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Файл:Ординатура / Хирургия / Библиотека им академика М.И. Перельмана / Книга_6027_Библиотеки_им_академика_М_И_Перельмана.pdf
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- •Preface
- •Acknowledgments
- •About This Book
- •Contents
- •List of Figures
- •List of Tables
- •Editor and Contributors
- •1.3 EEG and Other Neuroscience Methods
- •1.4 The Future of EEG
- •1.1 EEG Technology: Past to Present
- •1.2 What Do We Know About the EEG Signal?
- •1.5 Conclusions
- •References
- •2.1 Physiological Origins of the EEG
- •2.2 Signals of the EEG
- •2.3 Concluding Summary
- •References
- •3.1 Introduction
- •3.2 General Organization
- •3.3 Finding Your Way Around: Brain Atlases
- •3.3.2 Talairach Atlas and MNI Coordinates
- •3.3.4 Accessing and Using Atlases
- •3.4 Putting into All Together
- •3.5 Conclusion
- •References
- •4.1 Introduction
- •4.2 Overview of the Peripheral Nervous System
- •4.3 Basic Anatomical Unit of the PNS: Ganglia and Nerves
- •4.4 Anatomy of Somatic Nervous System
- •4.4.1 Receptors
- •4.4.1.1 Vision
- •4.4.1.2 Audition
- •4.4.1.3 Vestibular System and Balance
- •4.4.1.4 General Sensory Modalities
- •4.4.2 Somatic Sensory System
- •4.5 Anatomy of the Autonomic Nervous System
- •4.5.1 Sympathetic Nervous System
- •4.5.2 Parasympathetic Nervous System
- •4.6 Cranial Nerves
- •4.7 Function of the PNS and CNS as a Unit
- •4.8 Concluding Remarks
- •References
- •5.1 Introduction
- •5.2 Head Anatomy and Signal Propagation
- •5.3.1 The Eyes and Ocular Potentials
- •5.3.2 Facial Muscles and EMG
- •5.3.3 Sweat Glands and Skin Potentials
- •5.3.4 Blood Vessels and Heartbeat
- •5.4 Conclusion
- •References
- •6.1 Introduction
- •6.2.1 What Is a Brain State?
- •6.2.2 Brain States Measured with EEG
- •6.3 Examples of Brain States
- •6.3.1 Awake State Sleep State
- •6.3.2 Consciousness States: Presence Loss (Anesthesia)
- •6.4 Pathological Brain States
- •6.4.1 Traumatic Brain Injury
- •6.4.2 ADHD
- •6.5 Framework of Brain States
- •6.6 Concluding Summary
- •References
- •7.1 Introduction
- •7.3 Identifying Task Processes Within a Trial Segment
- •7.3.2 Cross-Trial Comparability and Flexible Time-Locking
- •7.4 What Does Activity Look Like on the Timeline?
- •7.5 Conclusion
- •References
- •8.1 Introduction
- •8.2 Setting a Research Question
- •8.3 Setting a Hypothesis
- •8.3.2 Testing the Hypothesis
- •8.4 Design of the Study
- •8.4.1 Contextualization of the Hypothesis
- •8.4.1.1 Experimental Paradigm
- •8.4.1.2 EEG Index
- •8.4.1.3 Group/Sample
- •8.4.2 Implementation of the Study
- •8.4.2.1 Paradigm/Task Implementation
- •8.4.2.2 Measurement Precision
- •8.4.2.3 Experimental Protocol
- •8.4.2.4 Pilot Testing
- •8.5 Concluding Summary
- •References
- •9.1 Introduction
- •9.2 Why Is Statistics Needed in EEG Research?
- •9.3 When Is Statistics Applied During EEG Data Analysis?
- •9.3.1 Raw EEG Data
- •9.3.2 Individual-Level (First-Level) Analysis
- •9.3.3 Group-Level (Second-Level) Analysis
- •9.3.3.1 Statistical Hypotheses
- •9.3.4 Application of Statistical Inference
- •9.3.5 Interpretation and Inference
- •9.4.1 Hypotheses (Upper Plane of Fig. 9.2)
- •9.4.2 Population and Sample Data (Bottom Plane of Fig. 9.2)
- •9.4.3 Sample Statistic (Middle Plane of Fig. 9.2)
- •9.5 Conclusion
- •References
- •10.1.1 Why Pilot Testing Matters
- •10.2 How to Prepare and Run the Pilot Testing
- •10.2.1.1 Signal Quality
- •10.2.1.2 Task Parameters
- •10.2.1.3 Instructions
- •10.2.1.4 Participant Experience
- •10.2.1.5 Equipment Setup
- •10.2.1.6 Procedures
- •10.2.1.7 Questionnaires
- •10.1 What Pilot Testing Is
- •10.3 Concluding Summary
- •References
- •11.1 Introduction
- •11.2 Lab Management
- •11.2.1 Admin and Organisation
- •11.2.2 Hardware and Software Maintenance
- •11.2.3 Lab Logbook
- •11.3 Keep Your Own Lab Notebook
- •11.4.1 Pre-measurement
- •11.4.2 Measurement
- •11.4.3 Post Measurement
- •11.5 Conclusion
- •References
- •12.1 Introduction
- •12.2 Components of the System
- •12.2.1 Detecting the Signal: EEG Electrode Technology
- •12.2.1.1 Passive Electrode Plus Gel
- •12.2.1.2 Active Electrodes Plus Gel
- •12.2.1.3 Passive Electrode and Saline Soaked Sponges
- •12.2.1.4 Dry Electrodes
- •12.2.1.5 Electrode Positions
- •12.2.2 Detecting the Signal: Sensors for Other Measures
- •12.2.2.1 Bipolar Peripheral Electrophysiology
- •12.2.2.2 Peripheral Physiological Sensors
- •12.2.2.3 GSR
- •12.2.2.4 Respiration
- •12.2.2.5 Photoplethysmography (PPG)
- •12.3 Conclusion
- •References
- •13.1 Purpose and Features
- •13.2 Before Starting Your Study
- •13.2.1 General Parameters
- •13.2.2 Special Applications
- •13.2.3 Real-Time Processing
- •13.3 During a Measurement Session
- •13.4 Troubleshooting
- •13.5 Conclusion
- •References
- •14.1 Introduction
- •14.2 Importance of Triggers
- •14.3 Advantages of Triggers
- •14.4 Disadvantages of Triggers
- •14.5 Alternatives to Triggers
- •14.6 Good Practice for Using Triggers
- •14.7.1 Setup
- •14.7.2 Analysis
- •14.7.3 Interpretation
- •14.8 Conclusion
- •References
- •15.1 Introduction
- •15.2 The Idea of Signal-to-Noise Ratio (SNR)
- •15.3 Sources of Artifact
- •15.4 Common Physiological Artifacts
- •15.4.1 Eye Artifacts
- •15.4.2 ECG Artifacts
- •15.4.4 Other Physiological Artifacts
- •15.5 Common Technical Artifacts
- •15.5.1 Technical Artifacts
- •15.5.2 Electrode Artifacts
- •15.5.3 Gel-Related Artifacts
- •15.5.4 Movement Artifacts
- •15.5.5 Body/Head Movements
- •15.5.6 Cable Movement Artifacts
- •15.6 Artifacts in Advanced Applications and Multi-modal Recordings
- •15.6.1 EEG and Functional MRI
- •15.6.2 EEG and Non-invasive Brain Stimulation
- •15.7 Optimizing the EEG Recording Quality
- •15.7.1 Focus on the Cap Preparation
- •15.7.2 Optimize the Recording Environment
- •15.7.3 During the Recording
- •15.7.4 Post Recordings
- •15.8 Conclusion
- •References
- •16.1 Introduction
- •16.2 Lab Infrastructure
- •16.2.1 Signal Quality
- •16.2.2 Control Over the Experimental Environment
- •16.2.4 Safety
- •16.3 Position of the Equipment and Accessories
- •16.4 Lab Procedures
- •16.5.1 Mobile Setups
- •16.5.2 Electrode Types
- •16.5.2.1 Passive Sponge-Based Electrodes
- •16.5.2.3 Dry Electrodes
- •16.5.3 Special Populations
- •16.5.3.1 Children
- •16.6 Concluding Summary
- •References
- •17.1 Introduction
- •17.2 Common Preprocessing Steps: Data Transformation
- •17.2.1 Inspecting Data
- •17.2.2 Changing the Sampling Frequency
- •17.2.3 Re-referencing
- •17.2.4 Interpolating Channels or Data Portions
- •17.2.5 Segmenting Data
- •17.3 Common Preprocessing Steps: Artifact Handling
- •17.3.1 Filtering
- •17.3.2 Attenuating Artifacts
- •17.3.2.1 Independent Component Analysis (ICA)
- •17.3.2.2 Regression Techniques
- •17.3.2.3 Template Subtraction Methods
- •17.3.3 Rejecting Artifacts
- •17.5 Tools for Processing and Analyzing EEG
- •17.6 Concluding Remarks
- •References
- •18.1 Introduction
- •18.2 Characterizing an Oscillatory Process
- •18.2.1 Fundamental Characteristics of an Oscillatory Process
- •18.2.2 From Time to Frequency and Back
- •18.3 Foundation for Spectral Analysis: The Dot Product
- •18.4 Fourier Analysis
- •18.4.1 From Vectors to Sinusoids: The Fourier Connection
- •18.4.2 The Fourier Family
- •18.4.3 Discrete Fourier Transform
- •18.4.4 Power Spectrum
- •Further Readings
- •References
- •19.1 Introduction
- •19.2 How to Get from EEG to ERPs
- •19.2.1 How to Process Your ERP Data
- •19.2.1.1 Pre-processing
- •19.2.1.2 Trial Selection
- •19.2.1.3 Baseline Correction
- •19.2.1.4 Averaging
- •19.2.2 Interpreting ERPs
- •19.2.3 Group Analysis
- •19.2.4 Single-Trial Analysis
- •19.3 Characteristics of the ERP and Its Components
- •19.4 Commonly Investigated ERP Components
- •19.4.1 Early Sensory Components
- •19.4.2 Long-Latency Sensory Components
- •19.4.3 Later Cognitive Components
- •19.4.4 ERP Components in Multimodal Recording Scenarios
- •19.5 Extraction of ERP Features
- •19.6 Conclusion
- •References
- •20.1 Introduction
- •20.2 Fundamentals of EEG Source Imaging
- •20.3 Forward Problem
- •20.4 Source Estimation
- •20.5 Statistical Inference in the Source Space
- •20.6 Source Connectivity
- •20.7 Conclusion
- •References
- •21.1 Introduction
- •21.2 Raw Data Access
- •21.2.1 How to Get Raw Data
- •21.2.2 How to Work with Raw Data Online
- •21.2.3 What Factors to Consider for Online Processing
- •21.3 Designing an Online Processing Experiment, an Example
- •21.4 Conclusion
- •References
- •22.1 Introduction
- •22.1.2 Chapter Overview
- •22.2.2 Exactly What the SME Means
- •22.2.5 Why the Scoring Method Matters
- •22.2.8 Other Potential Uses of the SME
- •22.4 Metrics of Reliability
- •22.5 Final Thoughts
- •References
- •23.1 Cognitive Neuroscience
- •23.1.1 Neuropsychology and EEG
- •23.1.2 Mental Chronometry and EEG
- •23.2 Research on the EEG Signals
- •23.3 Conclusions
- •References
- •24.1 Introduction
- •24.2.1 Clinical Research
- •24.2.2 EEG in Research for Clinical Applications
- •24.2.3 EEG as a Biomarker
- •24.3 Examples of Clinical Applications of EEG
- •24.3.1 Epilepsy
- •24.3.2 Sleep and Sleep Disorders
- •24.3.3 Anesthesia
- •24.4.2 Brain-Computer Interfaces and Movement Disorders
- •24.5 Future of EEG in Clinical Applications
- •24.6 Conclusion
- •References
- •25.1 Introduction
- •25.1.1 Why Connect Brains and Computers?
- •25.1.2 What Is a BCI?
- •25.1.3 Types of BCIs: Active, Reactive, Passive
- •25.1.4 BCIs in Neuroscience and HCI
- •25.2 Signals and Sensors
- •25.2.1 Neural Signals for BCIs
- •25.2.2 Wearable EEG and Form Factors
- •25.3 The BCI Pipeline: From Raw Signals to Decisions
- •25.3.1 Overview
- •25.3.2 Experimental Design and Labeling
- •25.3.3 Preprocessing and Artifacts
- •25.3.4 Feature Extraction
- •25.4 BCI Types Illustrated
- •25.4.1 Motor Imagery as an Active BCI Paradigm
- •25.4.2 P300 and SSVEPs as Reactive BCI Paradigms
- •25.4.3 Workload, Error, and Other Passive BCI Paradigms
- •25.5.1 Mental State Assessment as a First Stage
- •25.5.2 Open- and Closed-Loop Adaptation
- •25.6 Practical Challenges
- •25.6.1 Mobility, Artifacts, and Non-stationarity
- •25.6.2 Cross-User and Cross-Session Generalization
- •25.6.3 Evaluation in Real Settings
- •25.6.4 Ethics, Privacy, and Neurorights
- •25.7 Conclusions and Outlook
- •25.7.1 Key Takeaways
- •25.7.2 Future Trajectories
- •References
- •26.1 Focal Epilepsy
- •26.2 EEG Manifestations of Focal Epilepsy
- •26.2.1 Ictal EEG Patterns
- •26.2.2 Interictal EEG Patterns
- •26.3 Localization of Ictal and Interictal EEG Events
- •26.4 Intracranial EEG in Presurgical Planning
- •26.5 AI in EEG Interpretation
- •26.6 Conclusion
- •References
- •27.1 Introduction
- •27.2 Neonatal EEG Applications
- •27.2.2 Somatosensory States Monitoring in Neonates
- •27.3 Paediatric EEG Applications
- •27.3.1 Sleep Monitoring in Children and Adolescents
- •27.4 Future Directions and Conclusion
- •References
- •28.1 What Is Sleep?
- •28.1.1 Stages of Sleep
- •28.1.2 How Sleep Changes with Age
- •28.2 Measuring Human Sleep
- •28.2.1 The Various Forms of Sleep
- •28.2.2 Unihemispheric Sleep
- •28.3.1 NREM Sleep and Learning
- •28.3.2 REM Sleep and Learning
- •28.4.1 Active Brain Networks During Sleep
- •28.4.2 Measuring the Balance of Excitation and Inhibition in the Human Brain
- •28.4.3 The Cerebrospinal Fluid Dynamics in Human Sleep
- •28.5 Conclusions
- •References
- •29.1 What Is Mobile EEG?
- •29.2 Range of mEEG Systems
- •29.3 Technical Considerations
- •29.4 Validation
- •29.5 Application
- •29.6 Mild Cognitive Impairment
- •29.7 mEEG and Health and Exercise
- •29.8 mEEG in Sports
- •29.9 Conclusions
- •References
- •30.1 Introduction
- •30.2 General Framework and System Overview
- •30.3 Spectrum of Studies
- •30.4 MoBI+ Framework
- •30.5 Processing Multimodal MoBI Data
- •30.6 Challenges and Limitations
- •30.7 Conclusion
- •Appendix
- •List: Traveling with MoBI Equipment
- •References
- •31.1 Introduction
- •31.2 Types of Electric Brain Stimulation
- •31.3 Online Effects
- •31.3.1 Conventional Artifact Removal Strategies
- •31.3.1.1 Transcranial Direct Current Stimulation (tDCS)
- •31.3.1.2 Transcranial Random Noise Stimulation (tRNS)
- •31.3.1.3 Transcranial Alternating Current Stimulation (tACS)
- •31.3.3 Innovative Approaches to Minimize Artifacts
- •31.3.3.1 Non-Sinusoidal Waveforms
- •31.3.3.2 Amplitude-Modulated tACS (AM-tACS)
- •31.3.3.3 Transcranial Temporal Interference Stimulation (tTIS)
- •31.3.3.4 Summary of Advantages and Limitations
- •31.4.1 Spectral Power
- •31.4.2 Phase Locking/Phase Coherence
- •31.4.3 ERPs
- •31.4.4 Further Measures
- •31.5.1 Rationale
- •31.5.2 Procedure
- •31.6 Closed-Loop Systems
- •31.7 Technical Requirements
- •31.8 Conclusion
- •References
- •32.1 Introduction
- •32.2.1 Equipment

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Chapter 32
TMS–EEG: A Tool to Probe Key Features
of Human Thalamocortical Circuits
Silvia Casarotto and Mario Rosanova
Abstract Transcranial magnetic stimulation combined with electroencephalogra-
phy
(TMS–EEG) offers a unique opportunity to investigate human thalamocortical
circuits non-invasively from a causal perspective. Delivering a controlled perturbation—the TMS pulse—to a targeted cortical region and recording the ensui ng
TMS-evoked potentials (TEPs) reveals physiological features that complement
those accessible through observation alone, such as through the resting EEG.
These features include cortical excitability, effective connectivity, oscillatory
dynamics, and recurrent interactions across cortical and thalamocortical circuits.
Methodological advances—ranging from TMS-compa tible EEG equipment to biological artefact-control procedures—now allow researchers to isolate genuine neural
responses from stimulation-related confounds with increasing reliability. These
insights are turning TMS–EEG into a versatile tool for basic and clinical neuroscience, enabling the characterization of circuit-level mechanisms across behavioral
states, from wakefulness and sleep to anesthesia and disorders of consciousness, as
well as fostering the development of biomarkers relevant for stroke, psychiatric
disorders, neurodegeneration, and recovery of function. This chapter provides an
overview of the instrumentation, experimental procedures, physiological principles,
and representative applications of TMS–EEG, highlighting both its strengths and
limitations. We argue that TMS–EEG holds substantial promise for advancing
mechanistic models of thalamocortical function in humans and for developing
clinically meaningful biomarkers.
Keywords TMS–EEG · Thalamocortical
circuits · TMS-evoked potentials (TEPs) ·
Effective connectivity · Neurophysiological biomarkers
S. Casarotto
Department of Biomedical and Clinical Sciences, University of Milan, Milan, Italy
IRCCS Fondazione Don Carlo Gnocchi ONLUS, Milan, Italy
M. Rosanova (*)
Department of Biomedical and Clinical Sciences, University of Milan, Milan, Italy
e-mail:
mario.rosanova@unimi.it
© The Author(s), under exclusive license to Springer Nature Switzerland AG 2026
Warbrick (ed.), The EEG Handbook,
T.
https://doi.org/10.1007/978-3-032-20450-9_32
479

480 S. Casarotto and M. Rosanova
32.1 Introduction
The thalamocortical system plays a central role in human perception, motor behavior, cognition, and consciousness. These functions arise from interactions between
local cortical circuits and long-range cortico-cortical and thalamocortical loops that
regulate excitability, synchronize oscillations, and sustain recurrent processing.
Dysregulation of these mechanisms characterizes many neurological and psychiatric
disorders, in which altered excitability, connectivity, or oscillatory dynamics impair
information processing and behavior. Understan ding how these processes fail—and
how they may be restored—requires tools capable of probing thalamocortical
function directly and non-invasively. Transcranial magnetic stimulation combined
with electroencephalography (TMS-EEG) is uniquely suited to this goal, as it
captures how neural activity elicited by a focal perturbation propagates through
thalamocortical circuits.
Early TMS-evoked potentials (TEPs) components (10–50 ms) primarily reflect
cortical excitability and the recruitment of intracortical and thalamocortical
local
circuits, indexing changes in membrane excitability, synaptic efficacy, or
neuromodulatory state. At longer latencies (>50 ms), TEPs reveal the propagation
of activity across connected regions, providing a direct measure of effective connectivity that is highly sensitive to brain state, despite preserved structural connectivity—for example, during NREM sleep, anesthesia, or disorders of consciousness.
In addition, TMS can elicit sustained oscillatory responses at frequencies characteristic of specific cortical targets, reflecting the natural frequencies of corticothalamic
modules and conveying information about excitation–inhibition balance and network organization. State-dependent modulation of these responses distinguishes
conditions with preserved causal interactions, such as wakefulness or the minimally
conscious state, from those in which large-scale dynamics collapse, as in deep sleep,
anesthesia, or unresponsive wakefulness syndrome. The degree of such complex
causal interactions can be quantified by TMS–EEG–derived measures such as the
Perturbational Complexity Index (PCI), which discriminates between clinically
similar conditions with high sensitivity.
This chapter provides a focused overview of the methodological and physiolog-
foundations of TMS–EEG , with particular emphasis on its application to
ical
thalamocortical circuits. We review the instrumentation and experimental procedures required for reliable data acquisition, summarize the physiological interpretation of TEPs, oscillatory responses, connectivity and complexity metrics, and
present representative applications in basic, clinical, and translational neuroscience .
32.2 TMS–EEG: A Brief Overview
TMS is a non-invasive brain stimulation technique based on Faraday’s law of
electromagnetic induction. A brief, high-intensity current pulse—typically several
kiloamperes and lasting a few hundred microseconds—is delivered through a stimulation coil (Ilmoniemi et al.,
1999), generating a rapidly time-varying magnetic

32 TMS–EEG: A Tool to Probe Key Features of Human Thalamocortical Circuits 481
field that penetrates the scalp and skull with minimal attenuation and induces an
electric field in a focal region of the cerebral cortex (Ilmoniemi et al.,
magne
tic field strengths usually range between 1 and 3 tesla and rise within ~50–100 microseconds, providing TMS with submillisecond temporal precision. This
temporal accuracy allows controlled perturbation of neuronal membrane potentials
and modulation of ongoing and oscillatory brain activity, enabling causal interrogation of human brain circuits that are otherwise inaccessible to purely observational
methods (Hernandez-Pavon et al.,
Electroencephalography (EEG) records the electrical activity generated by large
populations of cortical neurons via scalp electrodes, offering millisecond temporal
resolution and spatial resolution that depends on electrode density and head conductivity (Ilmoniemi & Sarvas,
inferen
ce, it yields a direct measure of brain dynamics as they evolve over time.
Because of the intense electric field generated by the TMS pulse, combining TMS
EEG required the development of dedicated TMS-compatible amplifiers. Early
with
attempts in 1989 demonstrated the feasibility of recording transcallosal and
cerebellar–cortical responses (Cracco et al.,
use
of conventional EEG amplifiers resulted in large pulse-related artifacts that
prevented reliable recording under the coil. Fully TMS-compatible amplifiers introduced about twenty years ago (Ilmoniemi et al., 1997; Virtanen et al., 1999)
ssed these electromagnetic artifacts and enabled the reliable recording of
suppre
artifact-free TMS-evoked potentials (TEPs) within a few milliseconds (Ilmoniemi
et al., 1997; Paus et al. 2001; Massimini et al., 2005). More recently, DC amplifiers
with
wide dynamic range and high sampling rates (
the ability to capture TEPs without long-lasting pulse artifacts (Bonato et al., 2006).
The concurrent application of TMS and EEG enables the recording of neural
respon
ses to a precisely timed external perturbation. This combined approach allows
investigators to (i) evoke and measure cortical responses, (ii) probe how ongoing
brain activity is perturbed by stimulation, and (iii) implement state-dependent or
closed-loop stimulation paradigms in which TMS is adapted to the current brain state
(Bergmann,
Together, these features make TMS–EEG a powerful framework for studying the
causal
and thalamocortical circuits in health and disesase (Ziemann et al., 2026 10.1016/j.
clinph.2025.2111487).
2018; Hernandez-Pavon et al., 2023; Zrenner & Ziemann, 2024 ).
organization, state dependence, and dynamical properties of human cortical
2023; Koponen et al., 2018b).
2019). While EEG alone provides limited causal
1989; Amassian et al., 1992), but the
5 kHz) have further improved
1997). Peak
32.2.1 Equipment
The acquisition of high-quality TMS–EEG data depends on the careful selection and
integration of dedicated hardware and software components (Fig.
typica
l system includes (i) the TMS stimulator and coil, (ii) a TMS-compatible EEG
amplifier, and (iii) a neuronavigation system for accurate, reproducible targeting.
Additional elements—such as a graphical user interface for real-time monitoring and
32.1, Panel A). A
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