid blue), from which we can now extract the corrected peak-frequency and peak-power values for further analysis of periodic activity. (d) The aperi odic t (red) can be used for the characterization of the pink-noise activity: We can extract the offset ( log
of the intercept in log-log space) and the
10
exponent (-slope in log-log space) of the pink-noise activity . . . . . 463